Literatureof Anthropological (Gouldand Eldredge 1977; Stanley Rnatics

/ /

Species of HCiTLinidS

Jon Marks

Despite some claims tothe time" (Gould 1982a:84)?Obviously, contrary(e.g.,Stanley 1981) ,paleo— we cannot, and the failure to anthropological scenarios have tended nize the severe limitations of toincorporate both anagenesis and hominid taxa has undermined the cladogenesis,though in a "transfor— attempts to impose a punctuational mational"framework. After the framework upon paleoanthropology. initialfocus on stasis andcladogen— esisin the fossil record by Eldredge For example,a chart of the and Gould (1972) ,theseminal attempt duration of hominid species by toplace hominid evolution in such a Stanley(1979:81) presents novel framework (Eldredge and Tatter— Leakeyet al. 1964as a sail1975)unfortunately lineageexisting independently of adiscussion of Dart theoreticalsystematics. 1925 ; and (Leakey 1959) as a lineage existing independently of In the last fewyears, however, (Broom 1938). But this there have beenseveralmajor seems less to demonstrate stasis attemptsby paleontologists to within species thanit does an present hominid.evolutionin a uncriticalacceptance ot taxa in trie punctuated equilibria framework literatureof anthropological (Gouldand Eldredge 1977; Stanley rnatics. 1979, 1981; Eldredge and Tattersall 1982). Punctuated equilibria is a If we wish to discuss tempo and theoryabout the tempo of modein hominid evolution, a funda— cal change; more specifically, it is mentalprerequisite is thedelimita— astatement about the duration of a tionof the fossil species on purely relative to the origin of grounds, in the absence the morphology (Gould l982a, b). A ofthe social, political, academic, key assumption is that the emergence and economic biases which frequently of a morphology is equivalent to the affectpaleoanthropology (Holden emergence of a species, and conse— 1981) . Thisaspect of the tempo—and— quentlythat species (i.e., morpho— modecontroversy has been overlooked species)are formed very rapidly in by both the "gradualists" et relation to their longevity. al.1981) and "punctuationists". The purposeof this paper is to demon— Ifspecies are the units of macro— strate that, given a temporal frame— evol uti on, and i f punctua ted equl I I b— work, the distribution of hominid na is put forward to describe the morphologiesthrough time reveals macroevolutionary tempo of a group of fourpersistent lineages over the species,itis, therefore, of para— last three million years,each mountimportance that the species be presumablyequivalent to Simpson1s clearlydelineated and unambiguously (1961)evolutionary species (Figure defined.For ifthe species them- I). Between lineaqes there are small selves are ambiguous, how can wemake but apparent discontinuities; how— sensible statements about "speciation ever, within each lineage consider— and itsdeploymentin ablemicroevolution and anagenesis, ______- - - - So)nt-Ces&re (Sate'?) Qafzeh Figure 1: Li:oeageg of ChoukoutienNclutuSwanscombe Lontian I MX OH -9 XromdroaiKNM-Ek 406 Stw—53KNM-(R1a13 KNM-EROjetis 3733 OH-5Swartkrans 014-24KNM-ER [I L,neage Neonderthol Erectus— 2 MX UneogeRobust Omo—Shungura E Omo — Shungura E Sterkfontein Omo— Usno MakoponsgatHodarOmo—Slwngura B Austrolopithecinel Radar Lineoaei Basafi 34

