Gocha Kiladze’s research while affiliated with University of Georgia and other places

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Publications (11)


Early Pleistocene stratigraphy, sedimentary environments, and formation contexts at Dmanisi in the Georgian Caucasus
  • Article

November 2022

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149 Reads

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4 Citations

Journal of Human Evolution

Reid Ferring

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Sebastien Nomade

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The Early Pleistocene site of Dmanisi is now well known for its large number of fossils of early Homo erectus as well as associated artifacts and faunal remains, recovered mainly in pipe-related geologic features. Testing in the M5 unit 100 m to the west of the main excavations revealed a thick stratigraphy with no evidence of pipes or gullies, indicating that the geologic record at Dmanisi included spatially distinct sedimentary environments that needed further investigation. Here we report the results of a geoarchaeological program to collect data bearing on contexts and formation processes over a large area of the promontory. That work has defined over 40,000 m² of in situ deposits with artifacts and faunas. Stratum A ashes bury the uppermost Mashavera Basalt, which we have dated to 1.8 Ma in the M5 block. The Stratum A deposits contain stratified occupations that accumulated quickly and offer good potential for recovery of in situ materials. Stratum B1 deposits above the A/B unconformity include all of the pipe and gully facies at Dmanisi, reflecting a brief but very intense phase of geomorphic change. Those deposits contain the majority of faunas and all of the hominin fossils. B1 slope facies offer excellent formation contexts away from the piped area, and all B1 deposits are sealed by Stratum B2 over the whole promontory. Strata B2 to B5 register a return to slope facies, with no further evidence of pipes or gullies. Those deposits also present excellent contexts for recovery of in situ occupations. Overall, Dmanisi's geologic history preserves an exceptional record of the activities and environmental context of occupations during the first colonization of Eurasia.


Author Correction: The bears from Dmanisi and the first dispersal of early Homo out of Africa
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  • Full-text available

February 2021

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187 Reads

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2 Citations

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

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Figure 1. Ursus etruscus upper dental series. D50-52-55 in (A) occlusal view, (B) lingual view, (C) buccal view. D2214-2215 in (D) occlusal view, (E) lingual view, (F) buccal view. D4713 in (G) occlusal view. D2533 in (H) occlusal view. Scale bar = 5 cm.
Figure 2. Ursus etruscus mandible (D355). (A) right buccal view, (B) left buccal view and (C) occlusal view. Scale bar = 5 cm.
Figure 3. Morphometric analyses of tooth measurements in extant and extinct bears. (A) Bivariate scatterplot of the specimens' scores on the two first principal components for the lower (mandibular) teeth. (B) Bivariate scatterplot of the specimens' scores on the two first principal components for the upper (maxillary) teeth. (C) Bivariate scatterplot of the specimens' scores on the two first canonical functions for discriminating among dietary types in ursids with the tooth measurements of the lower dentition. (D) Bivariate scatterplot of the specimens' scores on the two first canonical functions derived from the upper dentition. The points for U. etruscus and U. deningeri correspond to the sample means of the sites studied. Numbers indicate for the sites of the fossil specimens.
Figure 4. A microwear features of grinding and slicing facets of Ursus etruscus from Dmanisi. (A) Slicing facet of a left lower p4 (D36). (B) slicing facet of a left m3 (D355). (C) grinding facet of a left m1 (D1277) and grinding facet of left m2 (D4940).
The bears from Dmanisi and the first dispersal of early Homo out of Africa

November 2019

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810 Reads

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20 Citations

We report on the taxonomy and paleodiet of the bear population that inhabited the emblematic palaeoanthropological Early Pleistocene (1.8 Ma) site of Dmanisi (Georgia), based on a dual approach combining morphometrics and microwear of upper and lower teeth. Given that the teeth of Ursus etruscus Cuvier, 1823 from Dmanisi show considerable size variability, their systematic position has been debated. However, a comparative study of the coefficients of variation for tooth size measurements in several modern bear species shows that the variability in tooth size of the ursid population from Dmanisi could result from sexual dimorphism. The analysis of tooth microwear indicates that these bears inhabited a mixed environment of open plain with forest patches, where they had a browsing diet with a substantial contribution of meat and/or fish. Comparative tooth morphometric analyses of modern ursids and fossil U. etruscus indicate that this extinct species had an omnivorous behavior similar to that of extant brown bears. The ecological interactions of the Dmanisi bears with other members of the large mammals community, including the first hominins that dispersed out of Africa, are discussed in the light of this new evidence.


