A concise re-assessment of the spread of

               hominids into Eurasia from Africa circa 2 mya, taking

            into account the latest fossil evidence and molecular data

 

 

                                            R Walsh, G. Spiller, A. Harbin

                                      

                                                      Lewis Research Unit

                                  

Abstract: A revision of previously held views that hominids diverged from apes 5.0 -7.5 mya, thereby evolving in Africa before a migration into the rest of the old world circa 1.5 mya, needs to be made based on the latest fossil evidence, and molecular framework that has come to light at the beginning of this century. Evidence, both pathological and lithic assemblages, in areas of the world such as Dmanisi, Georgian Republic, and Mojokerto (Indonesia) dated to 1.8 mya, suggests a type of Homo sp. migrated from Africa to the old world earlier then previously believed. Indeed there may have been a type of convergent evolution happening in separate corners of the old world among hominoids that led to a much more complex scenario of hominid evolution, and migration patterns across the old world. In light of this newly proposed scenario for cross continental hominoid movement, comes the assertion that hominoids from Eurasia may have returned to Africa to become the ancestors of hominids. Recent fossil discoveries: Orrorin tugenensis in Kenya circa 6.0 mya, and Sahelanthropus tchadensis, Chad Republic circa 7.0 mya have pushed the back the dates of hominid origins and a new molecular framework has been devised which enables us to include these fossils in the hominid family. Therefore a revision of what is classed as a hominid is required to incorporate these new finds into the family tree, and a new progenitor for both Asian and European Homo erectus needs to be sought. One that left Africa prior to 1.5 mya.         

 

 

Gayle Spiller, B.A. (Hons), Head of Archaeology, Lewis Research Unit, Chinnor, Oxon, England     

Amy Harbin, B.A. (Hons), Sociology and Anthropology Dpt, Lewis Research Unit, London, England

Dr Robert Walsh, Director, Lewis Research Unit, Bournemouth, Dorset, England.

 

      

 Introduction

 As we take our initial footsteps into a new century, the science which is palaeoanthropology, is embracing an evolutionary, if not revolutionary change in itself, directly affecting those of us who are involved in this scientific discipline.   

Whether we accept all of, or part of the information that is being directed at us, it is important to occasionally take a pace back to remind ourselves what we already understand, and how each piece of new information received, can best be assimilated into our neural database of knowledge.

Some information can be ‘hard to swallow’, especially if ones entire perspective of human evolution, or a significant part of it, is challenged by the latest scientific thought or new discoveries (Andrews 1996, Arnason, 1996, Tobias 2003).

We only have to think back a few years, to the 1980’s and 1990’s, to the impact molecular biology had on palaeoanthropology (Cann, Stoneking and Wilson 1992)

For many years the multi-regional hypothesis of human evolution held sway (Thorne and Wolpoff 1992). A new breed of young scientists chose to challenge it with an ‘out of Africa’ (Srtinger and McKie 1996) (Stringer and Andrew 1988) slant to the evolution and dispersal of modern humans, Homo sapiens, replacing the Neanderthals in Europe, and other archaic hominids in south east Asia and China. (Tyler 1995)

Molecular biology in the form of mitochondria DNA testing has proven the ‘out of Africa’ theory of Homo sapiens movement circa 100 kya correct, (Cann, Stoneking and Wilson 1992) (Aitkin and Stringer 1993),  although a more complex story is now unfolding as to movements of an older ‘out of Africa’ migration by Homo ergaster (Rightmire 2001) (Tobias 2003)

It is precisely these earlier movements that this paper will examine in some detail, concentrating mainly on European hominids, and their origins. However it is prudent at this early stage to remind ourselves of the latest table of hominids phyletic evolution.  

           

                 Original African Dispersal with New Molecular Evidence

 

The old view that a direct linage, tracing hominid ancestry back to Pleistocene Africa, has been, in the main replaced with a more enlightened picture of several forms of hominids (Kenyanthropus platyops, A. afarensis, H.habilis/rudolfensis) contributing to what has been considered in most circles, the common progenitor of old world hominids post 1.5 -2.0 mya, namely Homo ergaster (Rightmire 1989, 2001) (Manzi et al; 2001).

 

                                   

The molecular evidence originating from Berkeley UCLA, has given human palaeontology a genetic framework into which the fossil assemblages could be arranged, in phyletic order (Cann, Stoneking and Wilson 1992). It also supplied a divergence date of hominids from our genetically nearest relatives, the Chimps at 5 mya to 7 mya.   

