sovaver April 4, 19 NATURE 7 & )tuapstxs v A NEW SPECIES OF THE GENUS HOMO FROM OLDUVAI GORGE By Dr. L. S. B, LEAKEY Coryndon Museum, Centre for Prehistory and Palzontology Prot. P. V. TOBIAS University of Wiewatersrand, Johannesburg Dx. J. R. NAPIER Unit of Primatology and Human Evolution, Royal Free Hospital Medical School, niveaity of London HIE recent disperfries of fossit hominid remains at Olduvai Gorge have strongthened the conclusions — which eqgh Of us had reached independently through our respective investigations —that the fozeil hominid reraains found in 1960 at site .DK.N.N. I, Olduvai, did not ropresent a eroaturo bolonging to the sb-family Australo- pithoeinae Wo sor proparing to publish the evidence for thie ‘conclusion antd to give a sci¢ntifie name to this new species ‘of Lhe gens Homo, when the new discoveries, which are teenie by 1.5.8. and MD. Leaky i tho proerhig, Article, wore made, ‘An examination of theee finds has onabled 1s to broaden tho basis of our diagnosis of the propased new spacies and: has fully confirmed the presence of tho genus Homo in the tower part of the Olduvai geological sequence, earl than, ‘contemporary with, aswell as later than, the Zinjanthsopus skull, which is cortainly an australopithe For the purpose of our deccription here. wo have scoapte the diagnosis of the family Hominidae, ag it was Proposed by Sir Wilfrid Lo Gros Clark in his book The Fossil Evidence for Human Evolution (10: 1955). Within Uhis family’ wo accept the genns Ausiralepithecres with, for the momont, three sub genors. (duatralepithecns, Paranihropnes and Zinjonthreyms) end the pone. Homo Wo regard Pithecanthropus and possibly also taanthrops GE it ts indood “distinet) a2 spacies of the yonus Homo, although ono of ws (L. 8. BL.) would be prepared to accope Sub generic. rane Tt haa long boon recognized that as more and more diveoverios were mado, it would beeomo necessary to sevise the diagnosis of the genus Homo. In particular, it has become clear that it is impossible to rely on ony one oF two characters, such as the cranial capacity or ar erect. posture, as the necessary eritoria for mombership of the fois Tastead, the total picture presented by) the material available for investigation must be taken inte We lee come to the conclusion that, apart from Australopihecus (Zinjanthropus), the specimens we are leaing with From Bod T and the lower part of Bod TT at Obinvai reprocent. single spaces of the genus Homo and not an aactralopitherine. "The *pecies in, moreover, clearly distinct from the previously recognized species of the genus. But if we are to include the new mitterial the genvis Homo (rather than sot up a distinet genus for it which we believe to bo unwise), 1t becomes necessary to roviee the diagnosis of this genus, Until now, the definition Of Homo has usually centred about a ‘cerebral Rubicon’ Variably sot at 700 ee. (Woidenroich). 750 ¢.c. (Keith) and. S00 e.c. (Wallais}. The proposed new definition follows: oni ax (as dofined by Le (ros Clark, 1955) Linnews, Family Genus Homo Revised diagnosis of the genus Homo. A genus of the Hominidae with the following charactors: the structare ‘of the pelvic girdlo and of the hindlimb skeleton is adapted to habitual eroct posture and bipedal gait; the forelimb is shorter than tho hindlimb; the pollex is ‘well developed and fully opposable and the hand is capable ‘not only of a power grip but of, at tho least, a simple and ‘usully well developed precision grip t; the cranial capacity is very variable but is, on tho average, larger than the range of capacities of members of the’ genus Australo- Pitheeus, although the lower part of Ue Farge of capacities In the genus Homo overlaps with the upper part of the range in Australopithecus; the eapoeity is (on the average) Tange rolative to body sine and ranges from about 600 ce. in earlier forms to more Uhan 1,600 ec.; the