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New hominid fossils from Member 1 of the Swartkrans formation, South Africa

New hominid fossils from Member 1 of the Swartkrans formation, South Africa
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  New hominid fossils from Member 1 of the Swartkrans formation, South Africa Travis Rayne Pickering a , b , c , * , Jason L. Heaton c , d , Ronald J. Clarke b , Morris B. Sutton b , e , C.K. Brain c ,Kathleen Kuman b , f  a Department of Anthropology, University of Wisconsin e Madison, 1180 Observatory Drive, 5240 Social Science Building, Madison, WI 53706, USA b Institute for Human Evolution, University of the Witwatersrand WITS 2050, Johannesburg, South Africa c Plio-Pleistocene Palaeontology Section, Department of Vertebrates, Ditsong National Museum of Natural History (Transvaal Museum), Pretoria 0002, South Africa d Department of Biology, Birmingham-Southern College, Birmingham, AL 35254, USA e Centre for Language and Culture, University of Johannesburg, South Africa f  School of Geography, Archaeology and Environmental Studies, University of the Witwatersrand WITS 2050, Johannesburg, South Africa a r t i c l e i n f o  Article history: Received 10 February 2011Accepted 24 February 2012Available online 21 March 2012 Keywords: Swartkrans Member 1Early  HomoParanthropus robustus TeethFemurBody size sexual dimorphism a b s t r a c t Member 1 of the Swartkrans Formation is comprised of two sedimentary in 󿬁 lls, the Lower Bank (LB) andthe Hanging Remnant (HR). Together, the LB and HR preserve fossils of early  Homo  and  Paranthropusrobustus , Earlier Stone Age lithic artifacts, purported bone digging tools and butchered animal bones.Collectively, this evidence was the  󿬁 rst to establish the co-existence of two early Pleistocene hominidspecies and also led to inferences of plant root harvesting and meat-eating by one or both of thosespecies.  P. robustus  is the more abundant of the two hominids at Swartrkrans, represented in Member 1by hundreds of fossils that derive from at least 99 individuals. Thus, Swartkrans Member 1 stands as theworld ’ s single largest repository of that extinct species. Here we add to the Member 1 sample of hominidfossils with descriptions of 14 newly discovered specimens.   2012 Elsevier Ltd. All rights reserved. Introduction Swartkrans Cave (Gauteng Province, South Africa) (Fig. 1) hasyielded the world ’ s largest sample of   Paranthropus robustus 1 fossils(Broom and Robinson, 1952; Grine, 1989, 2004), in addition tofewer d but important d early  Homo  fossils, such as the partialcraniumSK847(Clarkeetal.,1970;ClarkeandHowell,1972;Clarke, 1977a). These hominid fossils, numbering over 400 individualspecimens, were recovered from Members 1 e 3 of the SwartkransFormation initially through the work of Robert Broom and JohnRobinson (e.g., Broom, 1949; Broom and Robinson, 1949, 1950,1952) and subsequently through the long-term excavations(1965 e 1986) of C.K. Brain (e.g., Brain, 1981, 2004a; Brain et al.,1988). More recently, the Swartkrans PaleoanthropologicalResearchProject(SPRP), initiated in 2005byCKB (Coordinator) andTRP (Director), discovered two additional fossil-bearing deposits inthe northeastern portion of the site (Sutton et al., 2009). One of these deposits, the Talus Cone Deposit (TCD), has already yieldedthree  P. robustus  teeth (Sutton et al., 2009). The TCD overliesa second deposit(and is separatedfromit byan unconformity) thatwe infer to be an eastward extension of the Member 1 Lower Bank(LB) sedimentary unit. This LB East Extension has so far yieldedtypical early Pleistocene macrofauna (unpublished data) thatmirrors that of the LB (see, Watson, 2004).TheSPRPalsorecovered13hominiddentalfossilsandafragmentofhominidproximalfemur fromexcavationsof