Early Pleistocene enamel proteome from Dmanisi resolves Stephanorhinus phylogeny

September 2019

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1,333 Reads

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169 Citations

Nature

The sequencing of ancient DNA has enabled the reconstruction of speciation, migration and admixture events for extinct taxa¹. However, the irreversible post-mortem degradation² of ancient DNA has so far limited its recovery—outside permafrost areas—to specimens that are not older than approximately 0.5 million years (Myr)³. By contrast, tandem mass spectrometry has enabled the sequencing of approximately 1.5-Myr-old collagen type I⁴, and suggested the presence of protein residues in fossils of the Cretaceous period⁵—although with limited phylogenetic use⁶. In the absence of molecular evidence, the speciation of several extinct species of the Early and Middle Pleistocene epoch remains contentious. Here we address the phylogenetic relationships of the Eurasian Rhinocerotidae of the Pleistocene epoch7–9, using the proteome of dental enamel from a Stephanorhinus tooth that is approximately 1.77-Myr old, recovered from the archaeological site of Dmanisi (South Caucasus, Georgia)¹⁰. Molecular phylogenetic analyses place this Stephanorhinus as a sister group to the clade formed by the woolly rhinoceros (Coelodonta antiquitatis) and Merck’s rhinoceros (Stephanorhinus kirchbergensis). We show that Coelodonta evolved from an early Stephanorhinus lineage, and that this latter genus includes at least two distinct evolutionary lines. The genus Stephanorhinus is therefore currently paraphyletic, and its systematic revision is needed. We demonstrate that sequencing the proteome of Early Pleistocene dental enamel overcomes the limitations of phylogenetic inference based on ancient collagen or DNA. Our approach also provides additional information about the sex and taxonomic assignment of other specimens from Dmanisi. Our findings reveal that proteomic investigation of ancient dental enamel—which is the hardest tissue in vertebrates¹¹, and is highly abundant in the fossil record—can push the reconstruction of molecular evolution further back into the Early Pleistocene epoch, beyond the currently known limits of ancient DNA preservation.


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Early Pleistocene enamel proteome sequences from Dmanisi resolve Stephanorhinus phylogeny

September 2018

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1,153 Reads

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9 Citations

Ancient DNA (aDNA) sequencing has enabled unprecedented reconstruction of speciation, migration, and admixture events for extinct taxa. Outside the permafrost, however, irreversible aDNA post-mortem degradation has so far limited aDNA recovery within the ~0.5 million years (Ma) time range. Tandem mass spectrometry (MS)-based collagen type I (COL1) sequencing provides direct access to older genetic information, though with limited phylogenetic use. In the absence of molecular evidence, the speciation of several Early and Middle Pleistocene extinct species remain contentious. In this study, we address the phylogenetic relationships of the Eurasian Pleistocene Rhinocerotidae using ~1.77 million years (Ma) old dental enamel proteome sequences of a Stephanorhinus specimen from the Dmanisi archaeological site in Georgia (South Caucasus). Molecular phylogenetic analyses place the Dmanisi Stephanorhinus as a sister group to the woolly ( Coelodonta antiquitatis ) and Merck's rhinoceros ( S. kirchbergensis ) clade. We show that Coelodonta evolved from an early Stephanorhinus lineage and that this genus includes at least two distinct evolutionary lines. As such, the genus Stephanorhinus is currently paraphyletic and its systematic revision is therefore needed. We demonstrate that Early Pleistocene dental enamel proteome sequencing overcomes the limits of ancient collagen- and aDNA-based phylogenetic inference, and also provides additional information about the sex and the taxonomic assignment of the specimens analysed. Dental enamel, the hardest tissue in vertebrates, is highly abundant in the fossil record. Our findings reveal that palaeoproteomic investigation of this material can push biomolecular investigation further back into the Early Pleistocene.


