gibert et al 1999 - humeral fragments attributable to horno sp. from lower pleistocene sites at...

8/6/2019 GIBERT Et Al 1999 - Humeral Fragments Attributable to Horno Sp. From Lower Pleistocene Sites at Venta Micena (Or

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Humeral fragments attributable to Horno sp. from LowerPleistocene sites at Venta Micena (Orce, Granada, Spain)

Josep GIBERT 1 , Asumpci M AL GOS A2 ,Florentina SNCHEZ 1 , Francesc RIBOT 1 ,ichael J. WALKER3'Institut Paleontolgic "Dr. M. Crusafont"C/ de 1'Escola Industrial, 23, 08201 Sabadell, Spain2 Unitat d'Antropologia, Departament de Biologia Animal, Biologia Vegetal i d'EcologiaFacultat de Ciencies (Edifici C)Universitat Autnoma de Barcelona08193 Bellaterra, Spain'rea de Antropologa Fsica, Departamento de Biologa Animal, Facultad de BiologaUniversidad de Murcia, 30100 Murcia SpainABSTRACT

Analyses are presented of long bone fragments attributable to the genus Horno, from the southeastern SpanishLower Pleistocene site of Venta Micena (Orce, Granada). Morphological and morphometrical analyses indicatesimilarity to both modern human and fossil hominid humeri, whereas they demonstrate significant contrasts withhumeri of carnivores and cercopithecoid monkeys. Early Lower Pleistocene hominid presence in southwesternEurope would represent a singularly important step in human evolution and a noteworthy contribution to thepresent controversy over whether or not hominids were leaving Africa at that time. The two fragments,VM-1960and VM-3691, were excavated in the same closed faunal assemblage (Table 1) and stratigraphical level in whichthere was found the cranial fragment VM-0 which has been assigned to Horno sp. on both morphological andpalaeoimmunological grounds.The two fragments are considered from a wide range of complementary methodologicalapproaches which separate humeral fragments of Horno from humeri of corresponding size, not only of moderncarnivore and cercopithecoid species, but also of those fossil carnivore and cercopitheoid taxa identified in thesoutheastern Spanish Lower Pleistocene to which they might otherwise have been assigned.

INTRODUCTION

In this paper the anatomical data have been worked out by Drs. J. Gibert, A. Malgosa, E Sanchez, E Ribotand M.Walker. The tomographies have been done in the UDIAT by Dr. Valls, following Dr. Gibert's indications.With those results Drs. E Snchez and A. Malgosa designed figures 3a, b, c wich were drawen by R. Torrico.The measures in tables 2, 3 were taken by Dr. J. Gibert. Figures 4a, b, c and 7 were made by Drs. Gibert andSnchez. The data for the morphometrical calculations were sent to Dr. P. Palmqvist and J. A. Prez-Claros byDr. Gibert within the Research Project Spanish Guvernemant (DGICYT): "Estudio Paleontolgico, Tafonmicoy Paleoantropolgico de los yacimientos del Pleistoceno Inferior de la regin de Orce y Cueva Victria" whereDr. Gibert was the main researcher.An extensive paper about the morphometrical data has been published in the abstracts of the Orce Conference(Palmqvist et al, 1995).The fauna in Table 1 has been taken from B. Martnez's doctoral thesis, directed by Dr. J. Gibert andmodified by Martnez in 1992.From 1982, sistematic excavation at Venta Micena (VM) ,have providet the following three potentiallyhominid fossils which, in order of discovery, are as follows:(a) VM-O, a cranial fragment of occipital squame and adjoining parietal fragments, found in 1982. It hasbeen the subject of both bioimmune assays performed both at the University of Granada and by Professor JeroldM. Lowenstein (n.d.) of the University of California at San Francisco, that indicate its human affinity (Borja etal. 1992, 1995, 1997, 1998; Garca-Olivares et al. 1989, Lowentein 1995, Zihlman et al 1996), as well as ofdetailed morphological studies (Campillo 1989; Gibert 1986, Gibert, Campillo et al. 1989, Gibert, Ribot,Ferrndez, Martnez, & Caporicci 1989, Gibert, Ribot, Ferrndez, Martnez & Ruz 1989, Gibert & Palmqvist1992, 1995, and cf. Campillo & Barcel 1989, Gibert et al. 1998) in contra to the opinion of Moy-Sol &Agust (1989), Moy & Klher 1997, Palmqvist 1997.

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(b ) VM-1960, a humeral fragment, found in 1988 (Gibert ct al. 1991, Gibert et al, 1989, Gibert et al., 1992)(Plate la, b),(c ) VM-3691, a humeral fragment, found in 1990, described here (Plate 1c, d).That VM-1960 and VM-3691 are, indeed, limb bones, rather than perhaps rib fragmenta of large mammals,

is clear from the proportions of their well-developed cortical bone and medullary canal, the demonstrableabsence of intercostal grooves, and the presence of clear-cut diaphyseal torsion inVM-1960 moreover, the sectionof the diaphysis extremes and the spongious tissue distribution into them it's very different between VM-1960and the greats felids ribs. Their morphological characteristics share more in common with humeri than withother long bones, as can be inferred from the detailed morphological descriptions given here.Other pointers to hominid presence at Venta Micena are stone manuports of dolomitic limestone and struckflakes of flint (Gibert et al. 1992), breakage patterns of long bones (Gibert ct al. 1992), and cut-marks on bones(Gibert & Jimnez 1989).

