Bull. Soc.roy. belgeAnthrop. Préhist. 97 : 33-44,1986
Homo erectus: an old specieswith nervproblems
by
M.H. Dey
Summary
Thehistory of thediscovery of Homo erectusgoes back for almost100 years to the work of Duboisin Java.Subsequent finds in Chinaand North Africa expandedthe sample.Recent finds from EastAfrica (KNM-ER 3 733,3883 ; WTl5 000),Europe (Arago, Bilzingsleben)and India (Hathnora) haveprompted a reappraisalof the speciesHomo erectusand new work on the Trinil remainsfrom Javahas questioned theantiquity of theTrinil I femur.The questionraised by thesenew finds in relation to the origin, the distributionand the fate of Homo erectusare reviewed.
Résumé
L'histoirede la découvertede l'Homo erectusremonte à presquecent ans avec les travauxde Duboisdans l'île de Java.Les trouvaillesqui ont suivi, en Chineet en Afrique du Nord, ont étendul'échantillon. Les découvertesrécentes en Afrique orientale(KNM-ER 3733,3883; WTl5000), en Europe (Arago, Bilzingsleben) eren Inde(Hathnora) ont incitéà faireune réévaluation de I'espèceHomo erectustandis queles nouveaux travaux sur les restes de Trinil provenantde Java,mettent en doute l'antiquitédu fémurI deTrinil. Lesquestions soulevées par ces nouvelles découvertes et relativesà l'origine, I'extensionet le devenirdes Homo erectussont passéesen revue.
Thebeginning of the storyof the discoveryof whatis now knownas Homo erectus took place in Java in 1890-1891when EugeneDubois, the anatomist turned soldier/physicianrecovered the Trinil calotteand Trinil FemurI from thedeposits of the Soloriver. He later namedthe creaturePithecanthropus erectus (Dubois 1894) from his convictionthat the calotteshowed features intermediate between ape and manand that the functionalanatomy of the femurwas indicative of bipedalism.This latter point has beenconfirmed time and time again by anatomists(Manouvrier 1895;Hepburn 1897; Weinert 1928;Weidenreich l94l ; Le Gros Clark 1964;Day and Molleson1973 ; Lamy 1984).Later descriptions of the finds from China(Zhou- koudian)by Black( l93l ) andWeidenreich ( 1936 , 1937and 1943) confirmedrhe cra- nial featuresof P. erectusbut did not confirm the postcranialfeatures as exemplified by the Trinil femur (Weidenreichl94l ). 34 Sociétéroyale belge d'Anthropologie et de Préhistoire
q \
-6 -5 -3 -2
Fig. l. - Canonicalvariate plot of the relativequantities of heavyelements detected by X-ray microa- nalysisin fossil material from Javaincluding the HthecanthropusI calotte, the Trinil femora (I-VI), the Solo remainsand a sampleof Elephasfrom KedungBrubus (KD). Trinil femur I is highly significantly different from all of the other material sampled.
Recentlythe discoveryby Bartstra (1932) that the Kabuh depositsin Java that containedthe Trinil fauna are overlaidat Trinil by youngerTerrace deposits (c. 0. I m.y. B.P. ) providesa plausibleexplanation for this anatomicalconfusion. It seems clear now that the Trinil femur on which Dubois dependedfor his assertionof upright posture and bipedal gait was in fact an intrusive Homo sapiensfemur from a much more recentperiod. Analytical studieson the Javanmaterial confirms that the compositionof Trinil Femur I is different from that of other Javan remains (Day 1984,1986) (fig. I ). The descriptiongiven by Weidenreichfor Homo erectus femora from China (Weidenreichl94l ) hasbeen confirmed from Olduvai (Olduvai Hominid28) (Day l97l), KoobiFora (KNM-ER 737,t47z,l48l ) (Day 1978;Ken- nedy 1983)and now from WestTurkana (\ryT 15000).The pelvicfeatures described for O.H. 28 (Day l97l) have also been observedon material from Koobi Fora (KNM-ER 3228) (Rose 1984) and Arago (Arago XLIV) (Day 1982;Sigmon 1982). Togetheras a femoropelviccomplex it may be distinctiveof Homo erectusor at the Homo erectus: an old specieswith new problems 35
very leasttypical of a segmentof a palaeospeciesof that name. It is indeedironic that the specificname erectus given to Dubois' find Pithecanthropuserectus should be basedon a Homo sapiensfemur ! After the discoveriesof specimensnow attributed to Homo erectusin Java (Tri- nil, Sangiran),China (Zhoukoudian)and North Africa (Ternifine), a picturehas slowly emergedof a group of Asian and African homininesof the Lower and Middle Middle Pleistoceneperiod whosemorphology and temporalranges seemed to be clearly defined. Sincethat time the picture has changedradically with new finds attributed to Homo erectusfrom East Africa (Olduvai; Koobi Fora; West Turkana), North Africa (Salé), Europe (Bilzingsleben)and India (Narmada). More controversiallyother remains,both long known and relativelyrecent finds, from Mauer, Vértesszôllôs,Arago and Petralonahave entered the discussion.A detailedreview of much of the materialhas been given by Howells( 1980).No lon- ger is Homo erectusa clearly definedtaxon, temporally, morphologicallyor even geographically.A brief review of someof the new finds and the new researchis givenhere.
