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The status of heidelbergensis (Schoetensack 1908)

Article in Evolutionary Anthropology Issues News and Reviews · May 2012 DOI: 10.1002/evan.21311 · Source: PubMed

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ISSUES

The Status of (Schoetensack 1908)

Chris Stringer

The Homo heidelbergensis is central to many discussions about two clearly do not articulate . In recent . For some workers, it was the last common ancestor for the early 1980s, with a shift to cladis- the subsequent species Homo sapiens and Homo neanderthalensis; others tic thinking and influences, I began regard it as only a European form, giving rise to the Neanderthals. Following the to gravitate toward the idea that impact of recent genomic studies indicating hybridization between modern Neanderthals were, after all, a dis- and both Neanderthals and ‘‘’’, the status of these as sepa- tinct species from Homo sapiens rate taxa is now under discussion. Accordingly, clarifying the status of Homo sensu stricto, and that this implied heidelbergensis is fundamental to the debate about modern human origins. the existence of a distinct ancestral species, if neanderthalensis and sapi- ens were sister taxa, and erectus did not represent the last common THE SPECIES HOMO was little used during the earlier part ancestor. Through linking Mauer HEIDELBERGENSIS of the twentieth century and, by the with Petralona and Petralona with 1960s the lumping together of taxa Broken Hill, the concept of a Eura- In 1907, the robustly built mandi- often treated the as a European frican stem species named Homo hei- ble that was to become the holotype 2 form of . However, delbergensis began to develop.9 The of Homo heidelbergensis was discov- 3 Howell took exception to that, argu- following extract and accompanying ered in the Grafenrain sandpit at ing that the fossil probably was mor- figure (Fig. 1) summarize the cau- Mauer, near Heidelberg, , phologically distinct enough to repre- tious arguments made at that time: associated with what is now termed sent a separate species. ‘‘Because at present they cannot be a Galerian or Cromerian (early Mid- In 1974, I completed my doctoral defined satisfactorily by their own dle ) fauna. The species thesis, which concentrated on cranial distinctive within-group characteris- name was bestowed a later by shape comparisons of tics, it is difficult to justify creating a Otto Schoetensack,1 who noted in and modern humans but, along the separate taxon for the Petralona and the Mauer the combination way, I noted clear phenetic resem- Broken Hill on the basis of of primitive features (for example, blances between the Broken Hill characters they lack, or ones they high corpus thickness, very wide () and Petralona () fos- share with other taxa. Nevertheless, ramus, and receding symphysis) and sils, and considered both of these to given the need to recognize their more recent human features, such as be clearly distinct from Neander- similarities to each other, and to small dentition, particularly the can- thals.4,5 Rather than allocate either other Middle Pleistocene fossils, they ines and anterior teeth. The name of these specimens to Homo erectus, could be placed in a separate species, I preferred, at that time, to regard H. rhodesiensis or H. heidelbergensis them as related primitive forms of (if the Mauer mandible is also Homo sapiens sensu lato, eventually included), provided the distinctive- has worked at The Natural History Museum London since 1973, and assigning them to Homo sapiens ness of the Neanderthals from is now Research Leader in Human Ori- grade 1 in a gradistic scheme.6 ‘modern’ H. sapiens is also consid- gins and a Fellow of the Royal Society. He has excavated at sites in Britain and Following discussions with Bjorn ered worthy of specific recognition abroad, and is currently leading the Kurte´n, I became aware of biostrati- (Fig. 1a). Alternatively their possible Ancient Human Occupation of Britain graphic evidence that elements of the position as a ‘stem group’ for the project. Email: [email protected] Petralona mammalian faunas were Upper Pleistocene hominids could be of Cromerian age, potentially compa- recognized by the use of a subspe- 7 ?? rable to those from Mauer. Both cific name for the evolutionary grade Kurte´n and I considered the possibil- they are supposed to represent VC 2012 Wiley Periodicals, Inc. ity that the Petralona cranium could within H. sapiens (Fig. 1b). However, DOI 10.1002/evan.21311 Published online in Wiley Online Library represent a counterpart for the the other possibility that must (wileyonlinelibrary.com). Mauer mandible,6,8 even though the be considered (Fig. 1c) is that we 102 Stringer ISSUES