The newly published discoveries of the past few years, e.g. Ardipithecus ramidus kadabba (5.8-5.2 mya) from Awash River in Ethiopia (Wolde et al; 1994), Orrorin tugenensis (6.0 mya) Tugan Hills, Kenya, and Sahelanthropus tchadensis (7.0 mya) from the Sahel region of the southern Sahara Desert in Chad (Brunet et al; 2002), have given the scientists conforming to this molecular clock, something of a problem, as these older remains still betray, what are held to be hominid derived homologies, such as bipedalism (Lovejoy 1998).

This quandary has now been addressed by Ulfur Arnason and his team (1996) working at Lund in Sweden. They investigated the calibration points of the molecular clock, concluding they had to be re-interpreted in light of the latest fossil discoveries in palaeoanthropology. (see Arnason cited in Tobias 2003).

They used three mammalian calibration divergence points, which were 60 mya  = ruminant artiodactyls and cetaceans, 50 mya  = equidae and Rhinocerotidae, and 33 mya = Odontocetes and Mysticetes.

These calibration points were used to place the divergence between apes and hominids at 10.5 to 13 mya.

If this new molecular clock is adhered to, then the recent finds from Ethiopia and Kenya can be more confidently classed as hominids (Brunet et al; 2002).

 

 

  

                                 Movements of Hominids Around the Old World

 

The conception of this extended hominid linage, prises some interesting questions in exactly when hominids first left Africa (Bar-Yosef, 2001). In Fig 1., the old accepted chronology of hominid migration, we had H. ergaster at circa 1.5-2.0 mya as the initial migrant into the old world (Bosinski, 1995), (Larick & Ciochon, 1996).

If bipedal hominids were roaming south, east and central Africa, several million years earlier then H. ergaster, perhaps another hominid took the imitative to enter the old world, following either coastal routes, or river valleys (Bar-Yosuf, 2001) (Voris, 2000).

H. habilis (2.5 mya), would be an immediately suitable candidate, with

its slightly encephalized brain (Aiello and Wheeler, 1995), and ability to manufacture mode1stone tools (Potts, 1991), much published as being found in archaeological deposits outside Africa (Carbonne et al, 1999), e.g. Dmanisi (Brauer, 1996) (Lontcho, 2000), Longgupo (Larick & Ciochon, 1996), Riwat (Larick & Ciochon, 1996),Ceprano (Ascenzi et al; 1996, 2000)(Manzi et al; 2001), Atapuerca (TD-6) (Besco et al; 1995), La Vallonnet, Soleihac, and Isernia La Pineta (Sevink, 1982).

Once in Europe and Asia, the processes of evolution, would to an extent, customise a species with the plasticity of homo, within a suitable environment (Brauer and Schultz,1996) (Brauer and Mbua, 1992) ((Tyler 1995). The genetic distance between geographically isolated groups of hominids would have slightly grown, with a propensity for each homo sp. to develop along a similar evolutionary trajectory (Van der Made, 1992). The flow of genes between geographically separated hominids would have kept the gene pool from expanding too far between the groups (Thoene and Wolpoff, 1992). If this was the case, palaeoanthropologists will have to award new cognitive abilities and physical toughness to H. habilis. Cognitive abilities in the way of communication and scavenging techniques, that would have needed to be employed for longevity in a migration from Africa and protracted occupation of Europe and Asia (Huffman, 2001) (Sala,1996) . Physical toughness to sustain a relatively puny body, while still attaining a larger brain and smaller gut (Aiello and Wheeler, 1995), in order to process the high quality protein from meat, it was scavenging. 

The extended foraging periods necessary for survival, would have required an ‘Activity envelope’, or an expanded area to move around (Bar- Yosef, 2001) (C K Brain,1981), with relative safety from predation, all the time developing more muscle, social, sexual and reproductive conduct, and improving bipedalism. This scenario is also outlined in The Posture Hypothesis, (Hunt, 1996) and the Behaviour Model (Lovejoy, 1998)  To this end the palaeo-environments so far studied at Sangiran, Ngandong, and Sambungmachan in east Java, Indonesia (Santa Luca, 1980) (Barstra, 1982) (Barstra et al; 1988), Riwat in Pakistan, Lunggupo in China (Larick & Cichon, 1996), and Dmanisi in Georgia (Gabunia and Vekua, 1995), show environments well capable of sustaining groups of hominids in terms of camouflage, scavengable fauna, and importantly water, as bipedal hominids require on average 1 and a half litres of fluid a day, i.e. The Savannah hypothesis (Wheeler, 1991).