ANATOMICAL DESCRIPTIONS OF THE DIAPHYSESVM-1960 (Plate la, b, c, d). The specimen seems to be an immature, left humeral diaphysis. Proximally,

the medial and posterior parts of the anatomical neck are leen to be present. Distally, the upper part of thesupratrochlear crests are visible. That the bone belonged to a young child is suggested not only by its shaft-lengthof only 185 mm, but also by the way the radiologically visible medullary cavity is replaced superiorly bycancellous bone which fills the cross-sections at both ends of this short diaphysis, strongly suggesting metaphysealpresence close by.The diaphyseal surfaces are quite well preserved.The diaphysis is slightly bowed anteroposteriorlyand presents mediolateral flattening over its proximal half. A weak impression lies behind the anterior border,no doubt corresponding to the deltoid insertion. The medial surface of the shaft curves upwards towards asurgical neck which had undergone mediolateral compression probably caused by sedimentary pressure. Themedial and lateral lips of the bicipital groove are poorly marked and lie somwhat dorsally due to the aforementionedpost mortero deformation. A nutrient foramen is visible on the anterior surface of the medial border slightlybelow the midpoint of the shaft.Two more small foramina can be malle out on the upper surface of the diaphysis.In contrast to its proximal mediolateral flattening, the cross-section of the distal part of the shaft forms amediolaterally-elongated triangle. The distal half of the anterolateral surface shows damage, with loss of superficialbony tissue perhaps caused by scavenging animals. The shaft ends at a break just aboye the superior margin ofthe olecranon fossa. The medial border of the shaft begins to curve outwards towards the supratrochlear regionwhere the borders are well defined and clearly divergent.The medial and lateral surfaces twist in parallel togetherlaterally.VM-3691 (Plate le, f). This seems to be a distal fragment, 122 mm long, of a left humeral shaft. The distalbreak on the radial side occurred just aboye the superior margin of the olecranon fossa, lying 37 mm proximallyon the ulnar side. The shaft has three well-defined borders and surfaces. The borders are rounded, apart fromthe distal 60 mm of the mediolateral border which is sharper and shows impressions of muscular insertions.Viewed anteriorly, the distal part of the bone is broader mediolaterally than the proximal part, and has atriangular shape with an anterior border which twists laterally in a more pronounced fashion than do the otherborders. The posterior surface of the proximal part of the fragment is smaller than the anteromedial or anterolateralsurfaces both of which are similar in size. Viewed laterally, the bone seems to be tilted forwards with respect tothe longitudinal diaphyseal axis. The lateral part of the dorsal surface presents numerous grooves and some lossof superficial bony tissue; perhaps both owe to scavenging animals. At the proximal and distals breaks, a narrowmedullary cavity is visible, surrounded by a thick bony cortex that reaches maximal thickness of 9.7 mm at theanterior border proximally and 8.8 mm distally.

A N A L Y T IC A L M E T H O D O L O G YThe foregoing anatomical descriptions are incompatible with any other long bone than the humerus. Asalready mentioned, in order to distinguiste the correct large-mammal group to which the two humeri shouldbe assigned, it is necessary to bear in mind those carnivore and primate species, present in southeastern Spainduring the Lower Pleistocene, which had humeri of roughly comparable size and shape. The groups chosen forcomparison, therefore, are carnivores and large primates (cercopithecoid monkeys; Horno). Both modem andfossil forms have been considered and, where feasible, different age-categories have been used.

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Carnivores chosen for comparison wcre those with humeri comparable in size to human ones (Table 2),namely, Panthera pardus (male and female adults; adapted to woodland habitats), Acinonyx jubatus (adult male;adapted for running), Panthera leo, Felis lynx, Canis farniliaris, and an immature Ursus arctos. Cercopithecoidmonkey humeri were chosen from collections at the Zoological Gardens at Barcelona and Hamburg: Papiohamadryas (young and adult), Macaca sylvana (young and adults), and Mandrillus sphynx (male and female adults).We have directly studied fossil hominids of similar early Lower Pleistocene age are from Kenya (KNM-WT-15000, KNM-ER-1504, KNM-ER-1808), as well as modern human skeletons, particularly of children from theiron-age cemetery at Illot de Porros on Majorca (IP), the Mediaeval cemetery at Orce (CO), and the earlyChristian Roznan cemetery at Tarragona (T; N). Comparative published hominid data have also been considered,particularly from the Middle Pleistocene site of Atapuerca (near Ibeas de Juarros, in Burgos, Spain) and aprehistoric Holocene H. sapiens series from Seplveda (in Segovia, Spain).Wherever feasible the basis of comparison was sixfold, namely:

