Newly recognized Pleistocene human teeth from Tabun Cave, Israel

Seven human teeth from Tabun Cave, Israel, curated at the Natural History Museum London since 1955, are of uncertain provenance and identity. They are all from the upper dentition, without duplications, and are characterized by a similar preservation. The Catalogue of Fossil Hominids (1975) suggested that they might have derived from Tabun Layer A (Bronze Age to Recent). However, one of us (AC) noted some distinctive features of these teeth that warranted further study. They are here assigned to a single individual, Tabun BC7. Their morphology and metrics were then compared with the frequency of Late Pleistocene and Early Holocene groups from Europe, North Africa and Middle East. A fragment of the right M3 crown of Tabun BC7 was removed for ESR and U-analysis, and it was determined that only samples from Layer B have similar dose values. Using the sediment dose values of layer B, preliminary age estimates of 82 +/- 14 ka (early U-uptake) and 92+/-18 ka (linear uptake) were obtained. U-series disequilibrium determined from other samples attributed to Layer B resulted in a U-uptake history close to linear uptake, giving a very comparable age estimate of 90(+30)(-16) ka. The dose value previously obtained on an enamel fragment from the Tabun C1 dentition is nearly double the value measured for BC7, and tentative age estimates for C1 were in the range of 143+/-37 ka. However, due to uncertainties in the exact provenance of the human fossils, we cannot confirm that C1 is older than the new tooth sampled here, and both C1 and BC7 can be attributed to Layer B on chronological grounds. On the basis of chronology, dental morphology and metrics, the specimen named Tabun BC7 was identified as a probable Neanderthal.     

Neanderthal skeleton from Tabun: U-series data by gamma-ray spectrometry

The Neanderthal hominid Tabun C1, found in Israel by Garrod & Bate, was attributed to either layer B or C of their stratigraphic sequence. We have used gamma-ray spectrometry to determine the²³⁰Th/²³⁴U and²³¹Pa/²³⁵U ratios of two bones from this skeleton, the mandible and a femur. The ages calculated from these ratios depend on the uranium uptake history of the bones. Assuming a model of early U (EU) uptake the age of the Tabun C1 mandible is 34±5 ka. The EU age of the femur is 19±2 ka. The femur may have experienced continuous (linear) U uptake which would give an age of 33±4 ka, in agreement with the mandible's EU age, but implies marked inhomogeneity in U uptake history at the site. These new age estimates for the skeleton suggest that it was younger than deposits of layer C. This apparent age is less than those of other Neanderthals found in Israel, and distinctly younger than the ages of the Skhul and Qafzeh burials. This suggests that Neanderthals did not necessarily coexist with the earliest modern humans in the region. All of the more complete Neanderthal fossils from Israel are now dated to the cool period of the last glacial cycle, suggesting that Neanderthals may have arrived in this region as a result of the southward expansion of their habitable range. The young age determined for the Tabun skeleton would suggest that Neanderthals survived as late in the Levant as they did in Europe.     

U-series and ESR analyses of bones and teeth relating to the human burials from Skhul

In order to resolve long-standing issues surrounding the age of the Skhul early modern humans, new analyses have been conducted, including the dating of four well-provenanced fossils by ESR and U-series. If the Skhul burials took place within a relatively short time span, then the best age estimate lies between 100 and 135 ka. This result agrees very well with TL ages obtained from burnt flint of 119+/-18 ka (Mercier et al., 1993). However, we cannot exclude the possibility that the material associated with the Skhul IX burial is older than those of Skhul II and Skhul V. These and other recent age estimates suggest that the three burial sites, Skhul, Qafzeh and Tabun are broadly contemporaneous, falling within the time range of 100 to 130 ka. The presence of early representatives of both early modern humans and Neanderthals in the Levant during Marine Isotope Stage 5 inevitably complicates attempts at segregating these populations by date or archaeological association. Nevertheless, it does appear that the oldest known symbolic burials are those of early modern humans at Skhul and Qafzeh. This supports the view that, despite the associated Middle Palaeolithic technology, elements of modern human behaviour were represented at Skhul and Qafzeh prior to 100 ka.     