Fig. 1. Left: Location of Dmanisi and Kvabebi sites in the frame of the Neogene deposits of Southern Caucasus. Right: Eastern Paratethys stages and southern Kakheti litostratigraphy. Note Ackchagilian stage and equivalents in grey. 
Fig. 2. Lithostratigraphy and paleomagnetism of the Kvabebi section. 
Fig. 4. Magnetostratigraphic correlation of the Dmanisi and Kvabebi sites with other well-calibrated Pliocene-Pleistocene sections from Africa and Western Europe. Chronology for Kada Hadar Member by Renne et al., 1993. See text for references of the other sections. 
Fig. 5. Comparison of Genus-Level Faunal Resemblance Indexes (GFRI) illustrates the zoogeographical affinities of the Kvabebi large mammal fauna. GFRI are made using both Simspon's and Dice indexes. The Dice index is calculated as: 2A/(2A þ B þ C), where A is the number of taxa present in both faunas, B is the number of taxa present in fauna 1, but absent in fauna 2, and C is the number of taxa present in fauna 2 but absent in fauna 1. Simpson's faunal resemblance index and is calculated as: A/(A þ E) where E is the smaller of B or C (data source updated from Turner et al., 1999 and NOW database, public release 030717 available at http. //www.helsinki.fi/science/now/).
The Pliocene-Pleistocene succession of Kvabebi (Georgia) and the background to the early human occupation of Southern Caucasus

December 2009

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1,008 Reads

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53 Citations

Quaternary Science Reviews

This article analyzes and discusses the chronological and zoogeographic context of the Pliocene site of Kvabebi in order to shed light into the background of the early human occupation of Eurasia, as evidenced by the early Pleistocene site of Dmanisi. New paleontological and paleomagnetic research has allowed this site to be placed in a reverse interval which can be identified as chron 2An.1r. The age of this site is therefore close to 3.07 Ma, coeval to the Hadar beds of the Afar Depression and slightly older than the last hipparionine faunas in Europe, such as Villarroya in Spain. The fauna of Kvabebi is composed of a number of species common to Europe and Asia, but also includes others with African affinities, such as Kvabebihyrax kachethicus, Protoryx heinrichi, Parastrepsiceros sokolovi and Gazella postmitilinii. However, the presence these species cannot be explained as the result of a Pliocene dispersal from Africa, but rather as the last remnants of the so-called Greek-Iranian province.



Figure 2: Dmanisi postcranial elements.a, Remains of subadult individual. D2724, left clavicle; D2716/D2855, right/left first rib; D2717, eleventh rib; D2673/D2674/D2721/D2713/D2672, vertebrae C2/C3/Th3/Th10/L1; D2715/D2680, right/left humerus; D3160, left femur; D2679/D3480, distal phalanges of hand; D2671/D2669, right metatarsal I/IV; D2670, first distal phalange of right foot. b, Remains of large adult individual. D4166, right scapula; D4162/D4161, right/left clavicles; D4063, right second rib; D4507, left humerus; D4167, right femur; D3418, right patella; D3901, right tibia; D4110, left talus; D2021/D4165, right metatarsals III/IV; D4508, left metatarsal V; D3877, distal phalange of foot. c, d, Remains of small adult individuals. D3479, right metatarsal III; D4111, right medial cuneiform; D3442, right metatarsal I.
Figure 3: Long-bone shape and proportions.a, Tibial mediolateral distal width versus maximum length. b, Humeral versus femoral length. c, Tibial versus femoral length. Stars, Dmanisi Homo; X, AL288-1 (Australopithecus afarensis); diamond, BOU-VP-12/1; triangle, KNM-WT15000 (H. erectus); Z, recent Homo sapiens; plus signs, Pan troglodytes; Y, Gorilla gorilla; squares, Pongo pygmaeus.
Postcranial Evidence from Early Homo from Dmanisi, Georgia

