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Before the Emergence of Homo Sapiens: Overview on the Early-To-Middle Pleistocene Fossil Record by Giorgio Manzi (2011)

Before the Emergence of Homo Sapiens: Overview on the Early-To-Middle Pleistocene Fossil Record by Giorgio Manzi (2011)

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07/02/2012

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SAGE-Hindawi Access to ResearchInternational Journal of Evolutionary Biology Volume 2011, Article ID 582678,11pagesdoi:10.4061/2011/582678
Review Article
BeforetheEmergenceof 
Homosapien
:OverviewontheEarly-to-Middle PleistoceneFossil Record(withaProposalabout
Homoheidelbergensi
at thesubspecificlevel)
GiorgioManzi
Dipartimento di Biologia Ambientale, Universit`a di Roma “La Sapienza”, Piazzale Aldo Moro 5, 00185 Roma, Italy 
Correspondence should be addressed to Giorgio Manzi,giorgio.manzi@uniroma1.itReceived 15 September 2010; Revised 5 January 2011; Accepted 24 February 2011Academic Editor: Darren CurnoeCopyright © 2011 Giorgio Manzi. This is an open access article distributed under the Creative Commons Attribution License,which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.The origin of 
H. sapiens
has deep roots, which include two crucial nodes: (1) the emergence and di
ff 
usion of the last commonancestor of later
Homo
(in the Early Pleistocene) and (2) the tempo and mode of the appearance of distinct evolutionary lineages(in the Middle Pleistocene). The window between 1,000 and 500 thousand years before present appears of crucial importance,including the generation of a new and more encephalised kind of humanity, referred to by many authors as
H. heidelbergensis
.This species greatly diversified during the Middle Pleistocene up to the formation of new variants (i.e., incipient species) that,eventually, led to the allopatric speciation of 
H. neanderthalensis
and
H. sapiens
. The special case furnished by the calvarium foundnear Ceprano (Italy), dated to 430–385ka, o
ff 
ers the opportunity to investigate this matter from an original perspective. It isproposed to separate the hypodigm of a single, widespread, and polymorphic human taxon of the Middle Pleistocene into distinctsubspecies (i.e., incipient species). The ancestral one should be
H. heidelbergensis
, including specimens such as Ceprano and themandible from Mauer.
1.Introduction
The origin of anatomically and genetically modern humans(
H. sapiens
) from a small population of “archaic”
Homo
is anevent reasonably well set in sub–Saharan Africa around 200thousand years before present (or ka) [13]. Nevertheless, this event has deep roots in the Middle Pleistocene, primarily at the time of the divergence between the evolutionary lineage of our own species and that of the Neanderthals—between approximately 800 and or 520ka, according toBriggs and coworkers [4] (compare [5]), or between 538 and 315ka, according to Endicott and colleagues [6]—andeven earlier, in the late Early Pleistocene, when the commonancestor of both
H. sapiens
and
H. neanderthalensis
emergedand began to spread geographically.This paper aims at investigating such a new frontier forpaleoanthropology. It will focus on topology, chronology,tempo, and mode of the main evolutionary nodes before theappearance of 
H. sapiens
.When considering all the available data, we are con-fronted with a comprehensive scenario about the deep rootsof our species. At the same time, it becomes possible toapproach the issue from regional and/or local perspectives[7]. The special case study provided by a well-known fossilspecimen from Italy—that is, the calvarium from Ceprano(for a review, see [8])—may help to see the remote origins of 
H. sapiens
from an interesting and helpful perspective.
2.Old andNew Paradigms
 2.1. The Midcentury View.
Views about the origin of ourown species greatly changed during the last couple of decades, involving our interpretation of the evolution of the genus
Homo
as a whole. The paradigm of mid-20thcentury, shared among many scholars until recently, wasbased on the existence of a single human species that evolvedgradually and sequentially during the entire Pleistocene: itis the so-called “single-species hypothesis,influenced by 
 