Timing of the Lower to Middle Paleolithic boundary: new dates from the Levant

This paper reviews the numerical dates available for the late Lower Paleolithic and early Middle Paleolithic in the Levant. We also present here new electron spin resonance dates for the late Lower Paleolithic sites of Holon, Yabrud I and Oumm Qatafa. Irrespective of dating techniques used, the ages of these sites converge on oxygen isotope stage 7 at roughly 215+/-30 ka. Similarly, dates for early Middle Paleolithic sites in the region, with the exception of Tabun, fall within oxygen isotope stage 7, suggesting a relatively rapid transition from Lower to Middle Paleolithic. In the light of these findings, the "early chronology" for the region, based on the TL dates on burned flint from Tabun, is discussed.     

The Vindija Neanderthal scapular glenoid fossa: comparative shape analysis suggests evo-devo changes among Neanderthals

Although the shape of the scapular glenoid fossa (SGF) may be influenced by epigenetic and developmental factors, there appears to be strong genetic control over its overall form, such that variation within and between hominin taxa in SGF shape may contain information about their evolutionary histories. Here we present the results of a geometric morphometric study of the SGF of the Neanderthal Vi-209 from Vindjia Cave (Croatia), relative to samples of Plio-Pleistocene, later Pleistocene, and recent hominins. Variation in overall SGF shape follows a chronological trend from the plesiomorphic condition seen in Australopithecus to modern humans, with pre-modern species of the genus Homo exhibiting intermediate morphologies. Change in body size across this temporal series is not linearly directional, which argues against static allometry as an explanation. However, life history and developmental rates change directionally across the series, suggesting an ontogenetic effect on the observed changes in shape (ontogenetic allometry). Within this framework, the morphospace occupied by the Neanderthals exhibits a discontinuous distribution. The Vindija SGF and those of the later Near Eastern Neanderthals (Kebara and Shanidar) approach the modern condition and are somewhat segregated from both northwestern European (Neandertal and La Ferrassie) and early Mediterranean Neanderthals (Krapina and Tabun). Although more than one scenario may account for the pattern seen in the Neanderthals, the data is consistent with palaeogenetic evidence suggesting low levels of gene flow between Neanderthals and modern humans in the Near East after ca. 120-100 ka (thousands of years ago) (with subsequent introgression of modern human alleles into eastern and central Europe). Thus, in keeping with previous analyses that document some modern human features in the Vindija Neanderthals, the Vindija G₃ sample should not be seen as representative of ‘classic’ - that is, unadmixed, pre-contact - Neanderthal morphology.     

The Neanderthalian molar from Hunas, Germany.

In this paper, we present a well-preserved isolated human molar found in 1986 in the Hunas cave ruin, south-east Bavaria. The tooth was located at the bottom of layer F2, which belongs to a long stratigraphic sequence comprising faunal remains as well as archaeological levels (Mousterian). A stalagmite from layer P at the base of the stratigraphic sequence was recently dated to 79.373+/-8.237 ka (base) and 76.872+/-9.686 ka (tip) by TIMS-U/Th (Stanford University). We identified the tooth as a right (possibly third) mandibular molar. Characteristic parameters such as crown and root morphology, fissure pattern, enamel thickness, occlusal and interproximal wear, dental dimensions and indices, and radiological features indicate that the Hunas molar represents the tooth of a Neanderthal. This is corroborated by both the palaeontological and archaeological findings (Mousterian) of layer F2.     

Neanderthal parietal, vertebrate fauna, and stone artifacts from the Upper Pleistocene deposits of Contrada Ianní di San Calogero (Catanzaro, Calabria, Italy)

Human skeletal remains recovered at a new archaeological site, dating back to the Italian Mousterian age are analyzed. The finds consist of a left parietal of a child about 2-3 years old. The metrical, morphometrical, and morphological traits of this parietal are compared to those of other Neanderthal children. The geological features of the site, its fauna, and stone artifacts recovered there during quarry operations are also analyzed.     