muscular the cranium range from very strongly marked to imperceptible, but the temporal erests of lines never réach the midline; the frontal region of the eranium ie without, undue post-orbital constriction (uch as 1s common in members of the gents Awstralopithecus): the Supra orbital region of tho frontal bane is very variable ranging from a massive and very salient supra-orbital ton to «completo lack of any supra orbital projection and a sindoth brow rogion: the facial skeleton varies from. moderately propnathous to orthognathous, but it is not Concave (or dished) az is common in members of the Avstralopitheeinae: the anterior symphyzeal contour varies from a mirked rotrout to a forward slope, while the bony chin may be entirely lacking, or may vary from ‘alight to a very strongly developed mental trigono; tho ental areade is evenly founded with no diastema in most rember of the genic: the first lower premolar ts clearly Licuspid’ with a variably developed hingual cusp; the {Ber the dit of en gp an prose si’ see Nant 3B. ee ee Ba aes NATURE molar Wocth aro variablo in size, but in general are small Fan tion oF han thn ths tra Diliecur, tho saa of Uielact uppor mola is highly variable Int morally smnaltr than the second pe ml and commonly alo smaller than tho frst upper molar; the Tower third molar in sometimes appreciably larger than the fecond; in relation 10 the postion seen in ho Tominoidea 1 0 whol, ho cin sna with ble oro ower. Tapping after the initial stages of wear, but when eompared with thoos of mvynbors of tho xenus Australopihecus, tho miisurs ais! canines are not very small relative to the Inolars and premoiursy tho testh i gona, am partie. lurly the hiolars and. promolars,, are. not. enlarged Inuceo-Tingually ne Uy art in the genus Australopitics tho fist deciduous lower molar shows a variable degren of otarzation Genus Homo Linnwne Species habilie sp. now (Nowe: "The specific name ia taken from the Lal ‘meaning, ‘ablo, handy, mentally skilful, vigorous’. We as indebtel to Prof. Raymond Dart for tho suggestion that Jabilie wonkd be a suitable naine for the new species) ‘A species of the gents Homo characterized by the following Featiro: ‘A mean cranial capacity greater than that of members fof the genus Australopitheeus, but sinaller than that of Homo erectus; muscular ridges on (ho eraninira ranging “yom slight to strongly marked; chin region retreating, Sith slight or no dovelopment! of the mental Unigene; ‘masille and mandibles smaller than those of Austral pithecus and within the range for Homo ereetus snd Homo sapiens; dentition characterized by incisors. which ‘are rolatively large in’ comparivon with hore of both Australopithecus and Homo «rectus; canines which are proportionately large relative to the premolars; premolars ‘which are narrower (in bueco-lingusl breadth) than those of Australopithecus, but which fall within the rarige for Homo erictus; molars in which the absolute dimensions rango between tho lower part of the range in Auttrato pithecus and the upper part of Uh range in Homo erectus & marked tendency towards bueco Lingual narrowing nd mesiodistal elongation of all tho teeth, which is especially evident in the lower premolars (iehore it expresses ito ‘65 9 marked elongation of the talonid) and in the Tower molars (where itis accompanied by a rearrangement of the distal cusps): the sagittal curvature of the paristal bone varies from slight (within the hominine range) to moderate (within the anstralopitheeine range); the external sagittal curvature of the ‘cecipital bone is lighter than in Austratopithceus or in Homo erectus, and lies within Une Fungo of Homo supiens; im eareature as well as in some bother morphological traits, the clavielo resembles, buat ‘not identical Uo, that of Homo sapiens sapiens; the hand from those of Homo sapiens sapiens. in wss, in the dorsal curvature of the