(1 ) anatomical comparison with humeri of known species;(2 ) comparison of diaphyseal cross-sectional descriptive morphology;(3 ) comparison of measurements and indices;(4 ) canonical discriminant function analysis of Fourier series harmonic descriptors of diaphyseal cross-sectional outlines;(5 ) comparison of the angle of humeral torsion;(6 ) comparison of ratios of cortical thickness to cross-sectional diameters.Further comments are in order concerning each of these comparisons in turn:(1 ) Anatomical comparison. This involved both direct observation on specimens from different species andpublished accounts and illustrations in standard works of reference (Pales & Lambert 1971; Pomba 1950; Sisson1947).(2 ) Comparison of diaphyseal cross-sectional descriptive morphology. Cross-sections were taken, both usinga "Microtecnica" comparator and by computer-assisted X-ray tomography, in both cases at 15 mm intervalsnumbered upwards from the apex of the olecranon fossa to the surgical neck. Humeral orientation was keptconstant when cross-sections were taken, by keeping horizontal that plane formed at right-angles to the longitudinalaxis of the shaft which bisects the apex of the olecranon fossa. Because VM-3691 is broken aboye the olecranonfossa, tomography commenced at the distal end, since divergence of the margins indicated their proximity tothe olecranon fossa. The number of cross-sections feasible varied with diaphyseal length which, in turn, vares

with biological age at death.(3 ) Comparison of measurements and indices. Quantitative comparisons are in order, given that the VM-1960 diaphysis is similar in length and cross-sectional dimensions, not only to immature human humeri, but alsoto humeral diaphyses found in some adult and immature carnivores and cercopithecoid monkeys (mammaliangroups represented in the southeastern Spanish Lower Pleistocene). This was done by comparing its humeralcross-sections with those of 32 humeral diaphyses selected with a view to taking into account the range ofvariation and sexual dimorphism in humeral diaphyseal shape in those taxa (Cercopithecoidea n = 10; Carnivoran = 6; Homo n = 16). The cross-sections were digitalized for reproduction by means of computerized axialtomography (CAT) at 15 mm intervals from the proximal margin of the olecranon fossa. Classical parameterswere considered, such as diaphyseal length and cross-sectional arcas and circumferences.The Diaphyseal Index (Olivier 1951) was calculated at 15 mm intervals from the cross-sectional values forcarnivores, cercopithecoid monkeys, Horno, and the Venta Micena humeri, and used for graphical displays as wellas for regression analysis.Values for cercopithecoid monkeys, Horno andVM-1960 were graphed for the diaphysealndex (y-axis) against values of the measured height of each cross-section aboye the epiphysis (x-axis). Polynomialregression analysis was undertaken of the diaphyseal ndex (y variable) on the true height aboye the epiphysis(x variable) and interpreted in terms of a 2nd-order polynomial regression equation of the kind

y = a + bx + cx (see Results, below).(4 ) Canonical discriminant function analysis of Fourier series harmonic descriptors of diaphyseal cross-sectional outlines. Following Davis (1986), the cross-sectional indices of circularity considered appropriate forthis study were Grain Shape Index (GSI), Circularity 2 and 3 (C2, C3), and Form Ratio (FR). Where A andP are area and circumference of the cross-section, and L the length between the two most widely separated pointson its circumference, thenGSI = P/LC2 = 4A/P2C3 = 4A/LPFR = A/L2

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The Radial Variation (SR2) was calculated from radial lengths from the centroid of the section to thecircumference, as follows:SR2 =R,_ r R.

i = 1 irlwhere Ri is that radial length which joins the centroid to the ith point of the n circumferential pointsdigitalized, and Rnt is the arithmetical mean of the radial lengths.Size-free multivariate morphometrical descriptors have been derived which overcome difficulties caused bypossible incommensurability of measurements and indices, particularly those which could be due to comparisonin (2) of non-homologous cross-sectional levels where diaphyses were of different length.The size-free descriptorsused here, on which between-taxa comparisons and contrasta were then made, characterize the cross-sectionalgeometry of proximal and distal diaphyseal regions in terms of successive harmonic amplitudes of the Fourierseries fitted to diaphyseal cross-sectional outlines.Different modalities of Fourier series analysis can be applied to outlines (whether open or closed) ofbiological shapes, ranging from polar radii and elliptic analysis (Rohlf, 1990), or eigenshape analysis (Lohmann,1983; Lohmann & Schweitzer, 1990), to methods to obtain median axes or line skeletons (Straney, 1990). Fourieranalysis of closed outlines is used here, following the polar radii approach (Ehrlich & Weinberg, 1970), beingone of the most widely used techniques in morphometrical analysis (Gonzlez & Palmqvist, 1990, and refs.).For closed outlines, the Fourier series is given by trigonometrical equations, involving sines and cosines,capable of describing accurately any two-dimensional figure, so long as any radii from its centre of gravityintercept, once only, the periphery of that figure. The shape-outline is determined from the following equationwhich defines the expansion of a radius, running from the centre of gravity of the outline to its periphery, asa function of the angle of rotation 0 in a system of polar coordinates whose origin is located in their centroid,with the radius (R,O) being given by:

R (0) = Ro (1 +cos (nO) +n sin (nO) ),an equation which is normally transformed to:RO = R (1 +cos (n0 - P) )n=1with C. = (A2 + 13; ) 1 andn arctan (B,/A),where O is the polar angle formed by the radius R(0) with a horizontal referente-line which crosses thecentre of mass for the outline, R o is the radius of a circumference whose arc is equivalent to that of the figureanalyzed, n is the harmonic order number, C n is the harmonic amplitude of the nth-order harmonic, and P isits phase angle. These equations can readily be fitted to circumferential outlines of diaphyses (note 1).The accuracy of Fourier analysis in characterizing an outline depends on both the number of coordinatesinitially taken to determine its periphery and the number of harmonics used in fitting the series to it. As a broadrule, there have to be taken at least twice as many peripheral points as the highest harmonic number to besought. The analysis allows the figure to be split into its geometrical components, regardless of its size or a needto have homologous points.Characterization of the outline can be as detailed as is desired; amplitudes of low-order harmonics willaccount for overall geometrical aspects of the outline, whereas higher ones take increasing account of fine-scaledsculpture. Take, for instante, a figure-of-eight: here, the amplitude of the second harmonic contributes to thataspect of the outline which is elongation, whereas the third harmonic shows how far it approaches a clover-leafshape and thereby measures the triangularity of the outline. By and large, the amplitude of the nth-orderharmonic measures how far a shape approaches an n-leaved clover-leaf. The amplitude of the first harmonic canbe regarded as measuring die error produced in fitting the Fourier series to an outline. Phase angles, dividedby their corresponding harmonic oder, locate the whereabouts in the figure of the influence of harmonics. Theseparameters may be employed as multivariate descriptors (Younkerhrlich, 1977).Any proposed system of shape-analysis will be regarded differently by its detractors and its defenders. Fourieranalysis of closed outlines presents some theoretical limitations and sonie possible practical drawbacks (Bookstein

et al. 1982; Ehrlich et al. 1983; Lohmann, 1983; Rohlf, 1990), namely:(a) Fourier series of polar radii are only useful in expanding single value functions;90


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HUMERAL FRAGMENTS ATTRIBUTABLE TO Homo SP. FROM LOWER PLEISTOCENE SITES ATVENTA

(b ) the centroid of the outline (to which all radii are referred) cannot be regarded as occupying a homologouspoint when comparing two specimens which have been analyzed; and(c ) whereas the shape of an outline can be completely represented by its adjusted Fourier series, in somecases it is not easy to tell how individual terms of the Fourier series (apart from lowest-order harmonics) reflect

different morphological features in the original shape: simple relationships between shape and harmonic functionappear clearly in such periodic or radially symmetrical forms as echinoderm exoskeletons, but less clearly in morecommonly aperiodic outlines.Figure 1 illustrates how the analysis is applied to a computer-assisted simulation of a humeral cross-sectionaloutlines, by incorporation of successive harmonics into the Fourier series fitted to the original outline. Thelowest harmonics clearly describe such geometrical componente of shape as elongation, triangularity, squareness,etc..., whereas reproduction of more localized details requires a greater number of terms in the harmonic series.Althoufh relatively large numbers of the outline, the lowest harmonics describe its major features sufficiently wellfor the second, third, ad fourth harmonics to be useful as morphometrical variables for carrying out discriminantanalysis of the cross-sectional outlines.The relative size of each cross-section was normalized with respect to all other cross-sections of the samebone as follows:Ar . I " ' A

1= 1

isA. i the arca of the j-th cross-section and m the number of cross-sections analyzed (1 = < j =


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VM1960:PINCIPAL FINDINGS(1 ) Anatomical comparison with humeri of known species. Notwithstanding its poorly developed anatomicalfeatures, there is broad similarity in length of VM-1960 to humeri of such medium-sized carnivores as Acinonyx,

Pantheraor Ursus (Table 2), although the maximum and minimum diaphyseal diameters ofVM-1960 are noticeablyless (11 and 14, see Table 2) even with respect to those of a young female Panthera, which was the most gracilespecimen considered.Whereas Acnonyx and Panthera humeral shafts are anteroposteriorly broad and radioulnarly narrow throughouttheir proximal two-thirds,VM-1960 only presents mediolateral compression in the most proximal cross-sections.The humeral diaphyseal morphology ofVM-1960 is most definitely not that of normal, medium-sized carnivores.Infantile and juvenile cercopithecoid humeri present very sharp, crest-like, lateral borders, unlike hurnansor VM-1960 (Figure 3a). Furthermore, the medial and lateral lips of the bicipital groove of both mature andimmature cercopithecoid monkeys are prominent and sharply edged. However, in children they are poorlydeveloped and rounded. Those of VM-1960 are barely perceptible and, therefore, resemble those of children.The border of the deltoid impression in Mandrllus, Macaca, and Papio is V-shaped with a strongly-markedrim, whereas in both children and VM-1960 the muscular impression is very faint, flat and rimless.In the three monkeys, the medial humeral border forms a smooth, continuous curve in mature andimmature individuals alike. However, in both children and VM-1960 the ulnar border is straight throughout themiddle third, coming to lie obliquely to it both distally and proximally, but never presenting a smooth, continuouscurve throughout its entire length. This difference is fundamental.VM-1960 has both a diaphyseal length and maximum and minimum mid-shaft diameters which are similarto those of 7 year-old children (cf. Table 2). The anteroposterior flattening of the distal third of VM-1960 andmediolateral narrowing of its proximal third are matched in children (Figure 3b).