the LBdeposits andsurveys of the Member 1  ex situ  breccia dumps. Here we presentdescriptions of and taxonomic allocations for these fossils. Swartkrans Member 1 Brain (2004b) provides a detailed hypothesis outlining theformationof the Swartkrans Cave sedimentary deposits. The recentSPRP excavations at Swartkrans update this hypothesis (Suttonet al., 2009). Member 1 contains two depositional subunits, theLB and the Hanging Remnant (HR), which are, respectively, theoldest and second oldest known of the remaining strata of theSwartkrans Formation. Pickering et al. (2011) provide U-Pb ageestimates derived from speleothem samples identi 󿬁 ed andcollected by the SPRP, and which effectively bracket all of Swartk-rans Member 1. Fig. 1 illustrates the position of the northwestportion of the HR overlying a basal speleothem. This speleothem *  Corresponding author. E-mail address: (T.R. Pickering). 1 TRP and JLH prefer the use of   Australopithecus robustus  over  Paranthropusrobustus . However, to conform to the current inclination of many specialists,including RJC, we employ the designation  P. robustus  throughout this paper. Contents lists available at SciVerse ScienceDirect  Journal of Human Evolution journal homepage: 0047-2484/$  e  see front matter    2012 Elsevier Ltd. All rights reserved.doi:10.1016/j.jhevol.2012.02.003  Journal of Human Evolution 62 (2012) 618 e 628  continues southward across the west wall of the cave, and it is alsobelow the LB deposit in that position. Uranium-Pb subsample SWK12, taken from this basal speleothem, yielded an age of 2.249  0.077Ma. Another speleothem (a capping 󿬂 owstone), fromwhich U-Pb subsample SWK 9 was extracted, is exposed along thecave ’ snorthwestwall,betweentheHRandtheoverlyingsedimentsof Swartkrans Member 2. Subsample SWK 9 is dated to1.706  0.069 Ma. It is important to note that Pickering et al. (2011)are in error when they state that the SWK 9  󿬂 owstone caps the LB;it does not, but instead caps the HR, dating the formation of the 󿬂 owstone following the Member 1 in 󿬁 lling and prior to theMember 2 in 󿬁 lling. Thus, Member 1 is bracketed by two datedspeleothems, which together provide the estimated age range forthat unit ’ s fossil and artifact samples. Age estimates of 2.248  0.052 Ma. (SWK 7) and 1.800  0.005 Ma. (SWK 5) for twoadditional speleothems that, respectively, under- and overlie themain body of the HR (i.e., that portion of the HR adhering to thenorth wall of the cave)(Fig.1) are each within error, respectively, of SWK 9 and SWK 12 (Pickering et al., 2011). Recently, burial datingby radioactive decay of cosmogenic nuclides in quartz inclusionsderived from the Member 1 sediments has corroborated the broadU-Pb results (Gibbon et al., in preparation). Pickering et al. (2011)contend that faunal data narrow the age bracket for the accumu-lation of the Member 1 paleontological and archaeological mate-rials down to w 1.9 e 1.8 Ma. Materials and methods SPRP   󿬁 eld methods During the course of his 1979 e 1986 excavations, Brain estab-lished a metal square meter grid over the Swartkrans surfacedeposits. In 2005, the SPRP re-established this grid using an EDMTotal Station. At that time, the SPRP also reinitiated excavations inthe northeastern portion of the uncalci 󿬁 ed LB (12 m sq). Eachsquare was excavated by quadrant (50 cm    50 cm) at 5 cm and10 cm depths. Each quadrant level was excavated separately andthen sieved using a 5 mm mesh over a 2 mm mesh screen. Thematerial from the 5 mm mesh was then wet sieved. The 2 e 5 mmfraction material was sorted to ensure the collection of smallbone and tooth fragments of macrovertebrates, microfaunalremains and stone artifact chips. The SPRP excavation in the LB isongoing, and individual squares are currently at varying depths.The SPRP