REPORTS 15. P. Mansky, C. K. Harrison, P. M. Chaikin, R. A. Register, N. Yao, Appl. Phys. Lett. 68, 2586 (1996). 16. K. Shin et al., Nano Lett. 2, 933 (2002). 17. E. Huang, L. Rockford, T. P. Russell, C. J. Hawker, Nature 395, 757 (1998). 18. This work was supported by NSF Grant Opportunities for Academic Liaison with Industry award (DMI-0217816), the U.S. Department of Energy under contract DE-FG03-88ER45375 and the Materials Research Science and Engineering Centers at the University of Massachusetts (DMR-0213695) and Santa Barbara (DMR-0080034). D.Y.R. thanks the support by the Postdoctoral Fellowship program of Korea Science and Engineering Foundation in Korea. We thank C. Silvas of Dow Chemical for providing the brominated BCB. Estimating Duration and Intensity of Neoproterozoic Snowball Glaciations from Ir Anomalies Bernd Bodiselitsch,1 Christian Koeberl,1* Sharad Master,2 Wolf U. Reimold2 The Neoproterozoic glaciations supposedly ended in a supergreenhouse environment, which led to rapid melting of the ice cover and precipitation of the so-called cap carbonates. If Earth was covered with ice, then extraterrestrial material would have accumulated on and within the ice and precipitated during rapid melting at the end of the glaciation. We found iridium (Ir) anomalies at the base of cap carbonates in three drill cores from the Eastern Congo craton. Our data confirm the presence of extended global Neoproterozoic glaciations and indicate that the duration of the Marinoan glacial episode was at least 3 million, and most likely 12 million, years. The Snowball Earth hypothesis (1–3) states that the Sturtian Eabout 710 million years ago (Ma)^ and Marinoan glaciations (about 635 Ma) were of global extent and lasted for several million years each. A variation of this hypothesis, called the Slushball Earth, requires milder conditions without substantial equatorial sea ice (4, 5). The Snowball Earth glaciations would have ended abruptly in a greenhouse environment, whereas the Slushball would have experienced a slower deglaciation. Silicate weathering and photosynthesis, which are the major sinks for CO2 at present, would have been inhibited by the ice that covered both continents and oceans, and huge amounts of CO2 could have accumulated in the atmosphere. Greenhouse gases, particularly large amounts of CO2 (up to È0.12 bar), would have been needed to overcome the high albedo caused by the glaciation and would have caused rapid melting (6). This scenario has led to an estimate for the duration of a Snowball glaciation of È4 million to 30 million years (My), under the assumption of a modern rate for CO2 outgassing from subaerial volcanism, with no air-sea gas exchange, lower solar luminosity in the Neoproterozoic, and reduced pelagic deposition of carbonate to reduce the release of volcanic CO2 at con1 Department of Geological Sciences, University of Vienna, Althanstrasse 14, A-1090 Vienna, Austria. 2 Impact Cratering Research Group, School of Geosciences, University of the Witwatersrand, Private Bag 3, Johannesburg 2050, South Africa. *To whom correspondence should be addressed. E-mail: [email protected] vergent margins (7). This estimate is in broad agreement with a 96- to 910-My duration derived from the d13C isotopic excursion in the Otavi (North Namibia) cap carbonates (2). Multiple magnetic polarity reversals within the Elatina Formation (South Australia) suggest a minimum time duration of several hundred thousand to 1 million years for the Marinoan glacial epoch (8). If Earth was indeed covered by ice for long periods, extraterrestrial material would have accumulated on and within the ice and would have been precipitated at the base of the cap carbonates when the ice melted. To determine whether such an extraterrestrial signal is present at the base of the cap carbonates, we studied three drill cores that intersected diamictite/cap carbonate boundaries from the Lufilian tectonic arc in the Democratic Republic of the Congo and Zambia of the Eastern Congo craton (Fig. 1) (9). We measured the concentrations of 43 