(2 ) Comparison of diaphyseal cross-sectional descriptive morphology. Children and VM- 1960 do not havethe humeral cross-sectional form of carnivores, which is oval or triangular and has its maximum diameter lyinganteroposteriorly throughout the upper two-thirds of the shaft (Figure 3a, b). Unlike hurnans, carnivores oftenhave a supraepicondylar foramen and a sharp, crest-like, distal radial border to the shaft.There is a noteworthy difference in the proximal third of the humeral shaft between cross-sections takenon children and VM-1960 on the one hand, and those taken on mature or immature cercopithecoid monkeyson the other (Figure 3a, b, e). Whereas in the latter they tend to be more or less triangular with angular borders,with their principal cross-sectional axes lying more or less squarely to one other, in the former the upper thirdof the humeral shaft presents a more "compressed" radioulnar diameter and cross-sections are oval with roundedborders, their two principal cross-sectional axes (anteroposterior and radioulnar) lying obliquely to one another.The change in relationships of the two principal cross-sectional axes vis--vis the longitudinal diaphyseal axisreflects that humeral torsion which is a uniquely human skeletal feature.Over the middle third of the shaft, children present a humeral cross-section that ranges from pentagonal tocircular. That of a Roman child (specimen N-1) from the Tarragona cemetery (Figure 3b) mirrors well thesituation ni VM-1960.There is a significant difference in the distal third of the humeral shaft between cross-sections taken onchildren or VM-1960 on the one hand (Figure 3b), and those taken on mature or immature cercopithecoidmonkeys on the other (Figure 3a). In the former, cross-sections are roughly triangular, of comparable proportions,and their main axis is the radioulnar dimension which lies perpendicularly to the anteroposterior one, whereasin the latter they present an anteroposteriorly narrow radial border, not unlike that of tree-dwelling carnivoressuch as Panthera pardus (Figure 3a).(3 ) Comparison of measurements and indices. In carnivores, diaphyseal indices show values significantlydifferent from that ofVM-1960 (Table 3). Figure 4a highlights the similarity between the three carnivore specieschosen for comparison, especially as regards the proximal third of the humeral shaft. Their curves stand in markedcontrast to those for VM-1960 or human children, shown in Figure 4c. Curves for cercopithecoid monkeys differfrom both, as can be seen in Figure 4b. However, there are points of similarity between them and carnivoresas regards the more rounded cross-sections of the proximal third of the shaft (probably related to the quadrupedalismof both), which distinguish them from the narrower shafts of humans. The graph for VM-1960 differs in formfrom graphs for carnivores and cercopithecoid monkeys.The curves for children show a common pattern. Those for 7-to-8 year-old children closely resemble thegraph for VM-1960, (Figure 4c) although the greater length of VM-1960 may imply an older child. Its curveis very similar to that of the Tarragona Roman child N-1 mentioned aboye, although Holocene children havemore rounded humeral cross-sections.

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HUMERAL FRAGMENTS ATTRIBUTABLE TO Homo SP. FROM LOWER PI EISTOCENE SITES ATVENTA .

In general, low values, ranging from 50 to 75, are obtained for the diaphyseal index in proximal cross-sections of recent children, comparable to that forVM-1960 (Table 3), whereas adult and juvenile values are closeto 75. A value of 72, comparable to that of modern children, was obtained for the juvenile KNM-WT-15000(E. Mbua, personal communication) which dates from about 1.5 m.y.a.It is worth remarking that values for the three most proximal cross-sections are very similar in juvenile andadult modern humans, differing by between only 4 and 8 units. In both modern and hominid children, however,values for the same three cross-sections differ by from 12 to 19 units. The greatest diffcrence between them is16, in the case of VM-1960 (Table 3). To sum up, radioulnar "compression" of the most proximal 40 millimetresor so of the humeral shaft of humans produces a numerical deviation in the diaphyseal index with respect tovalues for comparable mammal species. Compression is normal in children's humeri, and is found in VM-1960which presents noteworthy values at its uppermost extremity. The values of the index from the correspondingregion of KNM-WT-15000 resemble more those of modern human or hominid children than they do those ofmodern juveniles or adults. That is consistent with the slight or nonexistent development both of the crests atthe lips of the bicipital groove and of the groove itself in KNM-WT-15000. Both KNM-WT-15000 and VM-1960 present anatomical characteristics seen in modero 6 or 7 year-old children.Presuming a second-order polynomial, of the kind y =a+bx+cx2 , regression analysis of the diaphyseal indices(y variable) on the measured height at which they were taken aho ye the distal epiphysis (x variable) gavedivergent values for Horno, cercopithecoid monkeys and VM-1960, as follows (where n is the total number ofsections per specimen multiplied by number of specimens):