As water was a precious commodity to locate in the old world, and hunter/gathering/scavenging was at best a lucrative and precarious exploit, why did these early hominids, living outside Africa, remain bipedal, and not take to the forest canopy? (Susman, 1984).   

Questionable is the palaeo-environment of south east Asia, Java (Tyler,1995), and some of the Chinese provinces (Qian, 1985), as they may have been tropical, with a fair amount of forest cover. So why didn’t hominids evolve into arboreal primates to avoid predation?, and live on tree fruits etc? (Koop 1989). The answer to this is in the advantages of bipedalism itself, and the encephalized hominid brain. Fig 1 summarises these advantages.


 

 

 

Fig 1                          Advantages of Bipedalism on the Hominid Brain

 

 

                                                        BIPEDALISM

                     (originally due to changes in environment and topography in Africa)                          

 

REDUCTION IN HEAD                                   REDUCTION IN BODY AND

AND NECK MUSCLES                                    FACIAL HAIR

(development of Homo)                                      (development of Homo)

 

INCREASE IN CRANIUM SIZE                      REDUCED WATER INTAKE

AND EMISSARY VEIN NETWORK              AND BETTER QUALITY DIET

(increased cooling for larger brain)                    (more energy for prolonged foraging

                                                                             leading to faster absorbed proteins                   

                                                                             and essential amino acids)

 

                             ENCEPHALIZATION QUOTA INCREASED

                        ABILITY TO HANDLE SOCIAL COMPLEXITIES

 

Figure 1 displays in visual context how the evolutionary advantages of bipedalism and encephalization, married in Homo sp. to give longevity, adaptability, physical toughness, and a higher cognitive awareness to develop complex social skills, such as better linguistical communication or proto language. The advantages of remaining bipedal, and not reverting to tree dwelling primates are thereby borne out by the sustained link in mental and physical biology, granting geographically separated hominids the ability of complex social interaction and to maintain a physique conducive to a terrestrial hunter/gatherer/scavenger way of life. Fig 1 brings together the three hypothesis for the evolution and continuation of bipedalism mentioned earlier.

 


 

Once we’ve established the advantages of bipedalism in the oldworld (Foley, 1987), the two-way flow of hominids from Africa to Asia/Europe and back again to Africa (Rightmire, 2001), and flow of hominids between Asia and Europe (Clarke, 2000), seems much more plausible, taking into account environmental, geographical and topographical factors (Wood and Turner, 1995). The movement of hyper carnivores in Eurasia, e.g. Homotherium sp., Magantereon whitei, Pachycrocuta brevirostris, demonstrates that these factors (Arribas and Palmqvist, 2000), and barriers could be transversed. Homo sp. must also be classed as a ‘cleaver carnivore’, who could scavenge as well as hunt and gather. This made Homo a formidable predator in the old world, even at the stage of development of H. habilis and H.ergaster.   

Continuing this hominid distribution hypothesis, by the time H. erectus in Europe emerged at Ceprano, 800-900 kyr (Manzi et al; 2001) and TD-6 Atapuerca (Aguirre and Carbonell, 2001) <780 kyr, Homo sp. had evolved into an experienced traveller of distance, carrying many genetic markers of its peripatetic ancestry (Cann, Stoneking, and Wilson, 1992).

Fig. 2 and 3 are a guide to the current knowledge, based on fossil and lithic assemblages, of geographically separated early hominids in Africa and Eurasia.


 

 

 

 

 

                          Hypothetical Results of Genetically Related Homo ergaster/erectus   

      

Fig 2.       Average linkage:

 

_____________________________

African                          Asian

ergaster                         erectus                      ______________________________________________

                                                                        European              Homo                     Homo          Homo 

                                                                        erectus                  heidelbergensis      neander.       sapiens

_________________________________________________________________________________

2mya                              1.8mya                      800 kya                 500kya                   200kya        140kya

                                          

                                           

Fig 3.        Neighbour joining:

 

                                           *

                                                Asian erectus (Dmanisi/Sangiran/Riwat/Longgupo)

*

African ergaster

                                                                            *

                                                                               European erectus

                                                                                                   *

                                                                                           European heidelbergensis

                                                                                                                   *

                                                                                                  Homo neanderthalensis

_____________________________________________________________________2mya                                                  1.8mya                       800kya           500kya         200kya

Figures 2/3 generalise on an African origin for the first hominids to migrate into Eurasia. The following hominid species to evolve, Asian and European H.erectus, H.heidelbergensis, and H.neanderthalensis follow the generally held view of hominid descent in Eurasia.