blance to the Elandsfontein calva- ria,11 and the possibility that Middle Pleistocene Chinese and Indian fos- sils might also belong in this group (Table 2) was raised.12,13 Rightmire has adopted a comparable Eurafri- can concept of heidelbergensis,14,15 while some have preferred to retain a more gradistic concept of Homo sapiens, arguing that fossils such as Broken Hill and Bodo are primitive examples of the modern human spe- cies.16 Other workers have used the informal term ante- or preneander- thal for earlier European fossils, including Mauer and Arago, some- times with a purely chronological meaning, and in other cases imply- ing an evolutionary relationship.17 This latter option has become increasingly popular with the recog- nition that the Sima de los Huesos (SH) material displays a mosaic of heidelbergensis and neanderthalensis features. For such workers, H. heidel- bergensis could represent an early stage in the accretion model of Ne- anderthal evolution,17,18,35 forming a heidelbergensis-neanderthalensis con- tinuum. I briefly considered this argument, going so far as to suggest that all heidelbergensis material might be lumped into Homo neander- 19 Figure 1. Illustration of the possible phylogenetic relationships of the Petralona and Broken thalensis, but I did not persist in Hill fossils. Redrawn, with permission, from Stringer.9 that view. However, if the European- only model of heidelbergensis is cannot at present resolve the exact heidelbergensis hypodigm. A separate correct, then the non-European com- phylogenetic position of these homi- study of the first Bilzingsleben cra- ponents assigned to heidelbergensis nids because they are close to the nial finds reinforced their resem- by workers such as Rightmire and point of divergence between Nean- derthals and ‘modern’ H. sapiens TABLE 1. Some Traits Observed in H. heidelbergensis Fossils (assuming that the Neanderthals are Endocranial volumes overlap those of H. erectus and H. sapiens/H. not directly ancestral to ‘modern’ neanderthalensis Torus often highly pneumatized laterally, and superiorly into frontal squama humans). At present I believe this to Vault shape parallel-sided in posterior view be quite likely, and that these fossils Strong and continuous supraorbital torus* are close to the morphotype expected Occipital strongly angled* in the common ancestor of Neander- Strong continuous occipital torus* thals and ‘modern’ H. sapiens. If this Wide interorbital breadth* is so, only further careful analysis Iliac pillar* will allow a decision about the clad- Elongated superior pubic ramus*# Femoral platymeria*# istic affinities, and thus the classifica- High arched temporal squama1# tion, of fossils such as Arago 21, Pet- Gracile tympanic1# ralona and Broken Hill.’’ Increased midfacial projection expressed through measures of midline nasal I began to develop a suite of traits prominence1# (Table 1; Fig. 2) for grouping Broken In large-faced specimens there may be lack of both canine fossa and Hill and Petralona.10 European and infraorbital retraction# African fossils such as Bilzingsleben, Reduced total facial prognathism1# Vertesszo¨ llo¨ s, Bodo 1, and Elands- *Found in Homo erectus; þ potential synapomorphies with H. sapiens; # potential fontein were added to an enlarged synapomorphies with H. neanderthalensis. ISSUES The Status of Homo heidelbergensis 103

Figure 2. Facial (A) and lateral (B) views of crania. Clockwise from top left: Homo erectus (replica, , ), heidelbergensis (Broken Hill, Zambia), sapiens (recent, ), and neanderthalensis (replica, , ). All pictures Ó The Natural Histroy Museum London. [Color figure can be viewed in the online issue, which is available at wileyonlinelibrary.com.] me, which show few or no Neander- at all. In effect, the morphology of ples of heidelbergensis, such as thal apomorphies, would require a certain specimens does not accord Thomas Quarry and Kapthurin, in separate species names. Homo rhode- with that of the specimen. Obvi- siensis 20 would have priority for that ously, the taxonomic identification of TABLE 2. Some Fossils That May grouping.9,21 skulls lacking a mandible is problem- Represent Homo heidelbergensis* atic, but even some give Western RECENT RESEARCH ON THE rise to uncertainty regarding their Mauer assignment to the H. heidelbergensis MAUER MANDIBLE, AND THE Petralona hypodigm. In our analysis, the Arago SPECIES HYPODIGM Mauer mandible stands alone in its Ve´rtesszo¨ llo¨ s Recently, welcome attention has morphology, which appears to be the ?Bilzingsleben ?Kocabas¸ been given to the Mauer mandible outcome of a unique constellation of characters.’’ This mirrored the view ?Boxgrove itself, including further dating ?Ceprano 22 23 of Hublin,21 who argued that the work. Schwartz and Tattersall ?Zuttiyeh primitive of the Mauer man- compared its morphology with that of other Pleistocene hominins, noting dible and the predominance of cra- Broken Hill 1; tibia E.691; E.719 its particular resemblance to the nial material in discussions about Bodo Arago 13 mandible. Via the Arago Middle Pleistocene hominins ren- Elandsfontein assemblage, they were able to extend dered it unsuitable as a holotype. Ndutu their heidelbergensis grouping to In the most extensive comparative Kapthurin other European, Chinese, and Afri- analysis to date, Mounier, Marchal, ?Hoedjies punt 24 ?Thomas Quarry can fossils, while aligning the SH and Condemi concluded that the ?Sale´ material with Neanderthals. How- Mauer mandible and H. heidelbergen- sis could be specifically distinguished ?Tighenif ever, in their view, the derived ?Berg Aukas from H. erectus, H. neanderthalensis, of both the Neanderthal Eastern Eurasia and Homo heidelbergensis and H. sapiens. They produced a di- ?Narmada exclude these groups from the ances- agnosis for the species hypodigm ?Dali try of Homo sapiens. based on mandibular fossils such ?Jinniushan Rak and colleagues41 noted the idi- Mauer, Arago, Montmaurin, and SH. ?Yunxian osyncratic features of the Mauer jaw Their inclusion of the Tighenif man- *In my view, the inclusion of fossils such as and argued that ‘‘it is not clear dibles extended the species to Africa, Steinheim and Montmaurin is still doubtful, whether lumping certain specimens but unfortunately they did not while Narmada, Dali and Jinniushan may together in this hypodigm is justified include other potential African exam- alternatively represent early ‘‘Denisovans.’’ 104 Stringer ISSUES their analyses. However, it is worth the Atapuerca team’s preference for ern humans and Neanderthals.... remembering that a fragment of assigning the material to heidelbergen- Here, our 95% credible intervals for ramus found at Elandsfontein, possi- sis. However, for some time I have the MRCA [most recent common bly associated with the ‘‘Saldanha’’ cal- preferred to regard the SH material ancestor] (315-538 ka) fall squarely varia, was noted as being remarkably as an archaic form of neanderthalen- within the proposed dates from similar to that of Mauer in its shape.25 sis,13 based on the presence of Nean- palaeoanthropology.... Our estimates As mentioned, researchers using derthal-like features such as an incipi- are also consistent with dates derived metric and multivariate techniques ent suprainiac fossa and midfacial from analysis of neutral morphologi- have noted metrical and shape simi- projection in the crania, dental and cal characters in both species, 182- larities between Petralona and Bro- mandibular traits, and numerous 592 ka (mean 373 ka).34’’ ken Hill.5,14,26 This has been con- postcranial characters. More recently, Recently, even stronger reasons firmed through the use of geometric Ihavealsochallengedtheolderage have emerged to place the SH mate- morphometrics.27,28 For example, estimates obtained for the SH sample rial within the Neanderthal clade Friess,28 excluding the Mauer mandi- from a dated .32 That chal- rather than within heidelbergensis. ble, commented that ‘‘the striking lenge was based on taphonomic Data from the large SH dental sam- affinities between the holotype of H. issues rather than the age determina- ple have long suggested Neanderthal rhodesiensis (Kabwe) and a Euro- tion itself, but dating work on the affinities.35 Most recently, Martino´n- pean fossil (Petralona), as demon- speleothem is continuing (J.-L. Torres and colleagues36 concluded: strated previously and confirmed Arsuaga, personal communication). ‘‘We find that SH dentitions present here, are unlikely to disappear, even Regardless of those considerations, all the morphological traits that, ei- if more complete specimens were to morphological and genetic data sug- ther in their degree of expression, be included in the analysis. This frequency, or particular combina- makes H. rhodesiensis an Afro-Euro- tion, are usually considered as typi- pean species that retains an erectus- cal of Homo neanderthalensis. This like plesiomorphic calvarial shape, Recently, even stronger study ratifies the deep roots of the but no /H. sapiens apo- Neanderthal lineage in the Middle morphies, unless one lumps it with reasons have emerged Pleistocene of . In addition, SH5 and Steinheim.’’ In a further to place the SH material SH teeth are morphologically ‘more study, Mounier, Condemni, and Neanderthal’ than other penecontem- Manzi29 added the Ceprano calvaria within the Neanderthal poraneous Middle Pleistocene sam- to their comparative analyses. They clade rather than within ples such as Mauer or Arago, and grouped the Ceprano specimen with heidelbergensis. even more derived than some classic Eurasian fossils such as Petralona, Neanderthal samples.’’ Because they SH5, Steinheim, Dali, and Jin- adhere to an age of >530 ka for the niushan, as well as with the Broken SH sample, these authors are forced Hill cranium. They concluded that in gest that the divergence of the nean- to add, ‘‘Thus, our study would not spite of its relatively young chrono- derthalensis and sapiens lineages very sustain the linearity of the accretion logical age (385-430 ka), the Ceprano likely postdates 530 ka. Therefore, process hypothesized for the origins fossil could represent a morphologi- Neanderthal apomorphies would not of the Neanderthals, and we suggest cally primitive example of the widely be expected at such an antiquity. that other evolutionary models and dispersed species heidelbergensis. Comparisons using the draft Nean- scenarios should be explored for the derthal produced the follow- Middle and Upper Pleistocene of ing estimate: ‘‘Assuming that human- Europe. We propose that more than ARE THE SIMA DE LOS HUESOS average DNA sequence one hominin lineage may have coex- FOSSILS PART OF divergence was 5.6 to 8.3 million isted during the Middle Pleistocene ago, this suggests that Nean- in Europe.’’ While I agree with that HEIDELBERGENSIS? dertals and present-day human pop- last statement, it is on completely Although Mounier and co- ulations separated between 270,000 different grounds. Martino´ n-Torres workers24,29 felt able to include the and 440,000 years ago.’’33 Using only and coworkers35 argue for this on SH material in their heidelbergensis complete mtDNA sequences, Endi- the basis of an (in my view errone- hypodigm, I believe that this inclu- cott, Ho, and Stringer32 argued that ous) early date for neanderthalensis sion has led to the most serious con- ‘‘our genetic date estimates are (SH) characteristics. I would instead fusion surrounding the status of the inconsistent with the late Early Pleis- argue for coexistence on the basis of species. From the first detailed tocene, early Middle Pleistocene, and late heidelbergensis age estimates descriptions of the Sima fossils, their late Middle Pleistocene models for within and outside of Europe.29,37–40 combination of heidelbergensis-like the divergence between H. sapiens In addition, it may well be that sig- and Neanderthal-like features has and H. neanderthalensis.... Rejection nificant Neanderthal-like samples been recognized.19 Increasingly old of these three models leaves just the such as , considered to be age estimates for the SH assemblage mid-Middle Pleistocene model for relatively late in the European (now >530 ka30,31) have reinforced the time of divergence between mod- sequence, are older than is currently ISSUES The Status of Homo heidelbergensis 105 believed (R. Gru¨ n and C. Stringer, that is ancestral to modern humans grounds, I believe that a pragmatic unpublished research). and Neanderthals, which, in my view, case can still be made for such spe- Rak and coworkers41 have recently is still most reasonably named Homo cies distinctions, especially bearing added to the data that the SH material heidelbergensis. However, new data on in mind the extensive evidence for is predominantly Neanderthal in its the possible eastern representatives of interspecific hybridization even in affinities: ‘‘The claimed similarities heidelbergensis have emerged from the living .49 between the characters of the Mauer genomic study of fragmentary fossils Genetic data can provide new per- specimen...and those of Neandertals at the southern Siberian site of Deni- spectives on the evolutionary history and the Sima de los Huesos mandi- sova. Initial mitochondrial DNA of heidelbergensis and its daughter spe- bles...cannot be considered homolo- (mtDNA) study of a large sug- cies, as can be illustrated by compar- gous, and hence, they are not synapo- gested an ancient lineage predating ing Figure 1c and Figure 3, based on morphies. Although some of the the divergence of Neanderthals and the mtDNA divergence data discussed Mauer characters superficially resem- modern humans, but genomic recon- earlier. Gene trees are not species ble the ones on the Neandertal and struction centered on a phalanx indi- trees, of course, but Figure 3 may Sima specimens, the Mauer characters cated that the ‘‘Denisovans’’ were nonetheless serve as a useful heuristic stem from a different morphological actually a subgroup of the Neander- device. First, it is evident that both the configuration. On the other , the thal clade.47 This finding has fueled modern human (A) and late Neander- similarities between the Neandertal speculation that fossils previously thal lineages (B) may have suffered characters and those of the Sima man- considered to be possible Asian repre- bottlenecking, perhaps during the dibles are the outcome of identical sentatives of heidelbergensis, such as harsh conditions of Marine Isotope configurations, making these charac- Dali, Jinniushan, and Narmada,13 Stage (MIS) 6; the slender evidence of ters true synapomorphies.’’ This was could in fact be Denisovans, but this mtDNA already suggests reinforced as follows: ‘‘The study of will remain uncertain until more com- greater diversity in that lineage.50 mandibles from Sima de los Huesos plete material yields DNA. More an- that would have been reveals an identical morphology to cient Asian specimens such as the represented within early or archaic that of the corresponding and unique Yunxian crania () might still members of the sapiens (C) and nean- region in the Homo neanderthalensis represent examples of heidelbergen- derthalensis groups (D) has conse- mandible.... This constellation of char- sis13,43 and potential ancestors of the quently been lost. The hypothetical acters is absent in other early and late Denisovans, although biogeographic mtDNA last common ancestor for hominins, including the type specimen and archeological arguments can be AþCandBþD is estimated at 338-538 Homo heidelbergensis (the Mauer man- made against such as assignment.44 ka (mean 407 ka), while population dible).... We conclude that the mandi- In addition, the presence of relatives divergence is placed at 315-506 ka bles of the Sima sample are virtually of the Neanderthals in the Far East (mean 345 ka). The predivergence seg- identical to the Neandertal mandible. forcefully reminds us how much our ment labeled F would have existed Thus, we regard this morphology as a views are biased by the attention paid duringthetimespanofHomo heidel- synapomorphy that the Sima fossils to the European and African records. bergensis. I argue that if we had fossils share only with H. neanderthalensis.In We cannot exclude an Asian origin for of the relevant ancestors, we would some other cranial morphologies, the heidelbergensis, given the similar ages recognize them as members of that Sima sample does not resemble Nean- (600 ka) assigned to the earliest (if species. While fossils such as Swan- dertals; hence, we place the Sima we exclude Tighenif) potential exam- scombe and SH might correspond specimens within the Neandertal clade ples in Germany (Mauer22), China with segments D or E, in my view we as a sister group to Neandertals. This (Yunxian45), and Ethiopia (Bodo46). have yet to identify specimens showing conclusion is inevitable regardless of The new Denisovan genomic data convincing apomorphies of Homo whether one advocates a cladogenetic are also consistent with previous evi- sapiens that could represent the ear- or anagenetic model.’’42 dence of gene flow between Homo liest stages of segment C, although fos- sapiens dispersing from Africa and sils such as those from Florisbad and native archaic populations (Neander- Guomde are possible candidates. 33 GENOMIC DATA AND THE STATUS thals). Limited but viable - As for what initiated the divergence ization events led to an input of ‘‘ar- of these lineages, one possibility is to OF HEIDELBERGENSIS chaic’’ genes into all non-African return to an idea suggested more than As I previously argued,32 reclassify- people in the case of Neanderthal 50 years ago by Howell,51,52 though ing the SH material as an early form DNA, and Australasian populations framed by him in the context of the of H. neanderthalensis on the basis of in the case of the Denisovans.47 Such last glaciations. There is evidence now its derived Neanderthal features and gene flow is bound to raise serious that glaciation in the Balkans was dating it to no earlier than 400 ka issues about the validity of specific much more severe during the Middle would remove most of the data sup- distinctions between heidelbergensis Pleistocene than at the Last Glacial porting a European chronospecies of and its daughter species, particularly Maximum.53 Also, pollen data from H. heidelbergensis-H. neanderthalensis. as such hybridization events may Tenaghi Philippon indicate that MIS This would open the possibility of a even have occurred in Africa.48,58 12 (450 ka) was particularly severe.54 less inclusive diagnosis for the species Nevertheless, on morphological If,atthattime,cold,aridconditions 106 Stringer ISSUES