shafts of the phalanges. in the distal attachment of flesor digitorum ‘superficial, in the strength of fibro-tendanous inarkingo, tn the orientation of trapezium in the earpus, i the form ‘of the seaphoid nd in the marked dopth of Uo carpal thunnels howovor, the hand bones resemble those of Home sapiens sapiens in the presence of broad, stout, terminal Phalanges on fingers and thumb, in the form of the distal articular surface of tho eapitate and tho ellipsoidal form of tho mets valangesl joint surfaces; in many of foot bones fio within the range of Variation of Homo sapiens sapiens; tho hallux is stout, addueted and plantigrado: dhoro are well-marked longi tudinal and transvorso arches; on the other hand, the Sed 'motatazsal js relatively more robust Uhan itis in. mode rman, sad there 15 no warked difference in tho radi of of the mestial and lateral profiles of the trode Geological horizon Middle Plistocrno, Upper Villafranchian and Lower April 4, 1964 vo. soe ‘ype. ‘ho mandible with dentition and the associated upper rolar, parietal ad hand hones of singh joven individual fom ste F-L-KCW.1, Olhuea, Bet “This i catalogued as Olduvas Hominsd Paratypes. (a) An incomplate craniutn, comprising fragment of tho frontal, parts of both pasetas, the greater part of tho deepal and parts of both temporal {ogether ‘with an sociated sandible, with conines Pimolars and molars compete on ether sido but with te Erowns of the incisors dataged, parts of both maze Kewing ail tho cheek oth exenpt the upper Tet four Piomolar. ‘Tho "conditon ‘of the teat suggest an fuloleseent. This specimen, from site MCV-K.TI Oluvas Bed Ti, i cataloged ss Olduval Horn 13 (0) Th asostated hand once foot bones and prob bly the Gavicle ofan iit’ indvicual fom! te FLAN. 1, Oldwesi, Bel 1. Tis catalogued Gidea Homa 8 {¢) Alower premolar, an upper molar and eranial fog roonts from sto Plo, Ty Ontaval, Bed T (ee ato it Hie als tho Auctratopthocus (Zinjanthromas) skull) ‘This i catalogued an Olden! Homa 8. (tis possible Uhat the tibia and fibula ound ths it belo wth Tome habikis rates tha ith luctrolopthece jam thropus). The lta hones have been ieprtel oh by De PIME"Davis ature, March 3, 1063 p- 967. 2) A manibatarrapment with solar tn position nd susowiatad witha fow fragment of other tect Irom ste SAT, Oldwvai, Bea "this specinn is catalogued as Oda Hominid & Description ofthe type. Preliminary descriptions of the specimens which have now been desist the type of {oma haiti for meant the parts ofthe javena Foe ae tle FLAN. hn 1960" have slteady been pub Hshed in ature by" ono ol vs 189, C40 191, 317; 1961, "A foreher dette Steeipton and report on the parila, thomanibieand the teeth aryin acto prepare, tion by one of us (P. V2) whe is reporton the eas apocity (prowaling aries) ne well as prelainary note gn th hand by another of ws (Natures 196, 409. 1803) iio eon pode Wo donot prays, thertors, of fo a mord detailed doserption of dno typ hare ‘Desert of the prays, "A prlinhacy note on the clavicle anon the foot st the able, which represents paratype (0) was published in Doture (188. 1050, 1901) nds Farther topert‘on the foot hy De M.-H. Dey an Dr. 3. Ke Napier wan published Nature of Mace M6 p68 "The Following addtional preiminary notes on the otter paratypes have ‘been! prepared) by ‘one. of us mv Description of Paratypes {o) Olds Hominid 13 from MAN-K. 11. An adolescent represented hy a early complete mandible with complete fally-erupted tower dentition. right maxillary fropment ‘ncn palate snl teth front to Bis the eto ok process "of erupting tho comeeponving left manny Fragment with 0 to BP, th later hkoreice eruptang. the Halated Toft -P2, parte ‘af th vault of a small duit iting. much of the octal, including port of the posterior margin of foramen maan, parts of Both panietals, right and lft teinporcephenord fragmenta, each Including he mandibular fossa and foramen