elements by neutron activation analysis and x-ray fluorescence spectrometry, and we measured the Ir content by multiparameter coincidence spectrometry after neutron activation (10). Ir anomalies were found at the base of all cap carbonates after the Marinoan and Sturtian glaciations in the Nguba and Kundelungu Groups at Kipushi, as well as after the Sturtian glacial deposits in the Nguba Group at Chambishi. Ir and other platinum-group elements are typical proxies for extraterrestrial material, in which they are much more abundant than in Earth_s upper mantle and crust. The average Ir concentration in cosmic matter is about 4.6 105 parts per trillion (ppt) (11), which is greater www.sciencemag.org SCIENCE VOL 308 Supporting Online Material www.sciencemag.org/cgi/content/full/308/5719/236/ DC1 Materials and Methods Fig. S1 21 October 2004; accepted 22 February 2005 10.1126/science.1106604 by a factor of about 104 than in crustal terrestrial rocks. The accretion of extraterrestrial material during the Neoproterozoic is thought to have been dominated by the delivery of interplanetary dust particles (IDPs) over a time scale of millions of years (12). IDP accretion rates vary over periods ranging from a few thousand years (13) to millions of years (14). During glaciations, IDPs, as well as extraterrestrial matter from asteroids and comets that struck the Earth during that time (9), would have accumulated on the ice sheet. Substantial Ir anomalies up to almost 2 parts per billion (ppb) mark the base of the cap carbonate deposits (Fig. 2) (9). Before ascribing an extraterrestrial origin to the Ir, possible terrestrial sources, including reduced sedimentation rates, increased meteor ablation rates, accretion of extraterrestrial material or of terrestrial dust (such as volcanic airborne particles), and anoxic conditions connected with sulfide precipitation in seawater need to be considered. Our geochemical data clearly indicate that the (substantial) Ir anomalies at the base of the cap carbonates are derived from extraterrestrial sources, whereas the other (smaller) Ir anomalies disappear or are greatly diminished when ratios with other elements are used Esee (9) for detailed considerations^. Thus, these variations can be attributed to changing deposition rates or to the dissolution of an extraterrestrial signal caused by the addition of sediment. Further confirmation that the Ir anomaly at the Kipushi Petit Conglomerat/cap carbonate transition (KPCCT) is of extraterrestrial origin comes from Cr/Ir and Au/Ir ratios of 2.9 103 and 0.11, respectively. These values are similar to those of C1 chondrites (carbonaceous chondrites type 1) (5.5 103 and 0.29, respectively) (15). Cr/Ir and Au/Ir ratios from other samples show higher values ranging from 8.4 104 to 1.8 106 and 1.1 to 80, respectively (Fig. 2). In contrast to the sharp Ir anomaly at the KPCCT, the Nguba Group Ir anomaly extends over a broader interval, between 236.91 and 237.93 m, with a maximum of 648 ppt of Ir at 237.93 m. The integrated Ir concentrations are, however, as high as those at the other location. This Ir anomaly does not occur exactly at the Kipushi Grand Conglomerat/cap carbonate transition (KGCCT), but 0.63 m above, at 238.56 m in the cap carbonate succession (Fig. 2). Ir/element ratios with Fe, Cs, and Al, compared with the Grand Conglomerat and cap carbonate succession, are much higher in 8 APRIL 2005 239 REPORTS B EL T K IB A R A N 100km n, ia nt b r me am di ec se d Pr lly vere ca o lo c T EL B 240 N IA D EN B U BANGWEULU BLOCK LUFILIAN ARC 10° Archaean to lower Proterozoic I Katanga Supergroup II III CHAMBISHI to R ec en t BAROTSE BASIN c oz oi Sinda Batholith IV Chipata Granite Hook Granite Complex ZAMBEZI BELT Zambezi Belt ae Z MS Zambezi Belt Archaean to lower Proterozoic CHOMA-KALOMO BLOCK 100km 15° MOZAMBIQUE BELT L N 0 IR U M ID E B EL T Lu an gw aV all ey KIPUSHI Pa l the Ir-enriched interval. Ni/Ir ratios of 1.1 104 at 236.91 m and 2.6 104 at 237.93 m, respectively, are similar to the C1 chondritic value of 2.3 104 and are distinctly lower than