also conducts casual searches of the srcinal Broomand Robinson calci 󿬁 ed breccia dumps. Most breccia blocks in thesedumps contain a few or no visible macrovertebrate fossils, but theoccasional interesting piece is discovered, such as the  P. robustus canine described below. Fossil analysis We provide descriptions of and taxonomic allocations for 13hominid fossils recovered from ourexcavations in the northeasternportion of the LB and one discovered by Isaac Makhele duringcasual searches of Broom and Robinson ’ s srcinal breccia dump,presumably composed of HR sediment blocks. Each dental spec-imen was examined using a low-power binocular microscope and,when possible, measurements were taken with a Paleo-Tech Con-cepts   Hillson e Fitzgerald dental caliper. Standard gross toothcrown measurements were taken to the nearest tenth of a milli-meter (e.g., Robinson, 1956; Tobias, 1967; Grine, 1984). When possible, we measured the cervico-occlusal (CO) height (measuredin the vertical plane) and labio/buccolingual interproximal (L/BL)breadth (measured in the horizontal plane) of interproximal facets Figure 1.  The left side of the  󿬁 gure is a plan view of Swartkrans Cave illustrating its major depositional units as exposed sur 󿬁 cially. Abbreviations: LB, Lower Bank of Member 1; HR,Hanging Remnant of Member 1; M3, Member 3; M4, Member 4. Diagonal hatching indicates the northwest extension of Member 1 deposits under the cave roof. Member 2 isremoved in this plan view in order to clarify positions of the Member 1 deposits. The approximate locations where U-Pb dating samples were obtained for dating by Pickering et al.(2011) are indicated by open stars and their sample numbers. On the right side of the  󿬁 gure are two photographs showing the  in situ  exposures of the sampled speleothems thatbracket the HR and LB in 󿬁 lls of Swartkrans Member 1 (M1), below Member 2 (M2), in the northwest portion of the site (U-Pb dating subsamples SWK 9 and 12) (top), and those thatbracket the HR in 󿬁 ll in the north-central portion of the site (U-Pb dating subsamples SWK 5 and 7) (bottom; adapted from Pickering et al. (2011; Fig. 5)). Subsample SWK 5 is dated 1.800    0.005 Ma.; SWK 7, 2.248    0.052 Ma.; SWK 9, 1.706    0.069 Ma.; SWK 12, 2.249    0.077 Ma. (Pickering et al., 2011). T.R. Pickering et al. / Journal of Human Evolution 62 (2012) 618 e 628  619  on dental specimens. Taxonomic determinations of teeth werebased on various features used previously to differentiate  Austral-opithecus ,  Paranthropus  and early  Homo  dentitions, including, butnot limited to: number of cusps; relative cusp proportions andocclusal wear pattern (e.g., Robinson, 1956; Tobias, 1965, 1967;Clarke, 1977b; Howell, 1978; Grine, 1984, 1985, 1989, 2005). Thesingle postcranial fossil was examined macro- and microscopicallyand measurements were taken on it using a Mitutoyo   digitalcaliper.Fossil specimens were labeled sequentially from 001 witha SWT1- pre 󿬁 x.  “ SW ”  refers to Swartkrans,  “ T ”  refers to theTransvaal Museum, the permanent repositoryof the fossils, and  “ 1 ” refers to the fact that the fossils derive from Member 1 of theSwartkrans Formation. Within that designation, an LB- pre 󿬁 xindicates fossils recovered from the Lower Bank, and HR- indicatesthose from the Hanging Remnant (e.g., SWT1/LB-1; SWT1/HR-2).The new system of designation was created to readily distinguishfossils recovered from Swartkrans by the SPRP from those recov-eredbyearlierresearchprojectsatthesite,whicharedesignatedbythe pre 󿬁 xes SK, SKW and SKX. Results Table 1 lists the newly discovered hominid fossils fromSwartkrans Member 1 and provides their taxonomic attributionsand excavation coordinates. Table 2 gives standard dimensions formeaningfully measurable dental specimens, along with