Horno (n = 98)y = 59.127 + 0.631(s.e. = 0.077)x - 0.0038(s.e. = 0.00049)x2r = 0.647; F = 34.239; p


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tendency towards the value of unity is largely responsible for a non-homogeneous spatial distribution of thediscriminant variables with respect to the 3 pre-established mammalian groups.This is corroborated by low valuesfor Wilk's lambda statistic (which compares between-group variance with total variance) for both functions (bothtogether and separately), which, when transformed into values of chisquare, give a probability of p < 0.00001for the null hypothesis that there are no significant differences between the multivariate centroids of the threegroups.All the same, despite the significance of the value for Wilk's lambda, this does not necessarily imply thatthere is effective discrimination, because there could nevertheless be partial overlap between values of thefunctions for the various groups.That is why the usefulness of canonical functions for between-group discriminationhas to be demonstrated by checking whether their use results in correct reassignment of those individuals whichwere originally chosen to define the pre-established groups. Bayes' rule was used to determine the probabilitythat each individual belongs to its correct group, of the three under review, in terms of the values taken by thediscriminant functions. In this step, a probability of 1 was found that each of the individuals belonged to thatgroup which it had originally been chosen to establish; hence reclassification was 100% correct. How do fossildiaphyses fare? The procedure classified as belonging to Horno humeri with a probability of one, not only theKNM- WT-15000 humerus, but also the VM-1960 and VM-3691 diaphyseal fragments. Figure 6 shows the valuesof the discriminant functions for these examples and shows that they do not overlap with those for humeri ofcercopithecoid monkeys or carnivores; they lie within the ellipse of 95% confidence about the group mean.

(5 ) Comparison of the angle of diaphyseal torsion. The method of computer-assisted tomography used hereto determine the angle of humeral torsion gave mean values for carnivores of 91 within a range of 82-100,and for cercopithecoid monkeys of 93.2 within a range of 90-96.8. For hominids the mean was 135 withina range of 130-137: the value for VM-1960 of 128.5 stands far closer to the hominid range than to the rangesfor either of the other two groups. Froni the standpoint of those values obtained by traditional osteometry (Table5a: values calculated by Senut, 1983, from data in Knussmann, 1967) the new methodolo-gy used here gavecomparable results, although numerical values are consistently somewhat lower for all species measured, due tolack of the humeral head (Table 5b). Children's values are below adult human ones (the angle being inverselycorrelated with humeral length: Olivier, 1951) but, nonetheless, are far aboye those of adult monkeys. The valueobtained for VM-1960 corresponds to human children's values. By contrast, determinations based on computer-assisted tomographs consistently gave somewhat lower values. Interspecific consistency, however, corroborates thereliability of the new method.(6 ) Comparison of ratios of cortical thickness to cross-sectional diameters. Cortical thickness of VM-1960was measured at mid-shaft prior to refitting the diaphyseal fragments. The medullary canal was found to be verynarrow (Figure 7, 8). This is a well-knowm feature of fossil hominids, such as those from Atapuerca (Arsuagaet al., 1991), East Turkana, and elsewhere (Tattersall et al., 1988). It is also sometimes found in recent humanskeletons, such as the prehistoric Holocene ones from Seplveda (in Segovia, Spain). By contrast, at mid-shaftit is wide in both carnivores and cercopithecoid monkeys (Burr et al., 1989).When cortical thickness is represented as percentages of anteroposterior and mediolateral diaphyseal diameters(Kennedy, 1973), the results show that VM-1960 has greater cortical thickness than specimens taken for comparison(Figure 7).VM-3691 resemblesVM-1960 notwithstanding the slightly lesser anteroposterior (AP) and mediolateral(ML) cortical thicknesses of the latter, no doubt a reflection of immature biological age at death. KNM-ER.-1808 has lower values still, and even though its cortical bone thickness could only be measured at the brokenends of the shaft, they nevertheless exceed the corresponding values from Atapuerca or Seplveda and thosefound in carnivores or cercopithecoid monkeys. Values for the robusticity ndex of VM-1960 follow those forchildren, not carnivores or monkeys.