2 International Journal of Evolutionary Biologthe gradualist perspective of the “modern synthesis[9,10]. Consistently with this perspective, the current humandiversity would have been the result of small and constantchangesamongpopulationsandwithinthespeciesasawhole(viewed as a single, extremely large population) taking placefrom the original geographical di
ff 
usion of the genus
Homo
,where the roots of the present human diversity would befound.According to this view, successive stages and/or regionalvariants were implied within a widespread archaic species—
H. erectus
—and subsequently among
H. sapiens
. The resultwas a model of either progressive or regional changes that,with the persistence of a single polymorphic humanity ineach span of geological time, was evolving towards variantsof the modern species and, ultimately, to the present human“races” [11]. As a consequence, the taxon
H. sapiens
includedthe extant humankind, but also extinct morphotypes likethe Neanderthals in Europe and the Near East as well as,more in the depth of the Middle Pleistocene, diverse archaichumans in Africa, and in eastern Asia, collectively referred toas “archaic
H. sapiens
.” Thus, under the same specific name,
H. sapiens
, each of these morphs was ascribed to a di
ff 
erentsubspecies, with the adoption of a trinomial nomenclaturethat gave rise to nomina such as
H. sapiens neanderthalensis
and
H. sapiens sapiens
.The theory called “multiregional evolution” is a goodexample of this view. It is well known that accordingto the model introduced by Wolpo
ff 
and colleagues [12]the anatomically modern populations of Africa, Asia, andEurope have been viewed as emerging in continuity withthe preceding archaic humans of the same geographicalarea. This hypothesis was based on the observation thata certain degree of “regional continuity” characterizes themorphologies of archaic and modern populations withineach geographical area. This observation, however, has beencontradicted by a number of works [13,14] and, more in general, the model is regarded as not in agreement with mostof the paleogenetic data (since the seminal work by Cannet al. [15]).Toalargeextent,thisviewhasnowbeenabandoned,aftera debate that lasted for more than two decades [16]. Someof the issues of this scientific revolution (
sensu
Kuhn [17])may be summarised with reference toFigure 1and are briefly discussed below.
 2.2. A New Scenario.
One of the consequences of the middle-century paradigm on the interpretation of the fossil recordwas that a single species,
H. erectus
, included a largeldistributed, archaic-looking, and polymorphic humankindthat was ancestral to modern humans. It was consideredto be the first hominid species that left Africa, equippedwith a brain of about 1,000ml and skilled enough toproduce the elaborated stone tools of the Acheulean, andable to face di
ff 
erent climatic and environmental conditionsbecause of its behavioural and technological—someone says“cultural”—aptitudes. In brief,
H. erectus
was viewed as thequasimodern “hero” who “conquered” the Old World: anauthentic “dawn of humanity.”However, an increasing number of data—including theevidence coming from the Georgian site of Dmanisi [18,19]—suggests a di
ff 
erent scenario. Those that were respon-sible for the first out-of-Africa di
ff 
usion were not derived,encephalised, and technologically advanced humans, butmore archaic hominins, with a brain just above the thresholdof 500–600ml (maximum encephalic volume of the aus-tralopithecines) and morphologically close to the changingdefinition and hypodigm of 
H. habilis
(after Leakey et al.[20]). Driven by ecological rather than behavioural or“cultural” motives, these hominins had a tendency—whichwas new with respect to
Australopithecus
,
Paranthropus
,and the Primates in general—to di
ff 
use and adapt tovariable, nontropical environments. The same corpus of datasuggests that this process should have started well before theappearance of either
H. erectus
or the Acheulean (which arenow viewed as separated and independent phenomena), thatis roughly between 2,000 and 1,500ka.Within this new approach,
H. erectus
may be viewed asa Far Eastern (Java and China) species only (
contra
Asfawand colleagues [21]) whereas their African counterparts areconsidered as a distinct species,
H. ergaster 
[22,23]. Alterna- tively, these two geographical variants are grouped under thedefinition
H. erectus
sensu lato, while
H. erectus
sensu strictowould be the Asian deme of such a multiregional taxon. Atthe same time, other species have been named or old nominahave been reconsidered, and they include (according to thechronological order of references that are pivotal for thepresent debate):
H. rudolfensis
[22],
H. heidelbergensis
[24],
H. antecessor 
[25],
H. helmei
[26],
H. mauritanicus
[27],
H.rhodesiensis
[28],
H. georgicus
[29],
H. cepranensis
[30], and
H. floresiensis
[31]. However, many of these taxa are debatedand/ornotwidelyacknowledged;thus,thegenerallyacceptedscenario is far less “speciose” than it might appear from sucha tentative list.Although the identification of all these di
ff 
erent speciesclearly implies an overestimation of interspecific diversity—that in many cases was more probably intraspecific (seebelow)—this plethora of nomina is in accordance witha scenario that foresees the geometry of an “adaptiveradiation,” describing the generation from a common stemof a great variability, in space as well as in time. Moreover,this confers a clearer, more intelligible significance to humanvarieties that were formerly hidden, being referred either to
H.erectus
(sensulato)ortotheinformalandconfusingentity called until recently “archaic
H. sapiens
” (which has beenabandoned and has almost disappeared from the literature).Looking at Europe, the relatively best known regionalexample, at least two distinct waves of immigrants seem tobe recognizable between the late Early and the early MiddlePleistocene. In terms of fossil record, the former wave isdocumented at present only in Spain and is referred to
H.antecessor 
. As a matter of fact, this species occurs in two sitesof the Sierra de Atapuerca, near Burgos: in the layer TE9of the Sima del Elefante, dated to about 1,200ka [32], andin the layer TD6 of the Gran Dolina, dated to more than780ka [25]. These humans are in association with the so-called “Oldowan” (or Mode 1 of the Lower Paleolithic [33]),largely di
ff 
used in a number of sites of Mediterranean and
 
