Thomas Hopkins introduction

ATY 555, Spring 2010 D2700 Homo erectus/Homo georgicus

May 6, 2010

Specimen D2700 is a cranium found in Dmanisi, Georgia in 2001 and appears to be intermediate between Homo habilis, or the Habilines more generally, and Homo erectus. discovery and geological context Georgian scientist David Lordkipanidze is the chief name among several in an international team that discovered the D2700 specimen in the year 2001. As the specimen name suggests, D2700 comes from the Dmanisi site in the eastern portion of the Republic of Georgia, known for dated remains that suggest a quick H. erectus or H. erectus-like migration out of Africa; dates for hominin remains at the site hover around 1.75 to 1.8 million years ago (mya). The Dmanisi site itself is located approximately 85 kilometers southwest of Tiblisi (coordinates 41 19 N; 44 21 E). Excavations there began in 1936 but, at the time, centered around the medieval ruins and artifacts located at the site. It was not until the mid-1980s that the site gained interest within paleoanthropology, after stone tools were discovered there in 1984. The first finds from the team that subsequently assembled to excavate further came in 1991, when David Lordkipanidze unearthed a well-preserved hominin mandible, specimen D211, that established the early African exodus of Homo erectus. Geologically, the site is composed of an 80-meter thick layer of basalt that is covered by a 4-meter thick layer of alluvial deposits. By potassium-argon dating, this basalt has been dated at 1.8 0.1 million years old (Gabunia and Vekua 1995). D2700 was found in square 60/65 of the site and is associated with a mandible, specimen D2735, found nearby in square 60/66. Both of these specimens were situated in black to dark-brown tuffaceous sand immediately overlying the 1.85-million-year-old Masavera Basalt, (Vekua et al. 2002) and, therefore, its discoverers date it at around 1.75 million years old with the following justification: The new hominid remains were associated with animal fossils that include an entire skull of Stephanorhinus etruscus etruscus, a skull of Cervus perrieri with a full rack of antlers, a Dama nesti antler, two crania of Canis etruscus, a complete mandible of Equus stenonis, and the anterior portion of a Megantereon cranium. Human occupation at Dmanisi is correlated to the terminal part of the (magnetically normal) Olduvai Subchron and immediately overlying (magnetically reversed) horizons of the Matuyama Chron, and is ~1.75 million years in age. Faunal remains also support the dating of Dmanisi to the end of the Pliocene or earliest Pleistocene (Vekua et al. 2002:85; see also Messager et al. 2010). Thus, the date for D2700 is derived from paleomagnetic correlation, associated faunal assemblages, and potassium-argon dating. These require further explication. Paleomagnetic dating relies on the fact that Earths centers of magnetic polarity shift at a more-or-less known rate and that sedimentary rock aligns with the planets polar orientation at time of deposition. Obtaining dates via this method necessarily requires the principle of superposition, which states that geologic strata, when flatly laid, increase in age with increasing depth. Paleomagnetic dating utilizes this principle to establish a local sequence of magnetic reversals and, thus, can yield absolute dates. Faunal assemblages at a site can be correlated with assemblages at other