rather than stasis, are evident, son and White 1979b; Day et al, Thus, punctuated uilibria is a 1980b; Leakey and Walker 1980; Johan- necessary,but not sufficient, sonand 4lhite 1980),the fossils explanation for hominid evolutionary themselves tell us of a bipedal, patterns: the general usage of the small—brained creature with large theoryshould be extended. canines,a primitive tooth—row, and diasternata(Johanson and White Itis not my intention to give a 1979a), The mode of locomotion was thorough review of the hominid fossil apparently of a somewhat different record here; for that, the reader is naturethan modern bipedalism referred to any good physical anthro- (Jungers1982; Stern and Susman pology textbook. Further,I am 1983), but presumably the capacity exploringthe of evolution as for bipedal ism I s synapomorphous in revealedin the hominid fossil re— and The cord; and patterns are frequently overallaffinities of difficult to discern unless one havebeen suggested as being closest stands back a from the data. to (R.Leakey 197 6) ; Consequently,I shall make use of (Tobias 1981); and second—order in this (Olson 1981). (forexample, using disagreements about thetaxonomicstatusof speci— Although the australopithe— ficfossils as data).The factthat cinedeposits fromHadar, Ethiopia giventwo fossils of complete- were originally dated to 3.0 MYA ness,taxonomic disagreementsmight (Aronsonetal. 1977), they arenow existover one fossil and not over considered to be 2.9—3.6 MYA (Walter theother,is a datuminitself. and Aronson 1982), This makes the Hadar assemblage somewhat younger ficininid Taxa than the Laetoli fossils (3,49—3,76 MYA), andmay throw into question the

I• BasalAustralopithecines homogeneity of the sample (Day 1982; Brown 1982; Boaz et al. , Afterthe eight—million--year—old 1982) . Theprimitive Laetoli ramapithecids from the Siwalik Hills, fossils(M, Leakey et al, 1976; White Pakistan,the picture of the 1977; 1980) certainly resemble some australopithecinetransition becomes of the Hadar remains, yet there seems exceedinglyblurry, Outside ofa little reason to regard the Hadar mandibularfragment from Lothagam, an fossils as a homogeneous assemblage arm fragment from (Patterson (e.g.,Behrensmeyer 1983). etal, 1970), a tooth from Lukeino (Pickford et al, 1975), and perhaps a Thatthe Laetoli and part of the facial bone from themeron (Bishop and Hadar finds represent a species Chapman 1970), the fossil record is ancestralto later australopithecines silent until later than 4 MYA. canhardly be disputed (White et al, 1981). Tobias (1981), however, Tobias (1973) had predicted a argues that the range of variation basal australopithecine lineage for will encompass the

( tohave collection, a point lived about 4 Mm, and this was deniedby Johanson and White (1979b), subsequentlyfulfilled with the dis— If we accept the inhomogeneity of the coveryof fossils from Ethiopia and Hadarsample (Olson 1981) andtake it Tanzania collectively referred to torepresent (with Laetoli and Omo— (Johanson Usno;Howell 1978) and et al. 1979), Although considerable (withMakapansgat [Boaz et al. 1982]) taxonorni c controversy has surrounded A.. much of the systematic these fossils (Anonymous1979;Johan— ambiguity would probably vanish. 35

2. Gracile Australopithecines guishthe two genera. consequently, identification of the skulltends to Dating from about 3.0—1.4 MYA, beconfusing: Walker and Leakey thereap ars a collection of fossils (1978)call the skull australopithe— stilldebated taxonornically, but cine, and Delson (1979:511) broa dly conti nuous I n rn ol ogy ,and call attention to both the and probablyrepresenting a single affinities of the evolutionary skull, and Wolpoff (1980:164) identifies the skull as a female Gracileaustralopithecines are dated from Tuff B of the Shungüra formation in theOrno at 3.0MYA Finally,the fragmentary OH—13 (Curtis 1981), The South African fromlower Bed II, Olduvai Gorge is a australopithecines are slightly paratype for (L.Leakey younger, but notorious in their et al. 1964). Nevertheless, it has inabilityto be securely dated.Most been seen as similar 'tothe very recentestimates, however, have the earliestdental fragments fossils from Makapansgat at nearly thropus")from Swartkrans (Tobias and 3.0 MYA, and those from Sterkfontein Von Koenigswald 1964). Similarly, about MYA (Tobias 1981). the dentitionhas been likened to the Asian "Meganthropus" Slightly later, fossils attributed (Tobias and Von Koenigswald 1964), to appear in EastAf rica which others have called a robust (L.Leakey et al.1964), Olduvai australopithecine(Robinson 1955). hominid24 isfrom Bed I, about 1.9 Thisambiguity, however, is most MYA, and East Turkana's KNM—ER 1470 likely illusory,for the total is 1.87 MYA (Gleadow 1980). morphological pattern of these The latter skull is variously hominidssets them readily into fled as or distinctlineages.The apparent depending upon the author (e.g., ambiguity arises from the fact "that Walker and Leakey 1978). Fragments teethandmandibles alone cannotbe of "early arealsoknownfrom used to distinguish australoçithe— later deposits at Sterkfontein cines and without (HughesandTobias 1977). points associatedcranial material, except are worth notingaboutthis taxon, at the extremes of the ranges" however: 1) nearly alläpecimens (Wolpoff 1980:189). placed in are either crushed, highly fragmentary, or There is thus a continuity of form ture;and 2) the signal difference withinthe gracile lineage, and a between this and is the smalldiscontinuity of form between larger cranial capacity in the gracile lineage and its aholdover from the antecedent and descendant taxa (cf, "cerebralRubicon" days of paleo- below).This has led Pilbeam (1975) anthropology. torecognize "a period of relative stasis...from3 to 1.5 million years" I do not wish to affirm or deny in this group. the validity of the taxon here, butmerely to emphasize 3.Robust Lineage thestrong morphological continuity of this with Thus, The robust and gracile lineages specimen —1813from Lake Thrkana is are readily separable in terms of arelatively complete skull showing theirtotal morphological b a—likedental—maxillary withminor exceptions.The relative- features,yet the cranial ly 3ate (1.6 MYA) deniicranium KNM—ER capacitygenerally used to distin— 732 has been called a female robust 36