October 2007

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2,965 Reads

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646 Citations

Nature

The Plio-Pleistocene site of Dmanisi, Georgia, has yielded a rich fossil and archaeological record documenting an early presence of the genus Homo outside Africa. Although the craniomandibular morphology of early Homo is well known as a result of finds from Dmanisi and African localities, data about its postcranial morphology are still relatively scarce. Here we describe newly excavated postcranial material from Dmanisi comprising a partial skeleton of an adolescent individual, associated with skull D2700/D2735, and the remains from three adult individuals. This material shows that the postcranial anatomy of the Dmanisi hominins has a surprising mosaic of primitive and derived features. The primitive features include a small body size, a low encephalization quotient and absence of humeral torsion; the derived features include modern-human-like body proportions and lower limb morphology indicative of the capability for long-distance travel. Thus, the earliest known hominins to have lived outside of Africa in the temperate zones of Eurasia did not yet display the full set of derived skeletal features.


Fig. 1. The map of the localities with the genus Simocyon; 1.Arkneti, 2.Dzedzvtakhevi. 
Fig. 2. Simocyon primigenius from Dzedzvtakhevi, left mandible (DZ. 351). A) labial view, B) lingual view, C) occlusal view. Scale bar 20 mm. 
Table 2 . Measurements of P 4 from Arkneti and of two other species of Simocyon, measurements are given in millimeters. Measurements are taken from Wang, 1997 and Peigne et al., 2005. 
Fig. 3. Simocyon sp. From Arkneti, right P 4 (ARK. 5/25 IPB). A) labial view, B) lingual view, C) occlusal view. Scale bar 20 mm. Conclusion-As remarked above, there are only two sites in the Trans-Caucasia where Simocyon fossil remains are found. One of them is Arkneti, its age is determined as the Late Meotian (Gabunia, 1959; Meladze, 1967, 1985). Revision of the P 4 from Arkneti reveals that this specimen has no diagnostic characters that could help the species determination, but by, size it is close to S. batalleri and S. primigenius from China (Yushe and Baode specimens) (Wang, 1997; Peigne et al., 2005). Taxonomic status of the Arkneti Simocyon remains unresolved until new fossils are found. It is not difficult to distinguish Dzedzvtakhevi material from most of the species of the genus Simocyon, as only one species S. primigenius has completely lost second and third premolars. By measurements the Dzedzvtakhevi mandible approximates to the European form of S. primigenius. Finding of S. primigenius in Dzedzvtakhevi confirms the Latest Miocene age of this locality proposed by Vekua & Trubikhin (1988), as this species is known only from MN 12-13 localities (Werdelin, 1996; Roussiakis, 2002; Peigne et al., 2005). 
Simocyon from the Late Miocene of Georgia

January 2007

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178 Reads

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2 Citations

saqarTvelos teritoriaze pirveli cnoba gvari Simocyon-is aRmoCenis Sesaxeb gvxvdeba 1954 wels, sadac l. gabunia gvamcnobs adgilsapovebel arkneTSi sxva hiparionuli faunis warmomadgenlebTan erTad am gvaris arsebobas. mogvianebiT g. melaZis mier aRweril iqna gvari Simocyon-is zeda ybis meoTxe premolari arkneTidan (zeda mioceni). ukanasknel wlebSi adgilsapovebel ZeZvTaxevSi avtorebis mier aRmoCenilia Simocyon-is qveda ybis marcxena toti. ZeZvTaxevisa da arkneTis masalebis Seswavlam da reviziam gviCvena, rom ZeZvTaxevis Simocyon-i miekuTvneba S. primigenius-is evropul formas, romelic ZiriTadad cnobilia gvian miocenur adgilsapovebel pikermiidan (MN 12-13). Simocyon primigenius-is aRmoCena adgilsapovebel ZeZvTaxevSi adasturebs 1988 wels a. vekuas da v. trubixinis mier am adgilsapoveblis asakis meotisad (MN 12-13) gansazRvras. samwuxarod arkneTis masala arasakmarisia imisaTvis, rom saxeobamde ganisazRvros, Tumca misi mcire zomebiT igi axlos dgas S. batalleri-Tan daS. primigenius-is Cinur formasTan (iuSe, baode).