the values throughout the cap carbonate succession. Also, Au/Ir and Cr/Ir ratios of 0.46 and 1.05 104, respectively, at 237.93 m are also similar to the corresponding chondritic values of 0.29 and 0.55 104 (15), whereas ratios below and above the Ir anomaly are comparatively much higher. No indicators exist for hydrothermal, hydrogenous, or volcanic influences or for strong anoxic or reducing conditions that could produce an Ir anomaly. Thus, this enrichment was also likely caused by admixture of extraterrestrial material. The fact that the Ir anomaly after the Sturtian glaciation is characterized by a broader peak of high Ir, Co, Cr, and Ni values near the base of the cap carbonates could be the result of a transgression and slightly delayed deposition of the cap carbonates (3). In the Chambishi core, the Ir anomaly is located closer to the top of the Sturtian glacial deposits (only 9 cm above the top of the Grand Conglomerat) at 147.72 m than in the Kundelungu succession. The Ir anomaly at the Chambishi Grand Conglomerat/cap carbonate transition (CGCCT) is distinguished from the other Ir anomalies by an Ir/Fe ratio that is twice as high (Fig. 2). Changes in debris rainout during the glaciation caused by basal melting of ice sheets, floating glacier tongues, or episodic migration of calved icebergs into regions of warmer water (16) would affect the relative abundances of Fe and Ir but not their ratio. Thus, the occurrence of new source material in the glacial deposits may indicate warming periods during the glacial episode. The Ir/Al ratio and element/Cs ratios for Ce, Co, Cr, Eu, Fe, Hf, Sc, Ta, and Th at all Ir anomalies in this succession show values that are similar to the background values, except the Ir/Cs ratio at the Ir anomaly at 170.70 m, which is twice as high as that at the other Ir anomalies. However, the carbonate succession over the CGCCT Ir anomaly shows very high Ir/Fe, Ir/Al, and Ir/Cs ratios and low Co/Ir and Cr/Ir ratios, as well as Ir abundances of 45 to 60 ppt. This is very high, if we assume that cap carbonates precipitated very rapidly (17–19). Furthermore, at 144.96 m, strong evidence for the input of volcanogenic material is signaled by the very high element/Al ratios of Th, Ta, Hf, K, and Ti, which indicate the inclusion of a silicic volcanic component and/or a Hawaiian-like volcanic component (20, 21) (Fig. 2). However, it does seem that the deposited material was mostly extraterrestrial. It was probably not deposited at once but rather swept into the sea from uplands after a long glaciation, which was recorded for the Sturtian period in the Nguba Group succession from Kipushi. ZIMBABWE CRATON 30° 25° Basement inliers in Lufilian Arc I - External fold-thrust belt Strike-slip fault II - Domes Region Thrust III - Synclinorial belt International boundary IV - Katanga High L - Lusaka Borehole MSZ - Mwembeshi Shear Zone Fig. 1. Geological map of the Lufilian Arc in Congo and Zambia and part of the Zambezi belt, showing location of drill holes [modified after (29, 30)]. Extraterrestrial influx rates (total mass) estimated from ocean sediment studies based on Ir and Os measurements from mid–Pacific Ocean sediment samples range from 78 109 T 26 109 g yearj1 in the time interval between 33 and 67 Ma (14) to 30 109 T 15 109 g yearj1 between 0 and 80 Ma (22). This agrees well with other flux estimations from different time intervals from today to 700,000 years ago equal to È40 109 g yearj1, derived from 3He analyses in Pacific, Atlantic, and Indian Ocean sediments (23–25). If we assume an Ir concentration of 500 ng per g of extraterrestrial material and an Earth surface area of 5.1 1018 cm2, Ir influx rates of 8 T 3 ng cmj2 Myj1 to 3 T 1.5 ng cmj2 Myj1 are implied. However, the Ir flux estimated by (26), which was calculated from fossil meteorites in Lower Ordovician marine limestones from Kinnekulle, Sweden, at È30 ng cmj2 Myj1, indicates that accretion rates of extraterrestrial material were an order of magnitude higher during the Early Ordovician than at present. We made the conservative assumption