Kinetics and mechanism of electron transfer between meso-tetra sulphonated phenyl porphyrin iron(III)-apomyoglobin and Fe(EDTA)2- complexes

The kinetics of electron transfer between Fe(EDTA)2- and meso-tetra sulphonated phenyl porphyrin iron(III)-apomyoglobin have been studied by applying stopped-flow mixing and monitoring photometric changes at soret band (429 nm). The studies were carried out at pH's 6, 6.5, 7, 7.5, and 8 and at temperature between 10 and 40 degrees C. The mechanism proposed on the basis of the dependence of kobsd on Fe(EDTA)2- concentrations at various pH's, followed the rate equation: kobsd = ka[H+] + Kakb/[H+] + Ka.[Fe(EDTA)2-] The values of rate parameters calculated using a weighted non-linear least-squares analysis were: ka, 528 +/- 2 sec-1; kb, 25 +/- 1 sec-1; and Ka, 2.0 +/- 0.1 microM at 25 degrees C and 0.5 M sodium phosphate, and those of thermodynamic parameters calculated by the Eyring equation were: delta H*, 8.1 +/- 0.3 kcal mole-1 and delta S*, -23.4 +/- 1.1 eu at pH 7 and 0.5 M sodium phosphate.     

Does Homo neanderthalensis play a role in modern human ancestry? The mandibular evidence

Data obtained from quantifying the upper part of the mandibular ramus (the coronoid process, the condylar process, and the notch between them) lead us to conclude that Neanderthals (both European and Middle Eastern) differ more from Homo sapiens (early specimens such as Tabun II, Skhul, and Qafzeh, as well as contemporary populations from as far apart as Alaska and Australia) than the latter differs from Homo erectus. The specialized Neanderthal mandibular ramus morphology emerges as yet another element constituting the derived complex of morphologies of the mandible and face that are unique to Neanderthals. These morphologies provide further support for the contention that Neanderthals do not play a role in modern human biological ancestry, either through "regional continuity" or through any other form of anagenetic progression.     

Neanderthals

Neanderthals are a group of fossil humans that inhabited Western Eurasia from approximately 300 to 30,000 years ago (ka). They vanished from the fossil record a few millennia after the first modern humans appeared in Europe (ca. 40 ka BP). They are characterized by a unique combination of distinctive anatomical features, and are found with stone tools of the Mousterian stone tool industry. Current consensus views them as a distinct Eurasian human lineage isolated from the rest of the Old World and sharing a common ancestor with modern humans sometime in the early Middle Pleistocene. The extreme cold of the European Ice Ages is considered at least partly responsible for the evolution of some of the distinctive Neanderthal anatomy, although other factors (functional demands, effects of chance in small populations) were probably also important. The causes for the Neanderthal extinction are not well understood. Worsening climate and competition with modern humans are implicated. Neanderthals were our sister species, much more closely related to us than the chimpanzees, our closest living relatives are today.     

Sapropels and the age of hominins Omo I and II, Kibish, Ethiopia

The provenance and age of two Homo sapiens fossils (Omo I and Omo II) from the Kibish Formation in southern Ethiopia have been much debated. Here we confirm that Omo I and the somewhat more primitive-looking Omo II calvariae are from similar stratigraphic levels in Member I of the Kibish Formation. Based on (40)Ar/(39)Ar age measurements on alkali feldspar crystals from pumice clasts in the Nakaa'kire Tuff, a tuffaceous bed in Member I just below the hominin levels, we place an older limit of 198+/-14ka (weighted mean age=196+/-2ka) for the hominins. A younger limit of 104+/-7ka (weighted mean age=104+/-1ka) is provided by feldspars separated from pumice clasts in the Aliyo Tuff in Member III. Geological evidence indicates rapid deposition of each member of the Kibish Formation, concurrent with deposition of sapropels in the Mediterranean Sea. The (40)Ar/(39)Ar age measurements, together with correlations with sapropels, indicate that the hominin fossils are close in age to the older limit. Our preferred estimate of the age of the hominins is 195+/-5ka, making them the earliest well-dated anatomically modern humans yet described.     