Hopkins 2 sites where absolute dates are known by other methods and can thereby yield effective date ranges. Finally, potassium-argon dating, an absolute radiometric method, assesses the ratio of potassium-40 to argon-40 within volcanic rock and correlates this with the half-life of potassium-40, which naturally decays to argon-40. A date of ~1.75mya puts D2700 on the cusp of the Pleistocene epoch, a time of climatic contrasts: periods of glaciation and glacial retreat. When assessing the Dmanisi environment at the time, Gabunia et al. (2000a:13) establish that [local] fauna reflect quite diverse landscapesan interpretation supported by paleobotanical evidence. The data reflect the prevalence of a moderately dry climate and the presence of fairly extensive open landscapes. A paper by Messager et al. (2010; in press at time of writing) clarify this statement with additional information from phytolith, pollen grain, and carpo-remain (i.e., fruit) analysis. They establish two phases of paleoenvironmental conditions at Dmanisi. The first of these phases, extending from ~1.85 to 1.77mya, was marked by both chestnut and riparian forests as well as grasses that require high moisture levels, which indicate that the Dmanisi environment during this period was relatively warm and humid. The second phase, after 1.77mya and called Matuyama after the type of basalt in which associated materials were found, was markedly drier, as hygrophilous foliage disappear entirely around this time, leaving only temperate grasses and oak forest, which require less water than, for example, beech forest (Messager et al. 2010; see figure 8 in particular). D2700 obviously falls within this second period of greater dryness. description The D2700 specimen is a cranium in fairly excellent condition. It shows signs of damage on its maxillae, zygomatic arches, orbital walls, and the interorbital region, as well as heavy erosion on both its mastoid processes (Rightmire et al. 2006). D2700 also exhibits at least one pathology; there is great resorptive bone loss at its molar roots, a sign of periodontal disease. (The associated mandible, D2735, shows less severe signs of the same pathology.) The specimen has a molar eruption pattern that indicates the individual had not reached adulthood at time of death. Its third molar is only partially erupted, and the occlusal surfaces are approximately level with the bases of the M2 crowns (ibid.:10, Vekua et al. 2002). This makes the D2700 individual ontogenetically older than the famous H. erectus skeleton, Nariokotome Boy (KNM-WT15000), which, by contrast, shows no signs third molar eruption. Furthermore, the synchrondrosis, i.e. cartilaginous joint, between the sphenoid and occipital bone has not fused, further underscoring D2700s juvenile status (Vekua et al. 2002). D2700 has a cranial capacity of approximately 600 cubic centimeters (cc). Per Philip Tobias (1991) correction of a similarly-aged habiline individual (OH 13), D2700 may have attained approximately 98% of its adult cranial capacity at time of death; its adult size would be approximately 612cc in this instance. Others suggest that the H. erectus pattern of brain growth was relatively ape-like, in which case the specimen would have already attained its adult brain size (Coqueugniot et al. 2004). Nevertheless, 600 or 612cc are relatively small sizes for an individual initially classified within the species H. erectus. Cranial capacity is one feature that presents issues for taxonomic placement of this specimen.

Hopkins 3 For the purposes of prefacing a discussion of taxonomy and situating D2700 within the hominin lineage without interpreting its features through the lens of a specific species designation, I will compare it with three other specimens whose taxonomic status I will consider irrelevant; consensus opinion is sufficient at this stage of my review for those fossils. I compare D2700 with specimens KNM-ER 1813, attributed to H. habilis, KNM-WT 15000, unequivocally considered to be a H. erectus skeleton, and KNM-ER 3733, another H. erectus cranium. The following table summarizes relevant anatomical characters in these specimens.
D2700 Geologic date ~1.75myo Brain size ~600cc Age at death Sub-adult; M3 partially erupted Projected adult brain size Cranial length Max. cranial breadth Postorbital constriction index Supraorbital torus thickness Occipital scale index Palatal Index (breadth/ length) Max. biparietal breadth Presence of a sagittal keel ~612cc 153mm 125mm 73.3 9mm 81.8 67.2 115mm Yes, but unique/ not fully erectuslike KNM-ER 1813 ~1.9myo ~510cc Adult; M3 erupted KNM-WT 15000 ~1.6myo ~810cc Sub-adult; M3 not erupted ~909cc ~175mm [not available] 76 [not available] 131.5 80.0 [not available] No KNM-ER 3733 ~1.8myo ~850cc Adult; M3 erupted, fused sutures N/A 182mm 142mm 72.7 8.5mm 92.9 69.3 131mm No

N/A 145mm 113mm 71.4 9mm 72.7 62.9 100mm No

On the whole, D2700 exhibits several features that are traditionally associated with H. erectus. These include the presence of supraorbital tori (thin but well-defined per Vekua et al. 2002:87), a slight sagittal keel, and a low and transversely flattened (Rightmire et al. 2006:125) appearance from the rear. Nevertheless, D2700 looks remarkably similar to ER-1813, a presumably more primitive specimen attributed to H. habilis or the habiline clade. D2700 and ER 1813 are similar in size, share similar supraorbital tori, have diminutive faces relative to other Homo specimens, and share similar palate dimensions. As the table above illustrates, the definite H. erectus specimens, WT 15000 and ER 3733, clearly have endocranial volumes above both D2700 and ER 1813; they are also, as figures for cranial length indicate, larger overall as well. D2700s pattern of keeling is unique at least for one reason; it is more prominent on the parietals relative to other Dmanisi specimens and indeed other H. erectus specimens. taxonomy and phylogeny Vekua et al. note the similarities that D2700 shares with ER 1813 but ultimately assign it to H. erectus:

Hopkins 4 A comparison of the new skull to other specimens from Dmanisi, Koobi Fora, and West Turkana suggests that it has a number of similarities to early H. erectus (or H. ergaster). The cranium is exceptionally small, with a rounded occiput, and its face is like that of KNM-ER 1813, especially in profile. The canine juga of D2700, however, are well defined, and the zygomatic root is very thick. Despite certain differences among these Dmanisi individuals, we do not see sufficient grounds for assigning them to more than one hominid taxon. We view the new specimen as a member of the same population as the other fossils, and we here assign the new skull provisionally to Homo erectus (=ergaster) (Vekua et al. 2002:88). In the same paper Vekua et al. make another revealing point: The Dmanisi hominids are among the most primitive individuals so far attributed to H. erectus or to any species that is indisputably Homo, and it can be argued that this population is closely related to Homo habilis as known from Olduvai Gorge in Tanzania, Koobi Fora in northern Kenya, and possibly Hadar in Ethiopia. It now seems more likely that the first humans to disperse from the African homeland were similar in grade to H. habilis (ibid.). In assigning D2700, associated mandible D2735, and, indeed, the Dmanisi hominins as a whole to erectus, Vekua et al. prioritized the well-defined canine juga, presence of a glenoid cavity, tympanic plate, petrous bone, noticeable though unique sagittal keel, and overall shape1 that the specimens exhibit. The literature on D2700 is somewhat inconsistent about its classification. The 2002 paper by Vekua et al., which seems to be the first in which D2700 is described, assigns it provisionally to erectus. However, a paper by Gabounia et al. (2002) defines a new species for the Dmanisi material, Homo georgicus, named for its country of origin. (Interestingly, this paper, published in a French journal, Human Palaecontology and Prehistory [Palontologie humaine et prhistoire] lists Vekua and Lordkipanidze among its authors. Both paleoanthropologists are also listed as authors on the initial D2700 paper published in Nature.) Earlier specimen D2600, a mandible unearthed in the year 2000, is the holotype for the species; the paper includes specimens D211 (a mandible), D2280 (cranium), and D2282 (cranium) within the new species. H. georgicus is, then, defined by the mosaic of Habiline and erectus-like features that the Dmanisi material displays: larger dimensions, unique sagittal keeling, pronounced protrusion of paralacteal internal torus mandibularis2, a particularly long alveolar arcade, large-sized canines, and P1 crown morphology (a disto-lingual bulge [talonide] and the oblique orientation of the principle axis, the presence of two clearly individualized roots from the neck [mesial and distal root], each of which bears a vertical line [Gabounia et al. 2002:244]). Gabounia et al. mention the D2700 cranium and associated mandible, D2735, when de1 Keeling along the sagittal midline, the generally depressed appearance of the parietal surfaces, the shape of the temporal squama, and the transverse expansion of the base relative to the low vault all make the skull look more like a small H. erectus than H. habilis (Vekua et al. 2002:88). 2 This feature is a small torus on the mandible, overwhelmingly bilateral in incidence, that is usually present near the premolars, above the attachment site for the mylohyoid muscle (per Saunders et al. 2002).