that all Ir at the KPCCT is extraterrestrial (thus, no background needs to be calculated and 8 APRIL 2005 VOL 308 SCIENCE removed) and that no extraterrestrial material was deposited by major impact events. Then we integrated the elevated Ir abundances that represent the peak and took a rock density of 2.5 g cmj3 into account, to obtain an Ir concentration of 93 ng cmj2 at the KPCCT. If the Ir influx rates discussed above are used, a maximum duration of the Marinoan glaciation of 41 T 20 My and a minimum of 12 T 4 My are obtained for the Ir flux during 0 to 80 Ma, and a duration of È3 My if we assume the Ir input flux determined for the Early Ordovician. Nevertheless, because there probably was an unusually enhanced flux of extraterrestrial material in the Early Ordovician, we view the normal (0 to 80 Ma) values as the most representative, leading to a most likely duration of the Marinoan glaciation of 12 My. A problem in Neoproterozoic geology is the lack of reliable radiometric dates. For the Marinoan glaciation, age limits between 610 and 575 Ma are based on assumed relationships of Marinoan-to-Varanger glacial intervals from different locations (27), as well as estimations of sedimentation rate for the time between glacial deposition and for the Precambrian/Cambrian boundary (28). Recent www.sciencemag.org REPORTS Co/Ir [x10 -3 ] Cr/Ir [x10 -3 ] Ni/Ir [x10 -3 ] Au/Ir Ti/Al Middle Kundelungu Group Ir/Al [x10 -9 ] Upper Nguba Group Limestone Calcaire Rose Limestone Kakontwe 50 150 50 150 500 1500 300 900 20 50 0.06 0.10 Dolomitc Shale Middle Nguba Group Diamictite Petit Conglomerat Diamictite Grand Conglomerat 40 100 100 400 200 700 100 300 20 70 0.07 0.13 Finely Laminated Limestone (mm Scale) Mwashya Subgroup Laminated Shale Breccia Laminated AnhydriteDolomite Upper Roan Subgroup Nguba Group Roan Group Sturtian Glaciation - Chambishi MJZC9 Sturtian Glaciation - Kipushi KHI 115034HZ-S Marinoan Glaciation - Kipushi KHI 1150PVSSW Core Fig. 2. Stratigraphic A columns showing Ir Ir/Cs Depth Strat. Ir Ir/Fe Gr. Lith. abundances and se[x10 -6 ] [m] unit [ppt] [x10 -9 ] lected element ratio 0 Ku 1.2.3 profiles. (A) The middle part of the Kundelungu Ku 1.2.2 Group to the top of the 30 Nguba Group, Kipushi, Ku 1.2.1 Congo, showing Petit 60 Conglomerat from the Ku 1.1 Marinoan glaciation overlain by the Calcaire 90 Rose cap carbonates. Ng 1.2.3 (B) The middle of the 120 Nguba Group, Kipushi, Congo, showing Grand 600 1800 100 300 1000 4500 Conglomerat from the Sturtian glaciation over- B lain by the Calcaire de Kakontwe cap carbon180 ates. (C) The Sturtian Ng 1.2.1 Grand Conglomerat 210 overlain by the Calcaire de Kakontwe cap car240 bonates from the mid270 dle of the Nguba Group, further to the Mwashya Ng 1.1 300 Subgroup at the base of the Nguba Group, and the Upper Roan 100 450 10 40 1000 5000 Subgroup at the top of the Roan Group, C Chambishi, Zambia. 100 For tables of elemental Ng 1.2.1 concentrations, see (9). Analytical uncertainty 150 for Ir is 95 relative % Ng 1.1 (rel%) for values 9500 ppt and 5 to 15 rel% for lower values. Ana200 lytical uncertainties for elemental ratios are between 5 and 15 rel%. 250 Ku 1.2.3, dolomitic shale; Ku 1.2.2, dolomitic shale; Ku 1.2.1, Calcaire Rose; Ku 1.1, Petit Conglom300 erat; Ng 1.2.3, dolomitic shale; Ng 1.2.1, Calcaire de Kakontwe; Ng 1.1, 350 Grand Conglomerat; Strat. Unit, stratigraphic unit; Lith., lithology. 400 The dip of the beds at Kipushi is not vertical 100 400 30 90 100 500 but is about 70-. Thus, borehole depths do not represent true stratigraphic thickness. U-Pb dating indicates that the Marinoan glaciation took place at 635.5 T 1.2 Ma (9). The duration of glaciation in the Neoproterozoic is difficult to quantify because of uncertain age limits and nonuniform, noncontinuous sedimentary records. Previous methods used to estimate the duration of Neoproterozoic glaciation are as follows: magnetic polarity intervals (8), calculation of CO2 amounts needed to overcome a totally frozen Earth (6), and thermal subsidence modeling (2). The use of Ir anomalies located