Extending the chronology of deposits at Blombos Cave, South Africa, back to 140 ka using optical dating of single and multiple grains of quartz

Optically stimulated luminescence (OSL) measurements are reported for both single aliquots (of two different sizes) and single grains of quartz from deposits within Blombos Cave. Ages have been obtained for six sediments from the Middle Stone Age (MSA) occupation levels and for two sterile sands, one underlying the archaeological sediment and one overlying the Later Stone Age occupation levels. The ages for the archaeological sediments were obtained from single-grain measurements that enabled unrepresentative grains to be rejected. The MSA occupation levels have ages that, within error limits, are in stratigraphic order and fall between the OSL age for the oldest dune sand (143.2+/-5.5 ka) and a previously published OSL age for the sterile sand ( approximately 70 ka) that separates the Middle and Later Stone Age deposits. The earliest MSA archaeological phase, M3, from where fragments of ochre were found as well as human teeth, is dated to 98.9+/-4.5 ka, coinciding with the sea-level high of oxygen isotope substage 5c. The cave then appears to be unoccupied until oxygen isotope substage 5a on the basis of four OSL ages for archaeological phase M2, ranging from 84.6+/-5.8 to 76.8+/-3.1 ka; these levels contained large hearths and bone tools. An age of 72.7+/-3.1 ka was obtained for the final MSA archaeological phase, M1, from which deliberately engraved ochre and shell beads were recovered along with bifacial stone points. We conclude that the periods of occupation were determined by changes in sea level, with abundant sources of seafood available in times of high sea level and with the cave being closed by the accumulation of large dunes during periods of low sea level, such as during oxygen isotope stages 4 and 6.     

Transport of butyryl-L-carnitine, a potential prodrug, via the carnitine transporter OCTN2 and the amino acid transporter ATB(0,+)

L-carnitine is absorbed in the intestinal tract via the carnitine transporter OCTN2 and the amino acid transporter ATB(0,+). Loss-of-function mutations in OCTN2 may be associated with inflammatory bowel disease (IBD), suggesting a role for carnitine in intestinal/colonic health. In contrast, ATB(0,+) is upregulated in bowel inflammation. Butyrate, a bacterial fermentation product, is beneficial for prevention/treatment of ulcerative colitis. Butyryl-L-carnitine (BC), a butyrate ester of carnitine, may have potential for treatment of gut inflammation, since BC would supply both butyrate and carnitine. We examined the transport of BC via ATB(0,+) to determine if this transporter could serve as a delivery system for BC. We also examined the transport of BC via OCTN2. Studies were done with cloned ATB(0,+) and OCTN2 in heterologous expression systems. BC inhibited ATB(0,+)-mediated glycine transport in mammalian cells (IC(50), 4.6 +/- 0.7 mM). In Xenopus laevis oocytes expressing human ATB(0,+), BC induced Na(+) -dependent inward currents under voltage-clamp conditions. The currents were saturable with a K(0.5) of 1.4 +/- 0.1 mM. Na(+) activation kinetics of BC-induced currents suggested involvement of two Na(+) per transport cycle. BC also inhibited OCTN2-mediated carnitine uptake (IC(50), 1.5 +/- 0.3 microM). Transport of BC via OCTN2 is electrogenic, as evidenced from BC-induced inward currents. These currents were Na(+) dependent and saturable (K(0.5), 0.40 +/- 0.02 microM). We conclude that ATB(0,+) is a low-affinity/high-capacity transporter for BC, whereas OCTN2 is a high-affinity/low-capacity transporter. ATB(0,+) may mediate intestinal absorption of BC when OCTN2 is defective.     

Pharmacology of the leukotriene antagonist verlukast: the (R)-enantiomer of MK-571.