australopithecine by Walker and when the variation is ina single Leakey (1978) and Wolpoff (1980); character! although Day (1977) has seen it as a gracile. The problem is that the Thus,Stanley(1981:143), specimen possessesa large supra- zingthe incorrecthess of seeingmore mastoidcrest, though little of it is than one distinct species of robust present; yet it has no sagittal crest australopithecines (cf, Stanley and only a single fragmentary tooth 1979:81), incorporates them as a crown —butthese are the features single lineage in his second punctua.— generally used to identify robust tionalschemefor the hominids, australopithecines. Alsoperhaps insecurein its identification is the Robust australopithecinesare Taungchild, because ofits known from thesowanja, about 1,4 MYA ininaturity. (Gowlettet al, 1981), and slightly later atEastTurkana, Throughout Theearliest robust their duration, they do notapar to cines seem to be present at about 2 demonstrate any anagenetic trends, MYA at the Orno(Curtis1981). Again, onthe other hand, consistent theSouth African fossils are evolutionaryvariation between the cult todate, but faunal correlations South and East African forms is suggest that the material from. apparent, Kromdraai andSwartkrans are contem— poraneous withthe anatomically more extremeEast African counterpart. Erectus—NeanderthalLineage Althoughthe East and South African are usuallyclassified as distinct at Theonset of the Pleistocene the species level, it is unlikely coincides with thedebut of thatsuch distinction correspondsto (Dubois 1892) aspointed out anysignificant biologicalreality — byDelson (1981). These horninids, thevariation is quantitative and quite distinct from the slight andcan scarcely be considered ecinesin skull morphology, never the— morethan ecotypic.One possible way less are broadly similar inm y ways of resolving this taxonomic problem tothe later Neanderthals, They are isto view the lineage as a bothcharacterized by a thick skull superspeciescomposed of seinispecies vault, pronounced post—orbital (Tobias1973; Delson et al.1977; constriction and massivesupra— Delson 1978). But if we apply orbital tori, skeletal robusticity, Simpson's (1961) definition of an anda characteristiclong, low skull evolutionary species ("a lineage with prominent occiput. evolving separatelyfrom others and with its own unitary evolutionary Thesederived characters (i.e. not role and tendencies"), it seems foundin australopithecines)are unlikely that more than one role and diagnostic of both and tendency is represented here. It is theNeanderthals and their collateral therefore unlikely that specific relatives might, with little danger discrimination within the robust of exaggeration, be consideredas a australopithecines is warranted, A with an inflated brain" contrary view, however, is expressed (Eldredge and Tattersall 1975). The by Grine (1981), who sees three minor shape changes which differen— species of robust australopithecines tiate from Neanderthals by basedsolely upon enamel prism criteria other than size may be patternsof teeth, However, the mere accountableas 1)allometric effects discoveryof between—qroup Variation resultingfrom the size changes, 2) is not sufficient to establish the anagenetic responses to natural groups as goodspecies especially selection, and 3) microevolutionary 37