compara-tivesummarystatisticsforpreviouslyrecoveredspecimensforeachrepresented element. Descriptions of the new specimens areprovided below, as well as justi 󿬁 cations for their taxonomicattributions. Upper central incisors (Fig. 2) One deciduous (SWT1/LB-15) and two permanent (SWT1/LB-13and SWT1/LB-16) upper central incisors were recovered from theLB; each derives from a separate hominid individual. All threespecimens are attributed to  P. robustus .SWT1/LB-15isarightdi 1 withacompletecrownandrootthatisnearly complete except for its chipped apex. From the cervicoena-mel junction to its superior termination, the root measures10.7 mm in length labially and 10.5 mm lingually. The root is broadmesiodistally (MD length maximum  ¼  4.8 mm) relative to its 󿬂 atness labiolingually (LL maximum breadth  ¼  3.7 mm). The rootwidens in MD diameter midway between the cervicoenamel junctionanditssuperiortermination.Thethick,shortrootistypicalof   P. robustus  central incisors. The incisal margin of the crownshows moderate wear and its labial and lingual faces have somescratches and developmental pitting. The crown possesses a largeinterproximalfacetmesially(CO ¼ 2.4mm;LL  ¼ 2.5mm)andatinyone distally (not measureable,  < 1 mm in each dimension).Lingually, the crown has a pronounced cingulum, which gradesocclusally along the lingual face into a prominent tuberculumextension. The mesial marginal ridge is partially obliterated by thelarge interproximal facet created by contact with the opposingcentral incisor. Upper central deciduous incisors are rare in the P. robustus  hypodigm, but SWT1/LB-15 conforms metrically andmorphologically to the other fairly complete and moderately worn  Table 1 New hominid fossils from Swartkrans Member 1.Specimen TaxonomicattributionElement DepositionalunitExcavation CoordinatesSWT1/HR-1  Paranthropusrobustus Right C HangingRemnantDump n/aSWT1/LB-2 cf.  Paranthropus  RightfemurLower Bank 1N 6E 4.4 mSWT1/LB-3  Paranthropusrobustus Leftdm 2  (?)Lower Bank 3N 6E 4.5 mSWT1/LB-4 Hominidae sp. indet.PostcaninetoothLower Bank 3N 6E 4.7 mSWT1/LB-5 cf.  Homo  Molar Lower Bank 3N 6E 3.4 mSWT1/LB-6 Hominidae sp. indet.UppermolarLower Bank 1N 8E 4.5 mSWT1/LB-8 Hominidae sp. indet.Left C Lower Bank 1N 6E 3.4 mSWT1/LB-10  Paranthropusrobustus Left M 1 Lower Bank 2N 5E 5.4 mSWT1/LB-11  Paranthropusrobustus Left uppermolarLower Bank 2N 6E 5.5 mSWT1/LB-12  Paranthropusrobustus Right P 3 Lower Bank 2N 7E 5.6 mSWT1/LB-13  Paranthropusrobustus Left I 1 Lower Bank 1N 7E 5.8 mSWT1/LB-14  Paranthropusrobustus Left P 4 Lower Bank 2N 7E 6.0 mSWT1/LB-15  Paranthropusrobustus Right di 1 Lower Bank 3N 7E 6.3 mSWT1/LB-16  Paranthropusrobustus Right I 1 Lower Bank 4N 6E 6.5 m  Table 2 (a) Mesiodistal (MD) diameters of newly recovered  Paranthropus robustus  fossilsfrom Swartkrans Member 1 and comparative statistics for previously analyzed P. robustus  teeth from Swartkrans. a (b) Labio/buccolingual (LL/BL) diameters of newly recovered  Paranthropus robustus  fossils from Swartkrans Member 1 andcomparative statistics for previously analyzed  P. robustus  teeth from Swartkrans. a (a) New specimens Published statisticsTooth Specimen MD n x S.D. S.E. C.V. % Rangedi 1 SWT1/LB-15 6.1 2 6.78 6.75 e 6.80I 1 SWT1/LB-13 8.9 11 9.04 0.61 0.18 6.75 8.20 e 9.80SWT1/LB-16 7.0SWT/TC-4 b 9.4C SWT1/HR-1 8.4 13 8.80 0.40 0.11 4.55 8.3 e 9.7P 3 SWT1/LB-12 9.2 23 10.06 0.64 0.13 6.36 9.3 e 12.2P 4 SWT1/LB-14 10.0 23 10.81 0.57 0.12 5.27 9.8 e 12.1UM SWT1/LB-11 c 13.4M 2 21 14.14 0.77 0.17 5.46 12.8 e 15.6M 3 21 14.89 0.98 0.21 6.58 12.9 e 16.7(b) New specimens Published statisticsTooth Specimen LL/BL n x S.D. S.E. C.V. % Rangedi 1 SWT1/LB-15 4.5 3 4.60 0.53 0.31 11.52  e I 1 SWT1/LB-13 7.2 12 7.24 0.55 0.16 7.60 6.3 e 8.1SWT1/LB-16 6.3SWT/TC-4 b 7.6C SWT1/HR-1 8.4 16 9.41 0.74 