VM-3691: PRINCIPAL FINDINGSFor the fragrnent VM-3691, interspecific comparison is only possible for regions of the humeral shaft belowthe deltoid insertion, where it could be regarded as similar to humeri of cercopithecoid monkeys although theirlateral border is sharper in the distal third (Figure 3b, c). The cross-section of VM-3691 changes from triangularat its distal end to a mediolaterally compressed oval shape, with rounded borders, higher up. This morphologyis in stark contrast to carnivore humeral oval cross-sections which show angular borders of larger proportions,as well as to the rounded cercopithecoid cross-sections which become triangular proximally with sharp borders(Figure 3a):VM-3691 is quite unlike any of the adult carnivore and cercopithecoid humeri chosen for comparison.

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Mor-phology and cross-sections of human humeri, on the other hand, strongly resembie those of VM-3691.Values for the diaphyseal index show that VM-3691 follows the pattern of other fossil humeri of adult Horno(Table 3). Its thick cortex and indices of cortical thickness are comparable to those of VM-1960 (Figure 7).

CONCLUSION

The two fossil humeral fragments from the Venta Micena Lower Pleistocene site show significant contrastsnot only with respect to carnivore and cercopithecoid humeri from European Pleistocene deposits, but also withrespect to modern human and fossil hominid humeri.From comparative analysis, it is inferred that VM-1960 belonged to a hominid child, some differences frommodem children's humeri notwithstanding.Those differences included both proximal diaphyseal flattening (possiblyaugmented by post mortero taphonomical agents) and the slight nature of some anatomical features that are morepronounced in modem children's humeri of comparable size, which might hint at a difference in rates of growthbetween hominid and modern children.Although fewer comparative methods of analysis could be applied on the smaller humeral fragment ofVM-

3691, in its anatomical features it stands nearer to human humeri than to those of carnivores or cercopithecoidmonkeys, the only ones with which it can be compared given the particularities of morphology and biologicalage. It is proposed that that VM-1960 and VM-3691 have much more in common with humeri of fossil andmodern Horno than with those of other mammalian contenders. Together with the Horno sp. neurocranialfragment VM-0, not to mention the associated Venta Micena chopping-tool assemblage, these new finds fromthe sealed lithostratigraphical lacustrine deposit point unequivocally to hominid presence in southern Spainduring the Lower Pleistocene between 1.5 and 1.2 m.y.a. in a faunal context characterized by both African andEurasian mammals, long before the onset of the Middle Pleistocene at 0.73 m.y.a.. These singularly importantfindings demand reconsideration both of a much-repeated opinion that hominids did not reach Europe (or, forthat matter, South-East Asia) before 1 m.y.a., and of a conjectural Eurasian evolutionary trajectory of the genusHorno outside Africa which is usually couched in terms of mammalian-community ecology, palaeolithic responsesand hominid palaeontology, all in a fundamentally Middle, rather than Lower, Pleistocene context, and almostnever in a Eurasian early Lower Pleistocene context.ACKNOWLEDGEMENTS

We are obliged to Emma Mbua, Palaeoanthopolgy Conservator at the National Museum of Kenya, forkindly taking measurements on the fossil Horno specimens KNM-ER-1808 and KNM-WT-15000, in responseto our request to Dr. Meave Leakey.We also thank R.Torrico for assistance with the illustrations and tomographicalrepresentations.

NOTES

(') Using a digitalizer, a set (100-500) of Cartesian coordinates is taken around each circumference, thenumber of points depending on the degree of accuracy required by the subsequent analysis. The location ofcoordinates on the outline is arbitrary, although more points are needed where there are abrupt changes incurvature than where the curving outline is smooth. Coordinates are taken clockwise and consecutively. Comparisonof cross-sections from the standpoint of the information obtained in the harmonics requires only two equivalentpoints in order for all figures to be rotated to the same position. After calculating the (x,y) coordinates of Lpoints, the outline is closed by assuming = X, andY L ,1 =Y . The next step is to calculate the area enclosedby the outline using the following equation:(Yi + 1 +Y j )

1 2

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GIBERT, J., MALGOSA, A., SNCHEZ, F., RIBOT, E AND WALKEK,

Then, the first total moments about the X-axis and the Y-axis are estimated by:(Y ,2 ,--Y + Y 2)Xj -X+ 1 )J+1 j 3

6

MX-Ei= 12 2(Xj+1 +Y +1 XXj ) (Y j -Yj+1 )

6and the Cartesian coordinates of the cent oid are given by dividing the corresponding total moments by the areaof the outline:

MXYX-, -AEach of the initial L points taken along the periphery of the outline is then expressed in polar coordinates(R,O) about the centre of gravity as:R i = ( (Yi - Y.) 2 +X,) )l/2( Y - Y )e=arctan 3(X-X)3 cand bearing in mind, once more, thatRTH-1 LT- eThe next step is to calculate the mean radius of the figure (R .), by means of the following equation:

R =I4o (Rifi +R i )e-e6in which it is to be noted that, if O i+ ,e located in the first quadrant (O to 90) and O. in the fourth (270to 360), thenust be added to the difference in the angles.The radii of the points that define the periphery of the object are then divided by the mean radius, (R;= R/R), which renders the analysis independent of size, and the tercos A, and of the harmonic series arethen calculated by:) (cos -cos (neo ))R,--1'cos (nej +1) -os (ne j ) )2_, ( -n 1 = 1 (e -en2j+1_ j