divergence of populations, These Jaeger (1981) alsopoints the features include maximum skull ambiguous nature ofthis specimen; breadthand greater facial height in yetit is late andvery small and theNeanderthals.Suchdifferences, strongly biases Rightmir&s however,are dwarfed by the coristel- caltreatment. Niother questionable lationof cranial features shared by inclusionis thatofHolloway's the two groups. (1973) low estimate of 727cc for the late and very fragmentary OH—12. Thesystematic problems in dealing Indeed, this is the seventh Rightmire withthistaxon may be related to the groups with the six from thoukoutien factthat theconcept of lumping many at0.6 MYA, which range form915-1225 groups of fossils into a single taxon Yet even Holloway (1973)is of althoughinformally unabashedlyskeptical of the estimate proposed by Weidenreich (1940), of 727 ccfor thisspecimen; and suddenly gained and uncriti- given that estimates for cal acceptance when proposed by Mayr ly more cqrnplete specimensmay very (1951) a neontological 10% (see Day 1977:63 for estimates gist. ofSteinheim),itis probably unwise tohave let the conclusions ofhis The earliest (and most complete) statisticaltreatment be unduly -. is (1.5 biased by such fragmentary material, MYA)from Lake Turkana, but the If the analysis is correctedfor earliest fossils from Javaare thesesproblems, an anagenetic trend probably nearly as old (Howelis in cranial capacity becomes highly 1980). Olduval hominid 9 is from evident, in fundamentalagreement UpperBed II, dating to about 1.2 witha previous study by Bilsborough MTh. An early fossil from (1976). Lantien has been dated to MYA (JiaLanpo1980). Later fossils from Atabout 0.4 MYA, we begin to theTrinil Beds inJava, choukoutien, encounter several fossils whichseem andLake Ndutu (Clark 1976; Howells to fall on the border between 1980)all seemtofall about and theNeanderthals.Thus, MYA. thelineage continuesin the large, heavy, specimens from These fossils show Ngandong. Although identified by variation geographically(Coon 1962), some workers (e.g., Von Koeningswald However,the evolutionary patterns 1962)as Neanderthal, they are also areasourceofcontention. widely accepted as Rightrnire (1981) has claimed that SantaLuca 1980),Similarly, the thereis statistically no anagenetic Petralona skull has affinities with trend evident in skull size in both groups (e.g.,Trinkaus and His treatment, however, is Howells 1979), as do Arago (de Lumley flawedinseveral First, andde Lumley 1974);Bodo(Conroy et Rightmireregressed skull size on al,1978); and other skulls suchas time yetrather thanentereach of Broken Hilland Salclanha(Coon 1963; the 24 points as data, he instead Howells1973).The major character regressed merely the mean values for linkingthese to isera— each time interval,reducing his nialcapacity, yetanatomically, analysisto6 points.Thus, the Sale theseare all continuous with one skull,as theonly one at 0.2 MYA, is anotherand with the givenweightequal to the 7 skulls at sample (cf. Krantz 1980;Merrill 0.6 MYA. Yet Wolpoff (1980:225) 1982), focusseson the rounded contour of the occiput as evidence that the Itwould seem, therefore,thatthe skull is an early archaic most parsimonious explanationforthe 38