International Journal of Evolutionary Biology 3
2.2 2 1.8 1.6 1.4 1.2 1 0.8 0.6 0.4 0.2 0Africa andNear eastEuropeAsia andAustralasiaMaDmanisi
H. heidelbergensisH. heidelbergensisH. heidelbergensisH. floresiensisH. erectusH. antecessor H. neanderthalensis
   H .  s  a   p   i  e  n  s
?“early 
Homo
H. ergaster 
Figure
1: Evolutionary tree of the genus
Homo
illustrating trajectories of the di
ff 
usion and/or tentative phylogenetic relationships betweena limited number of species. Note that, according to this scenario, only 
H. sapiens
and
H. heidelbergensis
(not
H. erectus
) have a widespreaddistribution in Africa and Eurasia. Also what is called here “early 
Homo
(
H. habilis
? Primitive
H. ergaster 
?) might have been largely distributed at the beginning of its dispersal (i.e., well before 1Ma). Legend. Continuous bold lines: time-span covered by species in di
ff 
erentregions; dashed lines: inferred extension of specific time limits, although the evidence may be missing, dubious, or controversial; dottedlines: phylogenetic links and/or trajectories of di
ff 
usion.
continental regions [34]. In addition, their settlements couldhave been discontinuous until about 600ka, being strongly influenced by ecological conditions [35].Seemingly, the latter possible dispersal into Europe wasmore recent than 700–600ka and related to morphologically derived hominids, with clear signs of further encephaliza-tion, which are well known from a number of sites. The mostnotable assemblage of fossil material is again in the Sierrade Atapuerca [36], namely, at the site with the evocativename of Sima de los Huesos, or SH, whose extraordinarily richandwell-preservedhumansamplehasbeenpushedbackto about 600ka [37]. These fossils are generally includedwithinthespecies
H. heidelbergensis
,whichisusually,butnotalways, associated with Mode 2 (or “Acheulean”) Paleolithicassemblages.
H. antecessor 
and
H. heidelbergensis
compete, one againstthe other, for the same phylogenetic position in currentevolutionary trees of the genus
Homo
, viewed as provisional,alternative models of human evolution.
H. antecessor 
isregarded by the Spanish workers (after [25]) as the stemspecies that was ancestral to the evolutionary divergencewithin the late natural history of the genus
Homo
, leading tothe evolution of the Neanderthals in Europe and to the emer-gence of our species in sub–Saharan Africa. Alternatively, itis
H. heidelbergensis
to be claimed for the same crucial role[24].Inasense,theSierradeAtapuercaprovidestheevidencethat could explain this ambivalence, either in the former orin the latter direction. At the same time, however, it shouldbe remarked that the material from Sima de los Huesosis clearly Neanderthal oriented, being characterized by anumber of features that, later in the Pleistocene, will becometypical of the Neanderthals [38,39]. Looking at the material from Atapuerca SH, therefore,
H. heidelbergensis
acquires theidentity of a European regional chronospecies in continuity with
H.neanderthalensis
,whichappearsinappropriateasalsoancestral to the African lineage leading to
H. sapiens
. Thereare, anyway, in Europe other fossil specimens—such as thecalvarium from Ceprano, Italy (see discussion below)—thatare penecontemporaneous with the material from AtapuercaSH and might represent (far better than the Spanish sample)a possible ancestral morphotype for
H. heidelbergensis
, if we look at this species as the evolutionary stem beforethe divergence between Neanderthals and modern humans(compare [6]).Moving to Africa, it has been shown [8,23] that specimens dated to about 1,000ka or slightly less—suchas Daka, Buia, and Olorgesaille—share phenetic a
nitiescloser to
H. ergaster 
than to Middle Pleistocene Africanhominins—like Bodo and Kabwe—which are referred to
H. heidelbergensis
or, alternatively, to
H. rhodesiensis
. Hence,these sub–Saharan specimens of the late Early Pleistocenesignal a morphological discontinuity with the subsequentfossil record and should be considered as late representativesof 
H. ergaster 
. In addition, as emphasized inFigure 1, the

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