ovale, ‘The distal halt of 8 humeral shalt (excluding, tho, distel ‘oxtremity) may also belong to Olduvin Hominid 13. Tw Corgan manditndae ie vorg smal both the height ana Uekness at Af, falling below the australopiheoino rans find sith tho'horsine rang All the teat av sl comprred with those oF Anstrslopitheense, mont of the “Timensions falling at or below the lower extreme ef the Sitealopithesine ranges, tn thi other hand, practealy the dental dimensions cin be eeonanorinced within the ange of fossil Homininan. ‘w Oktwrat Hominid 13 teatsnocavar April 4, 1964 show the eharacteristie mesiodistal clongation and inbolingwal ‘narrowing, in some. tooth the EB index ‘txcouding even those of the type Olduvai Hominid 7, and parstype Olduvat Hominid 8. "The occipital bone has Slight ragivtal curvature, the Oosiital Sagittal ng outside the range for abstealopithocines an {or Homo erecius pekinensia nl within the range for Home sapiens. On th cther hand, tho parictal sagittal curvature cre mark than it all but we austrafopithectne snd 1 thr Pekin fossis, Ue index falling ab the top of the sof popistion means for modern man, ‘oth parietal tind occipital bones aro very sina in size, being exceedod in some mensions by oneor tro australopithecine eran tii falling short in all dimensions of the Fango for Joma rectus pekinensis. ‘Tho form of tho pariotal-antero posteriorly clongated and bilaterally narrow, with & faily [Rbrupt lateral descent in the plane of the parietal bore — reproduces closely these features in the somewhat, larger pparotal of tho t3po specimen (Olduvai Hominid 7 from eK ND), (i) Oldurat Hominid Brom BLK. 1. An unworn lower loft premolar, Montifod se P,, an tanworn, practically com plete crown snd partly dovoloped roots of an uppe molar, Miker Af or Ie aa well a'« number of fragments of Cranial vault, "These remains were found at the Zinjan. Uiropussite and level, some in situ and some on the surface, oth testh are small for an australopithecine, eepecially in buccolingual Ureadth, but Tango for Homo erectus. ‘The Iniazked tandency to elongation and natrovring imparts 10 both teoth an L/H index outside the range for ail known sanstratopithocine homologues and even boyond the range for Homo erectus patinensia. "The longating narrowing, tendency ia tore sarked in thi molar than in the upper tui to se Oks Mod Dom PIN. (6) Oldweai Homnndd § from BLIK.N.N, 1. Remains of an adult individu! found on the sarno horizon a the typ Specimen, and represented. by” two complete, proximal Phalanges, a fragment of a rather heavily worn tooth {premolar or molar), and a ct of foot-bones possresing host of tha specializations associated with the plantigrads propulsive fect of modem man. Probably the clavicle found at this site belonga to this adult rater than to the jnvenile typo-speeimen; itis characterized hy clear overall simalaritiqs to the clavicle of Homo supine sapiens. {€)Olduve’ Hominud 4 from MOK. A fragment of tho posterior part-of the left corpus mondibulae, eontaining 2 rll fully eruptad molar, cither Mt, oF My ‘The width of the marutibte fe 192 m towel with the mesial half of the moter, but the maaisnum width must have tasen someoshat greater, ‘The moar in 15-1 sora in tore distal ength and 12-0 mum in buccolingwal breadth; if ‘hor a einatl and. narrow tooth by australopithecine standards, but large in comparison with Homo errctus molar. There are several other isolated dental Fragments Including # motorately wor molar fragment, "These ne ratigeaphically the okleschonsiaid remains yet discovered at Oke the Referred Material Olduvas Hominid 14 from MWK. TI (1) A juvenito topreented by i frapment of tho right parietal with ele lunfused situral issn; vo staallor Vault fragenents with, sutural margins: a lft and a Tight temporal Trazeent, ‘meh sneladings he mandibslar fos, A fragmentary skull with partsof the appr and Mention of a yonne