evolutionary patterns observed here Saint—Cesal re I inds (1ndeed, si ightly involve a single, widely distributed antedatingthem) are remains of lineage, changing anagenetically anatom I cally moder n m an w i thtypi ca 1— throughtime and exhibiting extensivelyNeanderthal (Mousterian) tools; microevolutionaryvariation. for example, at Qafzeh. This bio— logicallineage can be tracedback to Although Delson (1977 and pers. "archaic" modern forms from the comm.) has argued that the traits Kibish formation of the Omo (Howell shared by andthe 1978), the Ngaloba formationat thals are merely symplesiomorphous, Laetoli(Day et al. 1980a), and this is a tautology. The Border Cave, each of which are thalssucceed temporally whatolder than 100,000 years. withlittle, if any, ambiguity. Yet interestingly, there are no Specimens if two populations arephylogenetic— thrustback and forth between allyancestor and descendant, it Neanderthaland Modern Man in the necessarily follows that any same way thatspecimens are inheritedtrait they hold in common ambiguously identified as Neanderthal mustbe a conservative retention in or the descendants. However, those diagnostictraits which Neanderthals, Further, there are two independent share with are clearly suggestions of a rapid and abrupt derived in theancestralpopulation, formation of a new species slightly therefore, these traits before the dates attributed tothe canbe used to establish the two (or three early modern fossils. The more) populations as closely related firstis through a critical examina— phena. The view I am advocating tion of Middle Pleistocenehominid involves simply the recognition of morphologyby Rightmire (1981) ,who theunity of these thick—skulled, concludedthatthe morphological platycephalichominids as a single suite characteristic of ourspecies continuous lineage. Two factors, 1 wasformed "during a short pulse of believe, favorthis over alternative evolution, late in the Middle explanations: I) the relative ease Pleistocene." The second is through with which the earliest the examination of mitochondrial may be diagnosed from contemporaneous inliving populations, under the populations of australopithecines, assumption of a molecular clock and 2) the existence of biostrati— (Brown1980). That study concludes graphic and morphological inter— that a population crashoccurred mediates linking within the hominid lineage between Neander thai s. 0.36 and 0.18 MYA, whichmayroughly date the appearance of the lineageof The culmination of the microevolu— modernpeople, tionary trend would obviously be in the classic Neanderthals of Europe The relationship of modern man and Western Asia (O.l—.035 MYA). The with the Neanderthals andotherUpper recent discovery of Neanderthals Pleistocene hominids has traditional- associated with cultural implements lybeen the subject of extreme of Chatelperronian type atSaint— disagreement among anthropologists Cesaire (Leveque and Vandermeersch (e.g., Brose and wolpoff 1971; 1981) gives a last glimpse of this Howells 1974). I suggest thata lineage about 34,000 years ago. majorcause of this isthe failureto recognize that as currently defined with two sub— Modern species, is a grade,determined largely by cranial capacity. it is Roughlycontemporary with the also instructive to recallthat 8

historically the uncritical replacement through Tabun Cave, acceptance of Israel, Perhaps in the systematic and as framework I propose here, many of the comprising a single species is due to prOblemswhich hereto— Dobzhansky. (1944), Dobzhansky, like fore In the recent evolutionary Mayr for , wasa profes-- patterns of the Hominidae will be sional student of neither fossilsnor shownto be spurious. primates,Neither Doazhansky's nor Mayr's species were frameddiachroni— cally, in terms of the linkage of Discussion ancestral and descendantpopulations, asSimpson's species were, withan Apopulation crash leading to eyetowards linking populations severe genetic "founder effects" in temporallyseparated. outlyingareas of a sdes' range is and are the most likely causeof anappar— more closely linked by virtue of entlypunctuational event (Mayr sharing the same Bauplan,or total 1942:234ff.,1954; Carson 1970; morphological than are Eldredgeand Gould 1972). In the and absence of such demographicproces— ses,a widely—distributed species shouldtend to change gradually Both throughtime, in accordance with are descendants of a selectivepressures, without signifi— Pleistocene stock of cantmorphological discontinuitjes, "Neanderthalojds"or "Collaterals" — The hominids seem to display bcth of representedby such specimens as theseevolutionary patterns,On the Steinheirn,Broken Hill, Saldanha, onehand, there are lineages within etc.. Subsequently, a new lineage of which change gradually: thinners- these are marked throughout by vaulted, hominids "transitionalfossils,"whichhave arosenear the beginning of the Upper provendifficult for anthropologists Pleistocene, while the ancestral form toidentify unambiguously.On the continued, The divergence between otherhand, these lineages tend to thetwo lineages became more marked originatesuddenly, such that the throughtime,such thatStringer earliestmember of a new lineage can (1974)could argue forcefully that be readily distinguished from the there is no close cranial relation— later members of the ancestral ship between the late Neanderthals lineage. andmodernpopulations, However,among the hominids, itis It is important to recognizethat quite difficult to state that these whatis generally considered to be punctuati on events represent "rapid"speciation in paleontology is tion, in any biologically meaningful adivergence on theorderof 50,000 sense of the term, What ispunctua— years, The divergence, indeed the tionalis the appearance of anew duration, of the modern lineage is of morphology, not necessarily the the sameorderof magnitude as a emergence of reproductive isolation "rapid"speciation event in (cf, Delson 1981). Consequently, logy. As a consequence, it is likely although the Neanderthalsare that the hominid record would reveal from Moderns in their dataon the process of divergence of morphology, it is unlikely that such thes des and replacementof one by distinction merits the theother, Smith(1982)traces the of a biological species latterprocessfor Eas€ern Europe, (Trinkausand Howells 1979) : the andJelinek(1982) traces cultural emergenceof modern man is genìerally 40