Hopkins 5 scribing notable differences in robusticity among the Dmanisi cranial material. These differences, they contend, are well encompassed within one species and explain them through sexual dimorphism: The differences between mandibles D2600 and D2113 and do not determine the coexistence of two species in Georgia at the beginning of the Lower Pleistocene. A gracile group, with D 211, D2280, D2282 and the latest discoveries D2700 and D2735, are in contact with a more robust group including D2600. They are the expression of a marked sexual dimorphism among only one species Homo georgicus sp. nov (Gabounia et al. 2002:244). It is interesting to note that many of the other papers about Dmanisi paleoanthropological material, including those containing the same authors, do not mention this H. georgicus species definition. A 2007 study of postcranial remains from the Dmanisi site emphasizes that they, too, look remarkably intermediate, possessing some Habiline and some erectus-like features; this study nevertheless omits mention of H. georgicus.4 A 2006 paper by Rightmire et al. that summarizes data on Dmanisi cranial remains provides insight into the species definition and its followers or supporters: The Dmanisi paleodeme can be designated as a subspecies [of H. erectus]. If it is agreed that, despite large corpus size, the D2600 mandible can be accommodated within this population, then the appropriate nomen is H. erectus georgicus. However, Gabunia et al. (2002) have argued that D2600 documents a different species from H. erectus. If this interpretation is verified by new discoveries, then a subspecies name other than H. erectus georgicus will have to be selected. On morphological grounds, it can be argued that the group from which the skulls are drawn is close to a stem from which later more derived populations are evolved (Rightmire et al. 2006:140). There seem to be a few sorts of opinions within academic literature on D2700 and the Dmanisi material in general. There are those who consider the material to fall within H. erectus, those who consider that some or all of the material falls within H. erectus georgicus, and those who feel some or all of the material belongs within the species H. georgicus. The first of these is the most prevalent position.
3 D2600 is generally larger and more robust than D211 (per Vekua et al. 2002, Gabounia et al. 2002, and Lieberman 2007). 4 The morphology of the upper and lower limbs from Dmanisi exhibits a mosaic of traits reflecting both selection for improved terrestrial locomotor performance and the retention of primitive characters absent in later hominins. The length and morphology of the hindlimb is essentially modern, and the presence of an adducted hallux and plantar arch indicate that the salient aspects of performance in the leg and foot, such as biomechanical efficiency during long-range walking and energy storage/return during running, were equivalent to modern humans. However, plesiomorphic features such as a more medial orientation of the foot, absence of humeral torsion, small body size, and low encephalization quotient suggest that the Dmanisi hominins are postcranially largely comparable to earliest Homo (cf. H. habilis). Hence, the first hominin species currently known outside Africa did not possess the full suite of derived locomotor traits apparent in African H. erectus and later hominins (Lordkipanidze et al. 2007:309).

Hopkins 6 Phylogenetically, paleoanthropologists seem to agree that D2700 lies somewhere between H. habilis and H. erectus. Some consider that the Dmanisi material may be ancestral to Asian H. erectus populations in Java and China. There is currently no direct evidence for such continuity outside common geography, applicable dates, and similar morphology. Just as the transition from the australopithecines to genus Homo is not extremely well defined, with paleoanthropologists defining H. habilis and H. rudolfensis in some circumstances, the transition from early Homo to Homo erectus is not extremely clear. Nevertheless, paleoanthropologists seem to agree that D2700 and the Dmanisi material in general are in some way ancestral to more definite H. erectus populations, given the similarities it shares with later H. erectus. This interpretation remains consistent among those who call D2700 H. georgicus and those who call it H. erectus, though all parties note the similarities the specimen shares with earlier Habilines, KNM-ER 1813 in particular. discussion: d2700 and migration out of africa, the habilis/erectus transition Eugene Dubois defined Homo erectus in 1891 from fossils discovered in Java, Indonesia. Initially described as Pithecanthropus erectus based on a skull cap and pathological, though remarkably human-like, femur, subsequent discoveries would help flesh out the species definition in order that it could be situated within the hominin lineage, somewhere before Neandertals and anatomically modern humans. Later, some paleontologists made a distinction between Asian and African specimens and thus forged a division within specimens previously known as H. erectus; the Asian specimens would continue to be known as Homo erectus where the African specimens became known as Homo ergaster. There are still some paleontologists who consider the differences between regions to fit comfortably within a single species, H. erectus. While this debate does not directly involve D2700 or any other Dmanisi material, it is symptomatic of the paleontological climate into which D2700 came after it was discovered in 2001. No matter whether one considers the differences significant enough to warrant separating Asian and African specimens into different species, H. erectus is the first hominin species that we have indisputably found outside of Africa. For the purposes of my discussion, I will consider erectus and ergaster to be at least similar enough for a very close groupingthe earliest erectoid specimen is KNM-ER 3733, dated around 1.75 or 1.8myo. This is, you will surely note, coincides exactly with ranges proposed for all the Dmanisi material based on a variety of dating methods. This implies first that erectus or erectus-like species were able to travel extremely quickly as they migrated beyond the African continent. Transcontinental migration also has behavioral implications as well, though it does not make migrating hominins worthy of any new genius; they could have been following food sources as environments shifted, particularly considering that hominins around the erectus period lived in a climactic paradigm of unique dynamism. Nevertheless, there must be something unique about the mental abilities of migrating hominins relative to their forebears. Otherwise, there seems to be little outside of environmental conditions that would necessarily limit a hominin from migrating when environments nearby were sufficient. It is still unlikely that migrating hominins left Africa out of any sort of wanderlust. This interpretation ignores the pragmatic reasons that a group of hominins might want to migrate in favor of a more salient perception: a desire to see and know the world abroad.