adult from site FLA. H. Mako Gully, Okluvai. Red UL, ts abo. provisionally roferred. to Homo habilis. “his specimen is eatslogunt as Olduvai Homined If 1 is reprecented by the caraplete upper right dentition, as wells somo of the left inaxiltary tert lopether with Sone of the mandibular teeth. ‘The sul Fenmmonts chute patts of the frontal, witly bolle th esternal orbital angles perved, as well is ve spr NATURE 9 orbital rogion, except for tho glabellu; parts of both pariotale and the oceipital are also represented. Implications for Hominid Phylogeny In preparing our diagnosis of Homo habilis, wo have not overlooked the fact that there aro several othor African {and perhaps Asian) fossil hominids whose status may now roquire re-examination in the light of the new discoveries fand of tho setting up of this new species. ‘The specimens “originally deceribed by Broom and Kobinson as Telanthro- [pus capensis and which were later transforred by Robinson, to Homo erectus may sell prove, on closer comparative investigation, to belong to Homo habilis. The Kanara mandibular fragment, «iscovered by the expedition in 1022 by ono of ws (1.8. B. T.), and whieh has been shown ‘to possocs archaic foaturos (Tobias, Nature, 185, 946; 1960), may woll justify further investigation along these lines, ‘Tha Tako Chad craniofacial fragment, provisionally deseribod by M. Yves Coppone in 1062, as'an australe Pithecino, is not, wo ara convinesd, a moxnber of this fub-fomily. We understand that tho diceovorer himself, following his investigation of the australopithocine originals from ‘South fies” and. ‘Tanganyika, nov ‘shares our view in this respect. Wo Deliove that it is very probably a northern roprosentative of Homo habilis ‘Outside Afriea, the possibility will have to be considered ‘hat the tooth and eranial fragments found at Ubeidiyals fon tho Jordan Hivor in Taraol may also belong to Homo habilis rathor than to Australoprthcous Cultural Association When tho skull of Ausiralopithecus (Zinjanthropus) boiget was found on a living floor at F-L.K. I, no remains ‘of any other type of hominiel were known feom the oasly part of tho Olduvai sequonee. It seemed reasonable, {herefore, to ascamo that this skull represented the makers of the Ollowan culture, "The subsequent discovery of romaine of Homo habilis in association with the Oldowan a eae ete pd jon. While it is possible that Zinjanthropus and. Homo habile both made stone tool, iis probable that the latter was the more advanced tool maker and that the Zinjanthropus slesll represents an intracer (or & wietion) fon a Homo habiis living site. "Tho reoent-diseovery of a rough cirele of loosely piled stones on the Hiving floor at site DAT, in the lower part fof Bed TI, is noteworthy. ‘This sito is’ geologically eon temporary with MKT, toss than ono milo distant, where rimsins of Homo habilie have boon found. It seems that the early houninids of this period were eapable of making rough shelters or windbreaks and it i8 likely that Homo habilis may have been responsible Relationship to Austrafapithecus (Zinjanthropus) “Tho fossil human remains representing the new species Homa habitis have been foxnd in Bed Tand in. the tower nd misdte part of Hist TI. "Two of the sites, MK. Land PLAN. 1, are goologically older than that which, yielded the skall of the australopitheeine. Zinjandhroyne. One site, FulK. I, ha yielded. both Australopithecus (Zinjanthropus) and remains of Homoghabilis, whilo to ites age Tater, namely ALN-K. Hand PLR. 1 Maiko ullly. The new mandible of Anstralopsthecns (Zinjanthre. us) typo from Lake Natron, reported. in tho procoding, farticlo by Dr. and Mrs, Leakor, was associated with it fauna of Bod 11 affinities Tt thus seems elvar that two different branches of the Hominidan wero evolving side hy side inthe. Olen fasion durin te Upper Villatranchian and the Tower p Or the Mididly Pleistocene