A Fourth Hominin Skull From Dmanisi, Georgia

November 2006

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273 Reads

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125 Citations

The Anatomical Record Part A Discoveries in Molecular Cellular and Evolutionary Biology

Newly discovered Homo remains, stone artifacts, and animal fossils from Dmanisi, Republic of Georgia, provide a basis for better understanding patterns of hominin evolution and behavior in Eurasia ca. 1.77 million years ago. Here we describe a fourth skull that is nearly complete, lacking all but one of its teeth at the time of death. Both the maxillae and the mandible exhibit extensive bone loss due to resorption. This individual is similar to others from the site but supplies information about variation in brain size and craniofacial anatomy within the Dmanisi paleodeme. Although this assemblage presents numerous primitive characters, the Dmanisi skulls are best accommodated within the species H. erectus. On anatomical grounds, it is argued that the relatively small-brained and lightly built Dmanisi hominins may be ancestral to African and Far Eastern branches of H. erectus showing more derived morphology.


Citations (9)


... The first contributions deal with two crucial aspects for understanding the entire Dmanisi site and its paleobiological content: the geological and chronological background , and the taphonomy Margvelashvili et al., 2022). Ferring et al. (2022) provide an exhaustive summary of the absolute and relative ages of the Dmanisi serial occupations and associated paleontological remains, as well as of the key formation processes that bear on preservation patterns. The stratigraphic correlations between the excavation areas documenting an area of >40,000 m 2 with in situ deposits of artifacts and faunas are presented and discussed in detail. ...

Reference:

Introduction to special issue: The biotic context of the Early Pleistocene hominins from Dmanisi (Georgia, southern Caucasus)
Early Pleistocene stratigraphy, sedimentary environments, and formation contexts at Dmanisi in the Georgian Caucasus
  • Citing Article
  • November 2022

Journal of Human Evolution

... Apart from this large hyaenid, carnivores are well represented in BLD and FN3, including two mustelids (Martellictis ardea and Meles meles), three canids (Vulpes alopecoides, C. mosbachensis, and Lycaon lycaonoides), one ursid (Ursus etruscus), and three felids (Lynx pardinus, Megantereon whitei and Homotherium latidens) Palmqvist et al., 2023a;Rodríguez-Gómez et al., 2016a, 2016bRos-Montoya et al., 2021). This rich carnivore guild resembles those found in other late Early Pleistocene archaeological and paleontological sites of Eurasia, such as Dmanisi in Georgia, dated to ~ 1.8 Ma (Medin et al., 2019;Bartolini-Lucenti et al., 2021, Pirro Nord in Italy, dated to 1.6-1.3 Ma (Petrucci et al., 2013), and Vallonnet in France, dated to ~ 1.1 Ma (Moullè et al., 2006), or at the early Acheulian site of 'Ubeidiya in Israel, dated to 1.6-1.2 ...

The bears from Dmanisi and the first dispersal of early Homo out of Africa

... This is important as often ZooMS and other ancient proteomic analyses are used to make ecological inferences about ancient ecosystems. This can include proposing phylogenies (Buckley 2013;Welker et al. 2015Welker et al. , 2017Cappellini et al. 2019;, inferring anthropogenic effects on ancient biodiversity (Harvey et al. 2019a;Fossile et al. 2023), informing on hominin subsistence behaviours (Sinet-Mathiot et al. 2019Torres-Iglesias et al. 2024) including tool manufacture (Ashby et al. 2015;McGrath et al. 2019;Martisius et al. 2020;Bradfield et al. 2021;Evans et al. 2023), and the development of animal husbandry (Janzen et al. 2021;Elnaggar et al. 2022). By only sampling a small proportion of excavated faunal material, outliers and uncontrollable sampling biases can heavily influence research outputs. ...