Hopkins 7 A number of other assumptions about the traits necessary for transcontinental migration were prevalent in the academic literature prior to uncovering the Dmanisi material. D2700 and the other Dmanisi crania are collectively responsible for problematizing, if not debunking altogether, many of these assumptions. D2700 was instrumental in discrediting the idea that transcontinental migration would require a significantly increased brain size, a great jump in size over the Habilines (Vekua et al. 2002, Rightmire et al. 2006, Lordkipanze et al. 2006, Lieberman 2007). D2700, a cranium dated at ~1.75myo with a cranial capacity around 600cc, was clearly not such a large jump above the habilines. What this means, however, may depend on adherence with a certain interpretation of the Dmanisi material. Those who consider the Dmanisi hominins to represent a new species, e.g. Homo georgicus, tend to consider this proof that a Habiline population, particularly H. habilis because of the congruence between KNM-ER 1813 and D2700 (among others), migrated out of Africa prior to the advent of H. erectus, may have given rise to H. erectus populations in the region and in Asia and gave rise at least to the Dmanisi hominins. This position is not mutually exclusive with one that places the material within H. erectus, but it does have an implication upon the nature and efficiency of locomotion within these hominins. Later, more definite Homo erectus specimens exhibit a remarkably human-like bipedal pattern with a few notable differences that may signify functional differences useful for longdistance running; H. erectus generally narrower pelvis and not-quite-human femora may have allowed for more efficient running. Bramble and Lieberman (2004) believe that this characteristic may be a hallmark of the genus Homo rather than a synapomorphic trait for erectus and subsequent hominins. This interpretation may suggest that Homo habilis, for example, migrated out of Africa in part due to its increased ability, to be fully realized in erectus, for marathon running. Lieberman is, however, among the majority of paleontoligists that place Dmanisis fossils in Homo erectus (see Lieberman 2007). As mentioned previously, D2700 is among a group of crania that the Dmanisi team describe as relatively more gracile than the holotype for the Homo georgicus species, the D2600 mandible. If sexual dimorphism can account for the differences, which include marked disparities in overal robusticity and molar size, between D2600 and the other Dmanisi fossils, we must consider the possible range of variation for these fossil species. A quick inspection of any sizeable human population can establish that Homo sapiens is a variable species, ranging greatly in size and sometimes in minutial morphology. Some humans may even possess, for example, a sagittal keel. Without treating this issue for humans in a greater capacity here, it seems that individual life history and environmental conditions play a remarkably significant role in determining individual morphology. D2700 and, of course, all other fossil species are now extinct and therefore do not allow paleontologists to attain a representative sample of variation for a given species. It is therefore difficult to say what is and is not an acceptable amount of variation for any species. We can make estimates and construct various hypotheses around these estimates, but ultimately we must rely on the amount of variation we see within extant species to determine whether variation is too great for one species to contain. To do this, paleontologists use a statistical calculation called the coefficient of variation or cv on a trait-by-trait basis. For brain size (endocranial volume), modern human populations yield a cv of 8.5. A composite cv for brain size in Dmanisi