immediately at the transition from glacial deposits to cap carbonates is an another possible way to estimate the du- Laminated Dolomitic Shale 100 400 200 600 600 2000 500 1500 50 200 ration of long glacial epochs. The advantage of this method is that samples from only the transition region of an undisturbed stratum are needed, otherwise a permanent ice sheet must have existed during the glaciation. We calculated a minimum duration for the Marinoan glacial epoch of È3 My, assuming the same Ir flux as obtained for the Lower Ordovician, and a most likely duration of about 12 My, based on the average accretion rate. For the Sturtian event, no Ir anomaly was found directly at the transition from the Sturtian glacial deposits to cap carbonates in the Chambishi and Kipushi successions, but it www.sciencemag.org SCIENCE VOL 308 2 5 was found 9 and 63 cm above, respectively. Furthermore, the extraterrestrial Ir is distributed over a wider interval in the cap carbonate succession. This distribution implies that extraterrestrial material was accumulated in upland areas over thousands or millions of years and was then washed into the sea after deglaciation. Unlike the Marinoan cap carbonates, Sturtian cap carbonates lack a transgressive stage, which implies that seawater did not reach critical oversaturation with respect to carbonate until after the postglacial marine transgression (3). Therefore, the extraterrestrial Ir in a global ice cover would have been partly dispersed 8 APRIL 2005 241 REPORTS before the cap carbonate precipitated. This may explain the lack of a sharp Ir spike at the base of the Sturtian cap carbonate. Alternatively, during the Sturtian glacial epoch, Earth_s surface may not have been fully covered with ice on which extraterrestrial material could accumulate for a long time; however, the presence of banded iron formations in and below Sturtian glacials suggests that the ocean was ice-covered at that time (3). References and Notes 1. J. L. Kirschvink, in The Proterozoic Biosphere, J. W. Schopf, C. Klein, Eds. (Cambridge Univ. Press, Cambridge, 1992), pp. 51–52. 2. P. F. Hoffman, A. J. Kaufman, G. P. Halverson, D. P. Schrag, Science 281, 1342 (1998). 3. P. E. Hoffman, D. P. Schrag, Terra Nova 14, 129 (2002). 4. W. T. Hyde, T. J. Crowley, S. K. Baum, W. R. Peltier, Nature 405, 425 (2000). 5. T. J. Crowley, W. T. Hyde, W. R. Peltier, Geophys. Res. Lett. 28, 283 (2001). 6. K. Caldeira, J. F. Kasting, Nature 359, 226 (1992). 7. K. Caldeira, Geology 19, 204 (1991). 8. L. E. Sohl, N. Christie-Blick, D. V. Kent, Geol. Soc. Am. Bull. 111, 1120 (1999). 9. Materials and methods are available as supporting material on Science Online. 10. C. Koeberl, H. Huber, J. Radioanal. Nucl. Chem. 244, 655 (2000). 11. J. T. Wasson, in Meteorites: Their Record of Solar System History (Freeman, New York, 1985), p. 267. 12. Z. Ceplecha, Astron. Astrophys. 263, 361 (1992). 13. K. A. Parley, D. B. Patterson, Nature 378, 600 (1995). 14. F. T. Kyte, J. T. Wasson, Science 232, 1225 (1986). 15. E. Anders, N. Grevesse, Geochim. Cosmochim. Acta 53, 197 (1989). 16. H. Heinrich, Quat. Res. 29, 142 (1988). 17. J. D. Roberts, J. Geol. 84, 47 (1974). 18. I. J. Fairchild, in Sedimentology Review, V. P. Wright, Ed. (Blackwell, Oxford, 1993), pp. 1–16. 19. M. J. Kennedy, J. Sediment. Res. 66, 1050 (1996). 20. D. C. Noble, W. L. Rigot, H. R. Bowman, GSA Spec. Pap. 180, 77 (1979). 21. Basaltic Volcanism Study Project, in Basaltic Volcanism on the Terrestrial Planets (Pergamon, New York, 1981), p. 1286. 22. B. Peucker-Ehrenbrink, Geochim. Cosmochim. Acta 60, 3192 (1996). 23. F. Marcantonio et al., Nature 383, 705 (1996). The Brain of LB1, Homo floresiensis Dean Falk,1* Charles Hildebolt,2 Kirk Smith,2 M. J. Morwood,3 Thomas Sutikna,4 Peter Brown,3 Jatmiko,4 E. Wayhu Saptomo,4 Barry Brunsden,2 Fred Prior2 24. F. Marcantonio et al., Geochim. Cosmochim. Acta. 62, 1535 (1998). 25. F. Marcantonio et al., Earth Planet. Sci. Lett. 170, 157 (1999). 26. B. Schmitz, B. Peucker-Ehrenbrink, M. Lindström, M. Tassinari, Science 278, 88 (1997). 