Early Pleistocene enamel proteome from Dmanisi resolves Stephanorhinus phylogeny

Nature

... The two species composing the genus Rhinoceros are also closely related (Tougard et al., 2001); the bones of R. unicornis and R. sondaicus having sometimes been confused with each other (Groves & Leslie, 2011). Conversely, the phylogenetic position of Dicerorhinus sumatrensis remains debated (Willerslev et al., 2009;Gaudry, 2017), this species being considered alternately as sister taxon of the two African species (Antoine, Duranthon & Welcomme, 2003;Cappellini et al., 2018), of the two Rhinoceros species (Tougard et al., 2001;Welker et al., 2017) or of all four other rhino species (Fernando et al., 2006;Piras et al., 2010). Our analyses reveal equally contrasting relationship patterns, with Dicerorhinus sumatrensis more closely resembling African species for some bones (radius, ulna and tibia) and Asiatic ones for the others (humerus, femur and fibula). ...

Early Pleistocene enamel proteome sequences from Dmanisi resolve Stephanorhinus phylogeny

... Four land mammal sites are known from the Shiraki Formation: Dzedzvtakhevi, Vashlovani, Tetri Udabnos Seri, and Dzhaparidze. Brief publications exist only for the Dzhaparidze and Dzedzvtakhevi faunas (Orlov 1941;Vekua and Trubikhin 1988;Vanishvili et al. 2007). ...

Simocyon from the Late Miocene of Georgia

... Depositional environments represented by strata exposed in the KFTB vary from shallow marine to terrestrial, and the strata are predominantly siliciclastic, with general coarsening-upward trends observed throughout most stratigraphic sections (e.g., Agustí et al., 2009;Forte et al., , 2015aVan Baak et al., 2013;Lazarev et al., 2019Lazarev et al., , 2021). Strata exposed in the KFTB were deposited in environments influenced by both the development of the GC and KFTB (e.g., Forte et al., , 2015a and large-magnitude (~1000 m) base-level changes of the Caspian Sea during the late Cenozoic (e.g., Popov et al., 2006;van Baak et al., 2017;Krijgsman et al., 2019;Lazarev et al., 2021). ...

The Pliocene-Pleistocene succession of Kvabebi (Georgia) and the background to the early human occupation of Southern Caucasus

Quaternary Science Reviews

... Ma (ref. 18), has yielded five well-preserved crania, associated mandibles and various postcranial elements [19][20][21][22][23] . Together with a rich record of faunal remains and lithic artefacts, the Dmanisi ensemble has provided unique insights into the palaeobiology of an early Homo population outside Africa, its life history, social organization and cognitive capabilities 24,25 . ...

Anthropology: The earliest toothless hominin skull

Nature

... 18), has yielded five well-preserved crania, associated mandibles and various postcranial elements [19][20][21][22][23] . Together with a rich record of faunal remains and lithic artefacts, the Dmanisi ensemble has provided unique insights into the palaeobiology of an early Homo population outside Africa, its life history, social organization and cognitive capabilities 24,25 . Here we focus on a subadult individual that is represented by cranium D2700, mandible D2735 and various postcranial elements 20,22 . ...

A Fourth Hominin Skull From Dmanisi, Georgia
  • Citing Article
  • November 2006

The Anatomical Record Part A Discoveries in Molecular Cellular and Evolutionary Biology

... The crania indicate a very early form of H. erectus, with a cranial capacity as low as 775 cc (Rightmire et al., 2006). The post-cranial bones indicate that it was fully bipedal but with a slightly different walk from ours and was perhaps adapted for endurance running (Lordkipanidze et al., 2007;Lieberman, 2007). This would have been advantageous by increasing the chances of fleeing from a carnivore, or of chasing a prey animal until it was exhausted. ...

Postcranial Evidence from Early Homo from Dmanisi, Georgia

Nature