0.12 7.86 8.5 e 11.1P 4 SWT1/LB-14 (13.6) 21 15.11 0.86 0.19 5.69 13.5 e 16.3UM SWT1/LB-11 c 14.4M 2 19 16.07 0.80 0.18 4.98 14.4 e 17.2M 3 23 16.81 0.71 0.15 4.22 15.8 e 18.2 a All measurements arestandard(Robinson, 1956;Tobias,1967; Grine, 1984) and in mm. The value in parentheses is an estimate. Measurements for the new speci-mens described in this paper are not corrected for tooth wear. Published statisticsare from Grine (2004) for  P. robustus  teeth recovered from the Swartkrans Forma-tion by Broom and Robinson (1948 e 1949 and 1950 e 1953) and by Brain(1979 e 1986). b SWT/TC-4 is a recently discoveredand published (Sutton et al.,2009)  P. robustus fossil from the newly identi 󿬁 ed Talus Cone Deposit in the northeastern portion of Swartkrans. c The authors disagree whether SWT1/LB-11 is a M 2 or M 3 , thus publishedstatistics for Swartkrans  P. robustus  samples of both elements are provided in thelast two rows of parts a and b of this table. T.R. Pickering et al. / Journal of Human Evolution 62 (2012) 618 e 628 620  Swartkrans  P. robustus  di 1 , SKX 16060, in possessing a short, thickroot and squat crown bulging from it (Grine, 1989, 2004).SWT1/LB-13 is a complete crown, with an attached root frag-ment,ofaleft I 1 .Metricallyandmorphologicallythetoothissimilarto the previously described  P. robustus  homologues from Swartk-rans (Robinson, 1956; Grine, 1989, 2004). The lingual face of thetooth is slightly shoveled, possessing a well developed cingulumandmarginal ridges.Thetooth ’ sincisal marginismoderately worn,with scratches. Its enamel displays pitting labially and lingually,a large interproximal facet mesially (CO  ¼  3.8 mm; LL   ¼  3.1 mm)andasmalleronedistally(CO ¼ 3.1mm;LL2.7mm).SWT1/LB-13isprobably the antimere of SK 3, a complete right I 1 (initially iden-ti 󿬁 edasanI 2 byRobinson(1956)andre-identi 󿬁 edasanI 1 byClarke(1977b)) recovered in 1948 by Broom and Robinson from the HR breccia.Inviewofoverallmorphologicalandsizeresemblancesandbecause of closely matching, opposing interproximal facets, Clarke(1977b) previously concluded that the HR central maxillary incisorSK 40(Robinson,1956) isthe antimere ofSK 3 (SK3: MD ¼ 9.0 mm,LL  ¼ 7.1 mm, mesial facet CO ¼ 3.7 mm, mesial facet LL  ¼ 3.2 mm,distal facet CO  ¼  3.7 mm, distal facet LL   ¼  3.1 mm; SK 40:MD ¼ 9.0 mm, LL  ¼ 7.5 mm, mesial facet CO ¼ 3.9 mm, mesial facetLL   ¼  3.2 mm, distal facet CO  ¼  2.9 mm, distal facet estimatedLL   ¼  2.2 mm). However, our current comparative analysis revealsthat while the opposing interproximal facets of SK 3 and SK 40 do,indeed, match, there are slight differences in the morphologies of their lingual faces, whereas SWT1/LB-13 and SK 3 are nearly exactmatches based on the morphology, metrics and interproximal facetshape (Fig. 3). The inferred association of SWT1/LB-13 and SK 3implies one of two things. Perhaps the cranium to which bothincisors belonged was srcinally deposited in the overhangingsediments of the HR. At some point in the cranium ’ s diagenetichistory, SWT1/LB-13 was dislodged and eventually becamereworkedintothe underlyingsediments of the LB. Alternatively, SK3 might actually derive from the LB (as does SWT1/LB-13), ratherthan from the HR; SK 3 was recovered in the early days fromblasting operations at Swartkrans, and thus its provenience isimprecise.SWT1/LB-16isarightI 1 withacompletecrownandrootmissingonly its tip. The root is 5.8 mm thick mesiodistally and 6.0 mmlabiolingually. The root is 11.4 mm in length labiallyand 11.5 mm inlength lingually. The robust form of the root of SWT1/LB-16 istypical of  P. robustus  permanentupperincisors (Robinson,1956). Itsincisal margin is heavily worn, so much so that it rounds over ontothe labial and lingual crown faces. This extreme wear, with thecrown tapering from the incisal