derthal features claimed for the spe- cies, simultaneously differentiating it from contemporaneous African fossils and linking it to the succeeding Nean- derthals. In this review, I have discussed the growing and, in my view, convincing evidence that the Sima de los Huesos material belongs to the Neanderthal clade, and perhaps represents a primitive form of Homo neandertha- lensis. Removing the extensive SH assemblage from H. heidelbergensis greatly clarifies the situation in also removing most of the unique links to the Neanderthals. This allows a reformulated heidelbergensis to ap- proximate more closely the plesio- morphous morphotype expected for the last common ancestor of Homo neanderthalensis and Homo sapiens. However, if genetic and morphologi- cal estimates for neanderthalensis- sapiens divergence at <530 ka are Figure 3. Reconstruction of mtDNA evolution in the sapiens and neanderthalensis line- accurate, the SH material must be ages, based on complete from 5 Neanderthals and 54 modern humans. Based on data in Endicott, Ho, and Stringer.32 younger, and perhaps considerably younger, than this date. The addition of ‘‘Denisovans’’ to the extended eastward across the high contributes selective pressure to hominin lexicon provides a further relief of the Taurus-Zagros mountain adapt and diverge.’’ dimension to these discussions. The systems, coupled with enlarged Cas- relationship of non-erectus Asian Mid- pian and Black Seas, European popu- dlePleistocenefossilstothosefurther CONCLUDING REMARKS lations could have been effectively iso- west has long been problematic, but lated from their African and Asian Clearly, many problems in Middle now we have the potential to properly counterparts. Moreover, increased Pleistocene human evolution remain integrate the hominin records from aridity in and the Levant unresolved. Some of these center on western and eastern Eurasia for the could have added to this paleogeo- chronological issues, others on the first time, and to see East Asian fossils graphic separation. Whether increased lack of data for some important fossils, like Dali and Jinniushan as counter- selection or drift then operated to dif- such as those from China. The rela- parts of the evolving Neanderthals - ferentiate these separated populations tionship of , still defi- ther west. Indeed, it may be just as log- progressively is still uncertain,55 nitely identified only from Atapuerca, ical to regard the Neanderthals as a but Neanderthal-derived features are to succeeding samples is also unclear, western subset of the Denisovan group evident in Europe from MIS 11 although it remains possible that this as to consider, as is usually done, the onward.13,52,56 derivative of Homo erectus went inverse relationship. The concept of A comparable speciation scenario, extinct during the early Middle Pleisto- Homo heidelbergensis remains at the using the mechanisms of refugia, has cene.59 Homo antecessor also seems an center of such discussions, as this spe- recently been proposed by Stewart unlikely ancestor for Homo heidelber- cies represents the probable ultimate and Stringer57: ‘‘When a lineage gensis, which means that the origin of ancestor of these three daughter allo- adopts a new (or changes its) refu- Homo heidelbergensis is obscure. How- taxa: sapiens, neanderthalensis,and gial area, and it survives for a num- ever, in my view, the main uncertainty Denisovans. ber of Milankovitch cycles, expand- about Homo heidelbergensis is much ing from and contracting into that more fundamental, concerned with its ACKNOWLEDGMENTS new refugium instead of its original very nature. The idiosyncratic mor- refugium, it is destined to evolve into phology of the type specimen is cer- I thank numerous colleagues, a distinct population. Given enough tainly problematic, but for me an even including those quoted here, for dis- time in isolation, it will become a more vexing issue is whether the spe- cussions about Homo heidelbergensis, new species.... A new refugium is cies existed only in western Eurasia and four reviewers for their helpful unlikely to have the same flora, and gave rise solely to the Neander- comments. I also thank Laura Buck fauna, and ecology compared to the thals. The main support for such a for her work on the manuscript and lineage’s original refugium, which view has come from the derived Nean- figures. Figure 1 was redrawn with ISSUES The Status of Homo heidelbergensis 107

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