 

 

The phylogenetic position of the Ceprano calvaria (Manzi et al; 2001) (Clarke 2000), is unique in that the calvaria displays both ancient H. erectus and later H. heidelbergensis / rhodesiensis features, thereby distinguishing itself as a pivotal bridge.   Is this hominid the descendant of one of the Indonesian populations migrating back into Europe, or are the hominids at TD-6, Gran Dolina, Atapuerca in Spain, descended from

the Dmanisi population. Such theories no longer seem so implausible. Many hypothesis can now be envisaged for the development and peopling of Eurasia by H. erectus. (Rightmire,

2001)  Recent studies hypothesise that   the ancestors of the Dmanisi hominids

 

(1.8 mya) were ancestral to the S.E. Asian lines of Pithcanthropus (1.8 mya), Longuppo, (2 mya?) and Sinanthropus (1 mya- 400 kya), all evolving separately, which explains the slightly different morphological anomalies, while the European H. erectus/H. antecessor came in a later migration circa 1.5 mya from Africa, either migrating through the middle eastern corridor, or by a direct route from North Africa via the straits of Gibraltar/Messina.  Morphological comparison and detailed cranial analysis can show similarities in many respects between European, Asian and African H. erectus (Ascenzi et al 1996, 2000), (Manzi et al 2001), (Brauer,1996), (Rightmire, 2001).


                                          The Role of Incipient Hominoids

 

In addition to this much more complex pattern of movement and cross movement of hominids circa 2.0 mya

is a theory proposed by Peter Andrews from London’s Natural History

Museum (Andrews,1996). He expands on this movement across the old world,

by suggesting old world Miocene apes (so called Incipient Hominoids), e.g. Ouranopithecus macedoniensis/Graecopithecus freybergi from Macedonia, and Samburupithecus kiptalami from Nachola and the Samburu Hills,  northern Kenya, are the progenitors of the homo lineage, after the divergence from the Pan lineage had occurred (Andrews, 1982), (Brain,1981).

The fossil record shows that circa 16 mya, Pliopithecus and Dryopithecus, two genera of primates left Africa and journeyed into Eurasia. Studies of dentition, show both were adapted as fruit eating vegetarians, and may have played an important role in the ancestry of Andrews Incipient Hominoids (Pickford, 1968).

A change of diet had occurred by the time of Ouranopithecus macedoniensis, as the surface of this species molars are heavy pitted, betraying a consumption of hard foods such as nuts and tubers (J Jacobs, 2000). 

As the fossil record in Africa between 10 mya and 7-6 mya is extremely sparse, Andrews believes that these Incipient Hominoids’ migrated back into Africa, their descendants thereby were Sahelanthropus tchdensis, Orrorin and Ardipithecus.

The morphological analysis for this hypothesis relies mainly on hemi-maxilla comparisons. Presumably then, the older lineages existing in Africa, relating to Proconsul, e.g. Proconsul africanus (18 mya from Rusinga Island, Kenya), and Kenyapithecus wickeri (14.5-12 mya from Fort Ternan (Solounias,1997), Kenya), may have led to other forms of old world monkeys, e.g. Gibbons. The Sivapithecines e.g Sivapithecus sivalensis (8 mya from Potwar Plateau, Pakistan) which bare such uncanny facial morphology to the modern orang-utans of Java and Sumatra (Pilbeam1984), presumably went extinct, or were already evolving into Orang-utans.

If Andrews is correct, then the migration by H. habilis or H. ergaster into Eurasia, completed a hominid lineage migratory full circle, with the decedents of the Incipient Hominoids returning to their ancestors former lands (namely Eurasia), with the palaeo-ecological factors having changed somewhat in the intervening time (Andrew 1996).

 

A hypothetical lineage to illustrate the previous information is herewith.