27. A. H. Knoll, M. R. Walter, Nature 356, 673 (1992). 28. J. P. Grotzinger, S. A. Bowring, B. Z. Saylor, A. J. Kaufman, Science 270, 598 (1995). 29. H. Porada, Precambrian Res. 44, 103 (1989). 30. R. M. Key et al., J. Afr. Earth Sci. 33, 503 (2001). 31. We thank H. Rice and F. Popp from the University of Vienna for discussions and comments. This work was supported by the Austrian Science Foundation. This is University of the Witwatersrand Impact Cratering Research Group publication no. 93. Supporting Online Material www.sciencemag.org/cgi/content/full/308/5719/239/ DC1 Materials and Methods Fig. S1 Tables S1 to S3 References and Notes 30 August 2004; accepted 18 January 2005 10.1126/science.1104657 tate shape comparisons (Fig. 1 and fig. S2). LB1_s shape most resembles that of ZKD XI, which is typical of classic H. erectus from China and Java (Trinil) (fig. S3). Both endocasts are noticeably wider caudally than rostrally (Fig. 1A), wider ventrally than dorsally (fig. S2), and relatively long and low in lateral profile (Fig. 1B). However, LB1 lacks the de- The brain of Homo floresiensis was assessed by comparing a virtual endocast from the type specimen (LB1) with endocasts from great apes, Homo erectus, Homo sapiens, a human pygmy, a human microcephalic, specimen number Sts 5 (Australopithecus africanus), and specimen number WT 17000 (Paranthropus aethiopicus). Morphometric, allometric, and shape data indicate that LB1 is not a microcephalic or pygmy. LB1’s brain/body size ratio scales like that of an australopithecine, but its endocast shape resembles that of Homo erectus. LB1 has derived frontal and temporal lobes and a lunate sulcus in a derived position, which are consistent with capabilities for higher cognitive processing. The type specimen of Homo floresiensis (LB1, female) (1) has a brain size of È400 cm3, which is similar to that of Australopithecus afarensis specimen AL 288-1 (Lucy) (2), who lived approximately 3.0 million years ago. Yet LB1_s species was associated with big-game stone technology, remains of Stegodon, and charred animal bones that hint at the use of fire and cooking. Its ancestors also had to cross the sea to reach the Indonesian island of Flores (3). Could a tiny hominin with an ape-sized brain really have engaged in such advanced behaviors? Some workers reject the notion that LB1 1 Department of Anthropology, Florida State University, Tallahassee, FL 32306, USA. 2Mallinckrodt Institute of Radiology, Washington University School of Medicine, St. Louis, MO 63110, USA. 3Archaeology and Palaeoanthropology, University of New England, Armidale, New South Wales 2351, Australia. 4Indonesian Centre for Archaeology, JI. Raya Condet Pejaten No. 4, Jakarta 12001, Indonesia. *To whom correspondence should be addressed. E-mail: [email protected] 242 represents a new species that was closely tied to H. erectus (1) and suggest instead that it was a pathological human microcephalic (4). To help address this debate, we compared threedimensional computed tomographic (3DCT) reconstructions of the internal braincase (virtual endocasts) that reproduce details of external brain morphology, including sulci, vessels, sinuses, cranial capacity, and shape (5–8), from LB1, an adult female chimpanzee, an adult female H. erectus (specimen ZKD XI), a contemporary woman, and a European microcephalic. To broaden taxonomic comparisons and supplement limited sample size, our analysis also included endocasts of the skulls of specimen Sts 5 (A. africanus), specimen KNM-WT 17000 (Paranthropus aethiopicus), 10 humans, 10 gorillas, 18 chimpanzees (9), an adult female pygmy, and five H. erectus. Our virtual cranial capacity estimate for LB1 is 417 cm3 (10). Virtual endocasts of the microcephalic, modern woman, H. erectus, and chimpanzee were scaled to 417 cm3 to facili- 8 APRIL 2005 VOL 308 SCIENCE Fig. 1. Comparisons of virtual endocasts of LB1 (center). (A) Dorsal views. (B) Right lateral views. Hs, H. sapiens; Pt, Pan troglodytes; mcHs, a human microcephalic; He, H. erectus. www.sciencemag.org
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