JAVA
Todaythe picturein Javaremains confused chronologically (Pope 1985)and fau- nally (Groves1985). It seemslikely that all the hominid materialfrom Javais less than 1.3m.y. old (Pope and Cronin 1984)and that thereis no tool culturefirmly associatedwith Homo erectusremains (Bartstra 1985).In addition the Ngandong fossils are regardedby some as belongingto Homo erectus(Santa Luca 1980). Taxonomicallythe hominid record of Java seemsas confusingas ever (Sartono, 1985)although many would attributeall of the hominid materialto one or other form of Homo erectus.
CHINA
The recoveryfrom Zhoukoudiânin 1966of new partsof a skull known to Wei- denreichis pleasing.The two frontal and parietalfragments fit on to castsmade by Weidenreichbefore the Zhoukoudianremains were lost. This is a completevindica- tion of the authenticityof the original finds and a confirmationof the reconstruc- tion on to which the new fragmentsfit perfectly. In 1980the recoveryof the Hexianskull confirmsagain the natureof the Chinese form of Homo erectusfrom a site dated at about 150-400000years B.p.
NORTH AFRICA
An important new skull was recoveredfrom Saléin North Africa from deposits datedat about400000 years B.P. (Jaeger1975,1981). Many of its featuresindicate 36 société royale belge d'Anthropologie et de préhistoire
that it belongsto Homo erectusbut it has a peculiarity of its occipital region that pro- bably precludesthis important areafrom considerationin termsof its taxonomicdia- gnosis.
EUROPE
In Europe remains of Homo erectushave beenreported from severalsites, some of them relatively new. A thick skull with an angulatedocciput was recoveredfrom Bil- zingslebenin Germany and is dated at about 228000years B.P.: it has beenattributed to Homo erectus(Vlëek 1983a and b). The Arago remains from southern France, perhaps400000 yearsold, show both Homo erectusand Neandertal featuresas does the Petralonaskull from Greeceof much the sameaee.
EAST AFRICA
EastAfrica hasproduced at l-1. 5 m.y. 8.P., Olduvai Hominid 9, a clearexample of Homo erectus; and also from Koobi Fora KNM-ER 3733 and 3883, two older skullsat 1.8 m.y. B.P. alsoclaimed to representHomo erectus.An interestingskele- ton KNM-ER 1808from 1.4-1.6m.y. B.P., alsofrom Koobi Fora, showshypertro- phic periostitis, which when stripped from the femur revealsthe Homo erectusmor- phology underlying the disease(V/alker, pers. comm.). The pelvic remains of Homo erectus from East Africa range in time from ' 1.9 m.y. B.P. (KNM-ER 3228)to c. 1.6m.y. B.p.(wr 15000)and to c.0.6-0.8m.y. B.P.(O.H. 28). The new skeleton from West Turkana ( WT l5 000 : fig. 2) is a remarkable find ar 1.6 m.y. B.P.: a skull reconstructedfrom 70 pieces,all the teeth presentand a vir- tually completepostcranial'skeleton. From a brief examinationof the castsof the pelvesand femora it seemsclear that the distinctive group of femoropelvic features outlined elsewherefor Homo erectusare present in this find, although not as well developedas they would have been had he reachedmaturitv.