Fig 4               Revised Hominid Linage Incorporating Andrew’s Incipient Hominoids                           

 

 

Ouranopithecus macedoniensis------

Graecopithecus freybergi-------------      

Samburupithecus kiptalami----------                                                          

                                            Sahelanthropus tchaensis

                                                  Orrion tugenensis

                                                       Ardipithecus ramidus kadibba

                                                                                     Ardipithecus ramidus ramidus

                                                                                        Australopithecus/kenyanthropus platyops

                                                                                               Homo habilis/rudolfensis/ergaster

 

 

____________________________________________________________________________________________

9.0 -8.0mya (fauna dating)           7.0-6.0mya              5.8-5.2mya         4.4mya            4.2-1mya         

 

The inclusion of Incipient Hominoids stretches the beginnings of the hominid lineage back to around 9.0mya, and the question of where the later African hominids ancestors originated is answered. If proconsul led to gibbons/monkeys, and sivapithecus to orang-utans, then the remaining afropithecines/dryopithecines either went extinct or evolved into these Incipient hominoids whom migrated back into Africa to give birth to all hominids leading to Homo sp.     

 

      

 

                                 So Where Did The European Hominids Evolve ?

 

The question we started with, still requires further research and fossil evidence to show beyond reasonable doubt, where the hominid ancestors of Dmanisi ,Ceprano, Guadix-Baza basin, and Atapuerca originated. The incessant nature of hominoids, such as Andrew’s Incipient Hominoids to move around, crossing continents (Andrews 1982, 1985), together with Arnason’s new molecular divergence time scale for hominids and other apes (Arnason 1996), provides the palaeoanthropological world with a fresh view on our own evolution.  If H. ergaster, was not the first hominid to take the initiative for mass across old world colonization (Rightmire, 2001) 2.0 -1.5 mya, as previously believed, then an earlier form of hominid must have made that unique step. If the dates for the assemblages at Dmanisi (Lontcho, 2000), and Mojokerto (Curtis et al; 2000) are correct at 1.8 mya, then their ancestor, if it was African derived as we believe, must have been this earlier form of hominid, e.g. H.habilis/H. ergaster. The morphological analysis bears this out (Curtis et al; 2000).

   

Some of the calvarias’ morphological aspects in profile compare well with H. erectus, e.g. low cranial vault, a capacious and continuous supraorbital torus, angulated occipital vault presenting distinctions between inion and endinion areas. Only by closer analysis can accurate measurements be obtained. Fig 5 gives more on these measurable comparisons.

 

 

Fig 5                    Table Comparing Selected Calvaria’s Frontal & Vault Dimensions

 

 

 

 

Supraorbital vertical thickness

Sangiran 2

 

 

(L) 12mm

 

Sangiran17

 

 

(L) 18mm

OH9

 

 

(L) 18mm

KNM –ER3733

 

(L) 8.5mm

 

 

Ceprano

 

 

(L) 21mm

Frontal breadth index

80.4mm

83.2mm

81.3mm    

82.7mm

89.8mm

Maximum frontal breadth

102mm

119mm

119mm

110mm

118mm

Minimum frontal breadth

82mm

99mm

100mm

91mm

112mm

Biauricular breadth

126mm

140mm

135mm

132mm

128mm

Supramastoid breadth

141mm

161mm

146mm

142mm

161mm

Mastoid height

(R)12mm

(R) 25mm

       ?  

        ?

(R) 27mm

Lamba-asterion chord

(L) 82mm 

(L) 87mm

       ?

(L) 78mm

(L) 92mm

Lamba-asterion arc

(L) 92mm

(L) 91mm

       ?

(L) 79mm

(L) 103mm

Parietal saggital chord

98mm

106mm

       ?

79mm

95mm

Parietal saggital arc

103mm

109mm

       ?

83mm

      ?

Biparietal maximum breadth

137mm

144mm

123mm

127mm

125mm

Cranial capacity

815ccm3

1000ccm3

850cc3

850ccm3

1057ccm3

 