Verlukast (MK-679) (3-[(3-(2-(7-chloro-2-quinolinyl)-(E)-ethenyl)phenyl)[3-(dimethylamino)- 3- oxopropyl)thio)methyl)-thio)propionic acid) is a potent and selective inhibitor of [3H]leukotriene D4 binding in guinea-pig (IC50 = 3.1 +/- 0.5 nM) and human (IC50 = 8.0 +/- 3.0 nM) lung homogenates and dimethyl sulfoxide differentiated U937 cell membrane preparations (IC50 = 10.7 +/- 1.6 nM) but is essentially inactive versus [3H]leukotriene C4 binding in guinea-pig lung homogenates (IC50 values of 19 and 33 microM). Functionally, when tested at 60 nM, it antagonized contractions of guinea-pig trachea (GPT) induced by leukotriene C4, leukotriene D4, and leukotriene E4 (respective-log KB values of 8.6, 8.8, and 8.9) and contractions of human trachea (HT) induced by leukotriene D4 (-log KB value 8.3 +/- 0.2). In contrast, verlukast (20-200 nM) failed to antagonize contractions of GPT induced by leukotriene C4 in the presence of 45 mM L-serine borate. Intravenous (i.v.) and aerosol verlukast antagonized bronchoconstriction (BC) induced in anaesthetized guinea pigs by i.v. leukotriene D4 but did not block BC to arachidonic acid or histamine. Intraduodenal verlukast (0.25 mg/kg) antagonized leukotriene D4 (0.2 micrograms/kg) induced BC in guinea pigs. Oral and aerosol administration blocked leukotriene D4-induced BC in conscious squirrel monkeys. Orally administered compound also blocked ovalbumin-induced BC in conscious sensitized rats treated with methysergide (3 micrograms/kg). The pharmacological profile for verlukast is similar to that of the racemic compound, MK-571. Verlukast is currently in clinical development for the treatment of asthma and related diseases.     

On the age of Border Cave 5 human mandible

An enamel fragment from the Border Cave 5 specimen was analysed with non-destructive ESR combined with laser ablation ICP-MS for uranium profiling. We obtained an age of 74+/-5 ka which fits exactly into the chronological framework that has been previously established for Border Cave by a variety of dating techniques. The result lays at rest the view that BC5 could be of Iron Age, as was implied by (Journal of Human Evolution, 31 (1996) 499) based on nitrogen contents and infra-red splitting factors.     

Direct ESR dating of a Pliocene hominin from Swartkrans

Two fragments of a hominin tooth (Australopithecus robustus) and two bovid teeth from the Hanging Remnant of the Swartkrans Formation were analysed with ESR. Research was complicated by the fact that the samples came from a curated collection and their precise provenance is unknown. The environmental dose rate was reconstructed by a series of in situ gamma spectrometric measurements and elemental analyses of a range of sediment samples. U-series isotopic analyses indicated that each of the teeth had a significantly different uranium uptake history, rendering the assumptive early U-uptake and linear U-uptake models ineffective. ESR and U-series data were combined to calculate open system ages, resulting in a best estimate of 1630+/-160 ka for the Hanging Remnant. An open-system model which provides the maximum age for given U-series and ESR measurements yielded an estimate of about 2100 ka.Two bovid teeth from Member 2, previously estimated to be between 1.0 and 2.0 Ma, yielded age estimates of between about 100 and 200 ka. No known geochemical processes are likely to explain this severe age underestimation. We conclude that these samples are of Middle to Upper Pleistocene age and their presence in Member 2 was either due to reworking or inadequate stratigraphical discrimination of these deposits.     

Correlation of the KHS Tuff of the Kibish Formation to volcanic ash layers at other sites, and the age of early Homo sapiens (Omo I and Omo II)

Hominin specimens Omo I and Omo II from Member I of the Kibish Formation, Ethiopia are attributed to early Homo sapiens, and an age near 196 ka has been suggested for them. The KHS Tuff, within Member II of the Kibish Formation has not been directly dated at the site, but it is believed to have been deposited at or near the time of formation of sapropel S6 in the Mediterranean Sea. Electron microprobe analyses suggest that the KHS Tuff correlates with the WAVT (Waidedo Vitric Tuff) at Herto, Gona, and Konso (sample TA-55), and with Unit D at Kulkuletti in the Ethiopian Rift Valley. Konso sample TA-55 is older than 154 ka, and Unit D at Kulkuletti is dated at 183 ka. These correlations and ages provide strong support for the age originally suggested for the hominin remains Omo I and Omo II, and for correlation of times of deposition in the Kibish region with formation of sapropels in the Mediterranean Sea. The Aliyo Tuff in Member III of the Kibish Formation is dated at 104 ka, and correlates with Gademotta Unit 15 in the Ethiopian Rift Valley.