Hopkins 8 fossils is 12.5, and the composite number for the entirety of erectus fossils older than 1myo is 17.5, both well outside the range of variation within H. sapiens (Lieberman 2007). Does this indicate that D2700 and its bretheren belong within a separate species? Not necessarily. Estimates of species-wide variation for fossil species through cv calculation are inherently problematic because, again, we can only see a subset of the entire species, particularly for hominins. We must also consider the significance of cranial capacity in our assessment of D2700 when mulling over its paleontological impact. Modern humans have a cranial capacity around 1350cc on average but exhibit a wide range of variation for this measurement. Even individuals significantly below this average cranial capacity possess modern human intelligence and, more generally, modern human intellectual potential. One might reasonably logic the correct position that overall brain size is a less significant determinant of intellectual faculties. Nevertheless, paleontologists use brain size as one of several diagnostic measurements for new hominin species. There are other means by which paleontologists infer intellectual capacity. The advent of stone tools in the archaeological record was one of the controversial diagnoses used to define Homo habilis, the first member of the genus Homo. Primatologists widely observe that many non-human primates use tools of various sorts; the diagnostic nature of the stone tools for habilis lies in the supposition that associated hominins were intentionally creating the tools in question. These stone tools, simply worked, belong to the long-standing Oldowan tool tradition. Homo erectus is the first species associated with the more complex Acheulean tool tradition, tools among which show signs of bifacial preparation. This Acheulean tradition likely requires a greater degree of spatial intelligence and some amount of imaginary intent, as such bifacial working is not necessary to make an effective tool; the earlier, simpler Oldowan tradition includes stone tools that were sufficiently useful for a number of tasks. The intelligence required to make an Acheulean hand-axe, which tend to maximize the available cutting edge, extending it fully around the perimeter of a stone tool, is therefore believed to be greater than for an Oldowan tool. We only find Oldowan tools at the Dmanisi site. This does not necessarily indicate that hominins could not or even did not produce Acheulean-like tools, but it does at least raise the possibility that Dmanisis hominins were not able to envision shapes or forms well enough to produce a hand axe, i.e. that they lacked the intelligence necessary to produce an Acheulean tool. D2700s small brain size would seem to be consistent with this notion, but it is, in my opinion, misleading to conclude much about the intellectual abilities of the Dmanisi hominins or any other hominin based on these methods. That they did or did not produce stone tools of a certain complexity says actually very little about their intellectual capacity. The level of sexual dimorphism that may be present among the Dmanisi hominins is an interesting issue that we may investigate along size this, comparing for example the D2600 mandible and D2700/D2735. The Homo georgicus definition stipulates that Dmanisi exhibits a relatively great degree of sexual dimorphism, but we could also account for differences in size through personal life history. Just as overall size can vary greatly among modern humans, it seems possible that nutritional intake could account for the differing specimen sizes at Dmanisi.

Hopkins 9 We thus have an issue of sample size when situating Dmanisi hominins and understanding the full extent of their significance for hominin evolution. There are a number of possibilities that further study may validate or debunk. Any more conclusive pronouncements can only come with continued investigation. Clearly, the situation is more complex than we can imagine. It remains unclear where D2700 exactly belongs within the hominin lineage. In the words of Rightmire et al. (2006:140), it [may be] close to a stem group from which later species of Homo are evolved. As of this point in time, conclusive evidence for an anagenetic relationship between Dmanisi hominins and later species has not been found. It seems likely that such a relationship exists, based on the presence of similar traits within D2700 in particular. In future expeditions at Dmanisi, paleontologists hope to find, among other things, the cranium associated with mandible D2600 in order that they can gain a better understanding of variation within local fossils. I am willing to place D2700 within Homo erectus and thus agree with the incredibly quick transcontinental migration that comes with the interpretation. It is important to remember that, while this migration did occur quickly by a geological perspective, that is not to say it occurred en masse. The Dmanisi hominin population could represent an early migration in a series of concurrent migrations. I admit that I am not completely confident that it is possible for all the regional erectus populations to evolve together so that certain traits would reach ubiquity or near-ubiquity within global populations. Nevertheless, we know that current human populations have remained a single species despite great geographic distance. It does not take many migration events, i.e. breeding between populations, to maintain species consistency. Still, my consideration of this issue quickly leads to ambivalence. What is the range of variation for a fossil species? We have no way of conclusively knowing this. Asking this question necessarily compels another: at what point should variation warrant establishing a new species? Indeed, this is a pressing question for paleontoligists and should not be taken lightly. To conclude on a more philosophical note, it is necessary to remember that human evolution is richer than we are able to know. Hopefully further study will produce a more conclusive understanding of D2700 and its relation to more primitive and more derived hominins. conclusion D2700 is a hominin cranium, dated around 1.75 million years old. It was discovered in Dmanisi, Georgia and exhibits a morphology that is, overall, a mix of Habiline and Homo erectus-like features. D2700 debunked the notion that the typical H. erectus brain size, around 900cc, was necessary for transcontinental migration. Taxonomically and phylogenetically, the cranium falls between the Habilines and H. erectus and may have been ancestral to other erectus populations in Europe and Asia.

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