Figure 5 compares frontal and vault measurements of one H. ergaster (KNM-ER 3733) and four H. erectus calvarias. A few measurements have not been recorded (OH9) and Ceprano, but the overall picture emerging is of H.erectus inheriting cranial morphological homologies, which are present to this species across geological time and space. The Asian H erectus have a few autapomorphic features not present in African H. ergaster (sagittal keel, and tympanomastoid fissure) which are probably due to  geological distance, and varying environmental conditions affecting post cranial development to a certain degree (this may be the case with the Zhoukoudian specimens which display a unique metric pattern in south east Asia, e.g. wide midvault and narrow frontal and occipital bones, compared with African and Indonesian specimens that have broad frontal, midvault and occipital dimensions). This can not detract from the fact H. erectus was a peripatetic species and the view that Asian H. erectus was quite capable of travelling in to Europe and Africa (OH9) is sufficiently demonstrated in Fig 7 and Fig 8 as a possibility, if not a certainty. Alternative hypothesis could be that the H. ergaster like Dmanisi specimens represent a migration out of Africa heading towards Asia, and H. Antecessor was a direct migrant from North Africa. Cranial morphology dictates the Ceprano calvaria (which may be an adult version of H. antecessor ), holds a unique place in bridging the gap between African H. ergaster and both African and European H. heidelbergensis. It displays features both H. erectus derived, and ancestral to H. heildelbergensis/ rhodesiensis.    

 

      

                         

Conclusion

 

In the light of the new fossil and molecular evidence, the path of migration, geographic distribution, and re-tracing ancestors footsteps into continents they previously inhabited (in the case of Africa), seems not only plausible, but quite possible. This is a conjecture that will perplex much of the established palaeoanthropological community who hold onto the steadfast  opinion that only a single track migration from Africa into the old world, with the migrated hominids remaining strictly sedentary, and confined within the geographical boundaries of their continent, was what must have happened.

So what species originally left Africa to start this confusing criss - cross of movement?

Who were the earliest hominid migrants?

Sites such as Barranco Leon 5, Venta Micena and Fuente Nueva 3, in the Guadix-Baza basin site, southern Spain (1.5-1.0 mya) (Palmqvist and Arribas 2001), TD-6 at Gran Dolina, northern Spain (0.8 mya) (Carbonell et al; 1997), Ceprano (0.8 mya) (Ascenzi et al; 1996, 2000),  Monte Poggiolo, and Isernia La Pineta in Italy (0.8 mya) (Sevink, 1982), Ubeidiya and Erq-el Ahmar in Israel (2.0 - 1.4 mya) (Bar-Yosuf, 1994), Dmanisi in Georgia (1.8 mya) (Brauer G, 1996), Riwat in Pakistan  (?2.0 mya), Sangiran, Mojokerto (Huffman, 2001) (Hyodo et al; 1993) and Flores (Van den Bergh, 1996) (Moorwood et al; 1999), in Indonesia (1.8 mya – 0.8 mya), and finally Lantain, Yuanmou (Hu, 1973) and Xihoudi in China (Qian, 1985), (?1.8 -1.0 mya) are just some of the best known examples of early hominid occupation outside the confines of Africa.

It seems more likely that a form of     H. habilis was the hominid to leave Africa and enter Asia (via Dmanisi)and then Europe. Which Asian hominid community crossed into Europe from Asia, if one did, we can only speculate on. Perhaps Dmanisi to Ceprano and Atapuerca, or did Asian H. erectus migrate back to Africa, and then took a sea crossing from north Africa to Calabria, (Ascenzi et al; 1996, 2000) transversing the straits of Messina, and to Iberia via the straits of Gibraltar (Carbonell et al;1996). Maybe populations in North Africa circa 1.4 mya, took the initiative for a direct crossing into Europe.

In Asia, H. erectus may have itself, become extinct circa 74 kya due to catastrophic volcanic events, or indeed survived to evolve into the Asian equivalent of the Neanderthals, the so named Homo Soloensis, as represented by the younger skulls at Ngawi, Sambungmachan and Ngandong (Widianto, Zeitoun, 2003). 

 

Not enough fossil evidence exists at this present time to make conclusive judgements, but perhaps as we continue to move into this new millennia, more archaeological evidence will be excavated to give a better idea of these post 2.0 mya African migration routes of early hominids. Finally Fig 6-8 gives a few possible lines of decent of hominids in Europe and Asia taking into account all that has been discussed in this paper.


Fig 6      


 

New phylogeny of Homo erectus (based on the evidence discussed)

 

(Cladistic analysis results on data obtained by use of multivarient programme: NUTS..from 52 cranial measurements. Originally documented in ‘Ceprano Calvarium’ research document by R M Walsh 2001. Revised 2003).  

 

 

                                 -------------KNMER3733

                                 ------------- KNMER3883         -----------African archaic/heid.  -----------Anatomically

                                 -------------KNMER1813                    e.g. Sadhana,Florisbad, Kabwe      modern sapiens

                                 ------------KNMER WT15000     

                        

                                ------------RIWAT              ------------------        ?