0 regarded isrecognized,it is diff i— as a product of cultto argue that the specimens tion, not macroevolution. attributed to it are not at least strongly continuous This is a problem not of (Cronin et al, logy, nor of the hominid fossil with 1981)and part of the same evolving record itis a problem stemming from thefundamentaldisharmony lineage. between the conceptsofthe "biologi— Thenecessity of seeing rapid cal" species and the "evolutionary" speciation in the fossil record species. The former is a reproouc- punctuation) as biological tive community, definedbyfertility (i.e., at a single time; the latter is an speciationis also called into defined by question.Although certain events ecological community, robust anatomical continuity through time. (e.g.,origin of the lineage) Although the Neanderthals and con— could well have involved the estab— lishrnent of a reproductive barrier temporaneousModernsmight have been inter—fertile, it is clear that a betweenthe ancestral and descendant other events (e.g., origin distinctecological gap separated the lineages, two populations.This should merit a of the modern lineage) are unlikely recognition of the two populations as tohave been speciation in the strict separateevolutionary species. On sense, or biological speciation. the other hand, no such ecological gap is evident between any two con— I therefore suggest that with teniporaneous populations of Middle systematic framework proposed here, thetempo and mode of hominid Plei stocene hominids. evolution may be explored with considerably ambiguity. My Further, it is difficult to argue less acknowledgmentofrapid andabrupt forstasis being the rule within each change between lineages isin persistentlineage of horninids. Although the establishment of a disagreementwith the extreme pattern seems to occur gradualist analysis of Cronin et al. rapidly in this family, within each (1981);however, I believe there is lineagethere may be considerable clearly considerable microevolution— arydivergence andanagenesisvisible anagenetic change evident.Thus, we contrast with the havestrong morphological continuity withinlineages, in (but not stasis) within the viewsof lineageand Stanley 1979; 1981; Godfrey and Jacobs 1981;Rightrnire 1981). withinthe lineage despitea relative lack of If the lineagesI recognized continuitymarking the origin of each (as lineage. here are consideredto be Ibelieve themtobe),the proper nomenclatureisas presented in Figure Conclusions 2. It has obviously not been my intentionto provide an exhaustive treatment of the hominid fossil record in this Rather, I have inted out gradations and discontinuities in the hominid record,which I have triedto place inan operative systematic framework, Consequently,regardless of the taxonomic level at which / 1' 41

Horninidae(Blyth1875) •. . .

Genus (Dart 1925) (Johanson, White & Coppens1975) (Dart 1925) (Dart1925) (Leakeyet al,1964)

(Broorn 1 93 8) (Broom 1938) (Leakey 1959)

Genus (Linnaeus 1758) (Linnaeus1758) (King1864) (King1864) flit 1892)

Figure 2. Proposed nomenclature for family Hominidae,

I wish tothankthefollowing for theirhelpfulcomments onearly versiOns ofthis paper:Steve Zegura, Eric Delson, Todd Olson, LaurieGodfrey,Karl Flessa,andM,E. Morbeck. 'I'T;

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