margin to the cervicoenamel junction, probably accounts for the fact that the specimen ’ s MDdiameter (uncorrected for wear) falls below the published I 1 MDdiameter range for Swartkrans  P. robustus . Both faces of the crownof SWT1/LB-16 are scratched. The labial face shows especiallyheavy antemortem damage, including large chips of missingenamel mesially and distally. The enamel displays pitting. Upper canines (Fig. 4) SWT1/LB-8 is a mesial enamel fragment of a left C, withmoderate apical wear. The fragment lacks a preserved interprox-imal facet. Meaningful measurements are not possible. The spec-imen is allocated to Hominidae gen. et sp. indet.SWT1/HR-1, a right C, has a complete crown and mesial portionof its root (lacking the tip). It is the only reasonablycomplete toothrecoveredsofarbytheSPRPfromBroomandRobinson ’ sHRbrecciadump. Typical of   P. robustus  upper canine teeth, the specimen has Figure 2.  Paranthropus robustus  upper central incisor fossils from the Lower Bank of Swartkrans Member 1. Each specimen is show in labial (left of each pair) and lingual (right of each pair) views; SWT1/LB-15 is a right deciduous (a), SWT1/LB-13 is a left permanent (b) and SWT1/LB-16 is a right permanent (c). Bar scale  ¼  1 cm. T.R. Pickering et al. / Journal of Human Evolution 62 (2012) 618 e 628  621  a clearly de 󿬁 ned cervical line with enamel that bulges thickly fromthe root in this area (Robinson, 1956). Like other  P. robustus  upperanterior teeth, SWT1/HR-1 possesses a shovel-shaped lingual face,with two marginal ridges converging onto its gingival eminence(Robinson,1956). Its labial surface has two small pits and there areatleastsixsmallpitsonitslingualsurface.Thereisalargeovalfaceton its mesial surface (CO  ¼  2.9 mm; LL   ¼  3.3 mm) and a narrowelongatedcrescentfacetthatrunsalongthecrown ’ sdistalmarginalridge (chord of CO  ¼  5.8 mm; LL   ¼  2.2 mm). The tooth ’ s apicalsurface is moderately worn. Upper third premolar (Fig. 5) SWT1/LB-12isarightP 3 crown,whichismissingitsbuccalridgebut is otherwise complete. The specimen is attributed to  P. robustus because of the roughly oval shape of its projected occlusal outlineand because it possesses a prominent cingulum bulge. Its mesial-distal groove is deeply incised, as is typical of Swartkrans P. robustus  P 3 s (Robinson, 1956). It possesses a moderately devel- oped interproximal facet distally (CO ¼ 2.9 mm; BL  ¼ 3.6 mm) andnone mesially. The occlusal surface of the tooth displays light wearscratches and pits, and except for the primary H-shaped  󿬁 ssurepattern (formed by the intersection of the mesial-distal groove and Figure 3.  The  Paranthropus robustus  right upper central incisor, SK 3, compared to thenewly recovered left upper central incisor SWT1/LB-13 (a) and to the previouslyrecovered left upper central incisor, SK 40 (b). All views are lingual and, in bothcomparisons, SK 3 is the specimen on the right of the paired teeth. All three teeth arequite comparable, but based on their closer morphological, metric and interproximalfacet shape similarities, we hypothesize that SK 3 is the antimere of SWT1/LB-13,instead of the antimere of SK 40, as previously suggested by Clarke (1977b). Barscale  ¼  1 cm. Figure 4.  Hominid  upper canine fossils from the Lower Bank (a) and Hanging Remnant(b) of Swartkrans Member 1. SWT1/LB-8, a mesial enamel fragment of a left perma-nent, is shown in mesial view (left of pair) and the right image of the specimen isa distal view of the tooth ’ s internal surface along a natural fracture plane (a). SWT1/HR-1, a right permanent, is shown in labial (left of pair) and lingual (right of pair)views (b). Bar scale  ¼  1 cm. T.R. Pickering et al. / Journal of Human Evolution 62 (2012) 618 e 628 622
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