 

                               ------------YUANMOU

                               ------------UBEIDIYA         ------------------         ?

                               -------------G.Y.B.

 

                               -------------DMANISI          ------------------         Orce, Ceprano, TD6 (H. antecessor)

H.hablis/-----------


 

ruldofensis

 


 

         

                        ---------SANG 2, 12, 17                  Yunnan 1,2              Sinanthropus 10,11,12     Dali

                               ----------MODJOKERT 2    -------- Lantain 1,2  ---------Yunxian 2      -------------- Jinnu-Shah-?

                               ----------TRINAL 1,2                       Xihoudi                          

                                                                 

                                                                  Ngandong 3,7,10

                                                                  Samb 1             ---------Ngawi 1 (H. soloensis)

                                                                                        Flores                          Orce, Ceprano, TD6 (H. antecessor) ?

                                                                                        OH9 ?                                   

                                              

 

 

 

 

 

 

 

 

 

Fig 7     Possible descent of Homo ergaster/erectus (based on evidence discussed)

 

                                                    H.habilis/rudolfensis/ergaster

                                                        (Africa, 2.4 – 2.0 mya )

_____________________________________________________________________                    

                                                                                                                          

Homo sp.                H. erectus                    H.erectus                     H. ergaster                    H. erectus

(Riwat, 2.0 mya ?)    (Sang/ Moj, 1.8 mya)     Lantian/Xihoudi/      (Dmanisi,1.8 mya)   (Erq-el-Ahmar/                  

(Pakistan)              (Indonesia) Trinal (900 kya)  Yuanmou,             (Rep. Of Georgia)    Ubeidiya, 2.1- 

                                                                      (1.8-1.0 mya, China)                                 1.4 mya,                  

      ?                  Homo sp           Homo ant/erectus                          Homo ant/erectus       Israel)

                             Flores           Orce/Atap.TD-6/              ?             Orce/AtapTD-6/              

                           (Indonesia)     Ceprano (Spain/Italy)                      Ceprano (Spain/Italy)     ?            

                                0.8 mya     0.8 mya                                                       0.8 mya               

                                   ,                    ,                                                                ,                                                  

  Homo soloensis                Homo heidelbergensis                                Homo heidelbergensis

 (Ngawi/Sam/Ngd)       (Sima de los Huesos, Spain                         (Sima de los Huesos, Spain

          0.4 mya                     Boxgrove, England                                     Boxgrove, England

                                             Petralona, Greece                                        Petralona, Greece

                                             Arago, France………..etc)                          Arago, France…….….etc)   

                                             

                                                                                                                    

                              Homo neanderthalensis                                      Homo neanderthalensis

                             (Middle East and Europe)                                   (Middle East and Europe)

 

 

 

Fig 8

          Cladistic anaylsis for a European phylogeny (based on evidence discussed)

 

 

Dmanisi series                         

 

 

OH9 (African series)                                Ceprano

                                       

                                                     ---------                  ------ H. heidelbrgensis

                                                                                            (Sima, Boxgrove,----------H. neanderthalensis

                                                                     Orce/TD6        Petralona, Arago)           (Middle east and

                                                                                                                                      Europe)

Ngandong/Sambun. (Asian series 1)

 

Trinil/Sangiran (Asian series 2)

 

 

The Neanderthals are the terminus in Europe and the Middle east, of this early out of African lineage, being replaced themselves by anatomically modern Homo sapiens migrating in a subsequent African migration. The possible interplay of hominids from Africa, Indonesia and Georgia to produce the European hominids represented by Ceprano, Orce, and Gran Dolina, Atapuerca is far too complex for a definite solution to be made apparent. Only a plethora of hypothesis of this interplay and the descendents that may have been left behind can be offered. Figures 6-9 therefore represents three possible theories. More probable on current evidence that the Dmanisi hominids were an early 2ma migration of hominid from Africa, on route to South East Asia, and the European H. antecessor the result of a later direct crossing from North Africa. 

          

 

 

 

 

 

          Acknowledgments

 


 

We are grateful to Mr R. Kruszynski, curator of Anthropology at London’s Natural History Museum, for his allowing R.Walsh and G. Spiller access to castes of many of the calvarias mentioned in this paper, for them to study, photograph, and measure. Also for his advice on the first draft of this paper.


 

 

 

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