INDIA
On December5th 1982,Arun Sonakia recoveredthe first example of Homo erec- /as from the Indian sub-continent.The skull was found at Hathnora in the Narmada valley, Madhya Pradesh. The well preservedcalvaria shows numerous features of Homo erectusas well as somethat are advancedsuch as a tall cranial vault and a cra- nial capacitythat may lie between1155-1421cc. (de Lumley and Sonakia 1985).It is associatedwith an Upper Acheulian industry with hand axes and cleavers. What are the morphological characteristicsof Homo erectus that can provide some basis upon which to make a differential diagnosis? The early descriptions of what is now widely known as Homo erectuswere basedon the material known from Java (Pithecantropus), China (Sinanthropus) and North Africa (Atlanthropus). Homo erectus: an old specieswith new problems 37
Fig.2. - The skeletonof a Homo erectusboy of 12years of agefound at Nariokotome,West of Lake Turkana,Kenya (WT 15000).Photograph by courtesyof RichardLeakey, Director of the NationalMuseums of Kenva. 38 société royale belged'Anthropologie et de Préhistoire
These descriptions and others of later material from East Africa include the follo- wing featuresthat epitomise a Homo erectusskeleton : a low and long cranial vault with flattened frontal and parietal bones, an angulated occiput, a strongly marked and continuous supraorbital torus, a small mastoid processwith a marked supra- mastoid crest and a torus angularis on the parietal bone. The vault also shows a frontal and parietal sagittal keel with parasagittal flattenings, a ((tent-shaped > coro- nal section,a low maximum skull breadth betweenthe supramastoidcrests as well as a low triangular squamousportion of the temporal bone. The vault bonesare thick, there is pronounced postorbital constriction and the inion and the opisthocranion coincide. The cranial capacity is between 700-1225 cc. The face of Homo erectusis lesswell known but it is prognathic with a large inter- orbital breadth and broad nasal bones. The mandible has no chin, a narrow but rounded dental arcade, thickening of the mandibular body, a large bicondylar breadth and in many casesmultiple mental foramina. The teeth are of variable size but are in general robust with shovel-shapedincisors and basal tubercles. Molars and premolars have a cingulum and the molars have a dryopithecine cusp pattern that has a tendency to transformation to the < plus > pattern by metaconid reduc- tion. The third molar is often reducedin sizewhile the permanentmolars, premolars and milk molars show some degreeof taurodontism. The postcranial bones were poorly representedat first but later finds have impro- ved the sample. The femora are unusual in the great thicknessof the cortical bone, they are platymeric, have a low narrow point and a prominent convexity of the medial border of the shaft. The pelvic morphology is also distinctive in having a large acetabulum, a stout acetabulocristalbuttress and laterally rotated ischial tube- rosities. The recognition of this femoropelvic complex of features (Weidenreich, l94l; Day, l97l) has disposedof the view held earlier that Homo erectuswas essen- tially sapiensin its postcranial features. This general description includes many features referred to by others who have attempted a definition of the speciesHomo erectus (Weidenreich 1943; Le Gros Clark 1964; Howell 1978; Howells 1980; Day and Stringer 1982;Rightmire 1984; Stringer 1984; Wood 1984) but no attempt has been made here to distinguish bet- ween characters of common inheritance and new or derived features unique to Homo erectus.Those who have sought such unique featureshave not been too suc- cessful(Wood 1984; Andrews 1984; Hublin 1986; Bilsboroughand Wood 1986). The derived features that are usable for a diagnosis of the taxon are very few and their apparent concentration in the Asian specimenshas led to doubts about the existenceof Homo erectusfrom African sites(Andrews 1984); thesedoubts are not entirely sharedby Bilsborough and Wood ( 1986). Hublin goeseven further and sug- gests that from a purely cladistic point of view Homo erectus does not exist as a taxon, but as a grade it can be defined by featuresprimitive to Homo sapiensparti- cularly if evolutionary stasishas occurred. This agrees(perhaps for different rea- sons) with the view taken earlier by Jelinek (1978, l98l) and Thoma (1973) who argue that Homo erectus should be sunk as a taxon into Homo sapiens. Homo erectus : an old species with new problems 39
A more widely held view seesHomo erectusas a palaeospeciesor chronospecies that will show evidenceof evolution through time from a more primitive ancestor, such as Homo habilis, to a more advancedsuccessor such as Homo sapiens(Le Gros clark 1964; campbell 1972;wolpoff 1980, 1984; Howells l98l; Day 1984). The questionof evolutionary rates,of the punctualist versusthe gradualist models, in the hominine fossil record has been raised in relationto Homo erectusby those who support the punctuational model (Could and Eldredge 1977; Stanley 1979, 1981;Eldredge and Tattersall 1982). Homo erectusis seenby theseauthors as a ( true ) taxon that did not vary greatly in form during its existence.Some evidence has also been presentedfor this view from the postcranial skeleton (Kennedy 1983, but seeTrinkaus 1984; Day 1982). Wolpoff ( 1980, 1984) contends that significant evolution within the taxon can be determined by the examination of cranial, dental and mandibular features.The full descriptionand analysisof the new find from West Turkana will do much to underpin, or otherwise, the contention that Homo erectus was in existencefrom as early as 1.6m.y. B.P. at that siteand lasteduntil perhapsas recentlyas 0.3 m.y. B.P. at other sitesin both Africa and Asia - a spanof more than 1.0m.y. The erectus/sapienstransition has also been examined in relation to the Omo I and Omo II skulls from Ethiopia by Day and Stringer ( 1982) who conclude that these two, supposedlycontemporaneous skulls can be aligned with mod ernHomo sapiens and Homo erectus respectively, or Omo II included in an < archaic >>Homo sapiens group that displays a suite of mosaic or intermediate characters. The existenceof a <
It is clear, therefore, that the taxon Homo erectus is under intense inquiry in terms of its geographicrange, its temporal range, its origin and its evolutionary fate. It is also the subject of discussionin relation to lhe evolutionary models of punctua- tion and gradualism as well as the taxonomic approachesof the cladistsand the gra- dists. As in most debatesthat are pursued with great vigour and determination it is possible,indeed probable, that the truth will lie in part with all the contenders.Only time and researchwill provide the answers.
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Adressede I'auteur.' Prof. M.H. Day Departmentof Anatomy St.Thomas's Hospital medical School LambethPalace Road LONDON SEI 7EH GREAT BRITAIN