1 - )R.' -R ) (sin (nej+1) -sin (ne.) (Rj ' +1 cos (ne ) -R 1cos (n e o))j +1j_11E= _E ( - 3 )an (e1 + 1 -0 )n2 n( 2 )The analytical stage involves establishing two matrices: [B] (of dimensions mxm, where m is the numberof variables) for the estimates of between-group variance of the variables, and [W] (of identical dimensions) fortheir within-group variance. The analysis provides that series of vectors [A] of linear coefficients which weightsthe variables such that they maximize the ratio of between-group variance to within-group variance, accordingto the function [A]T[B]/[A]T[W][A]. If the denominator is equal to 1, the ratio attains its maximal value when[A] contains the eigenvectors of the matrix [W]'[B], which are called canonical discriminant functions: theirability to discriminate between pre-established groups is in proportion to their associated eigenvalues and,

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generally speaking, the number of eigenvalues greater than zero is equal to the number of groups less one (ornumber of variables if this is less than the number of groups). By definition, eigenvectors are orthogonal, hencethe discriminant functions are uncorrelated with one other and, consequently, account for independent aspectsof the variante. A fundamental condition for calculating these functions is, of course,-that these specimens usedfor this purpose must already belong to groups which have been established by independent criteria. It is onlyin the second, classificatory, stage that specimens of doubtful attribution are added -in this case fossil diaphyses-in order to observe their effect on the values of the canonical discriminant functions: SCij = [A]T[X] i ,whereSC.. represents the value due to the j-th observation of the discriminante function [A] i , and [X]i is a vector withthe values which that observation imposes on the several variables. The Stepwise Discriminant Analysis programof SPSS 4.0 (Norusis, 1988) has been used on the VAX computer of Mlaga University's Central ComputingService.R E F E R E N C E SANADN P., JULIA R., DE DEKKER P., ROSSO J-C., & SOULI-MRSCHE I. 1987. Contribucin ala paleolimnologa del Pleistoceno inferior de la cuenca de Baza (sector Orce-Venta Micena). In S. Moy-Sol, J. Agust, J. Gibert & J.A. Vera, Eds, Paleontolgia i Evoluci. Memoria especial nm. 1. Geologa

y paleontologa del Pleistoceno inferior de Venta Micena, pp. 35-72, Sabadell: Institut Paleontolgic Dr. M.Crusafont, Diputaci de Barcelona.ARSUAGA J.L., CARRETERO J.M., MARTNEZ I., & GRACIA A. 1991. Cranial remains and long bonesfrom Atapuerca/Ibeas (Spain). Journal of Human Evolution 20, 3: 191-230.AllAROLI A. 1983. Quaternary mammals and the "end Villafranchian" dispersal event. A turning point in thehistory of Eurasia. Palaeogeography, Palaeoclimatology and Palaeoecology 47: 117-139.AllAROLI A., DE GIULI C., FICCARELLI C., & TORRE D. 1987. Late Pliocene to early mid-Pleistocenemammals in Eurasia: faunal succession and dispersal events. Palaeogeography, Palaeoclimatology and Palaeoecology66: 77-100.BOOKSTEIN EL., STRAUSS R.E., HUMPHRIES J.M., CHERNOFF B., ELDER R.L., & SMITH G.R.1982. A comment upon the uses of Fourier methods in systematics. Systematic zoology 31: 85-92.BORJA C., GARCA-PACHECO J.M., RAMREZ-LPEZ J.P. & GARCA-OLIVARES, E., 1992. Cuantificaciny caracterizacin de la albmina fsil del crneo de Orce. Proyecto Orce- Cueva Victoria (1988-1992):Presencia humana en el Pleistoceno inferior de Granada y Murcia. pp. 415-424. Ed. Ayuntamiento de Orce.BORJA C., GARCA OLIVARES E. 1995. Detection and characterization of proteinas in fossils from VentaMicena and Cueva Victoria by inmunonological methods.Orce Conference. Abstracts. Orce, 1995BORJA C., GARCA-PACHECO M., GARCA OLIVARES E., SCHEUENSTUHL G., LOWENSTEIN, J.1997. Immunoespecificity of Albumin Detected in 1.6 Million-Year-Old Fossils from Venta Micena inOrce, Granada, Spain. American Journal of Physical Anthropology, 103: 443-441.BORJA C. 1998. Deteccin y caracterizacin de proteinas fsiles mediante tcnicas inmunes: The hominids andtheir environment in the middle and lower Pleistocene of Europe and Asia. (In Gibert et al eds.) Museo dePrehistoria y Paleontologa "J.Gibert" (Orce, Granada).BROCA P. 1881. La torsion de 1' humrus et le tropomtre. Rdig par L. Manouvrier d'aprs les notes deBroca aprs sa mort. Revue Anthropologque de Paris 4: 193-210, 385-425, 577-592.BURR D.B., RUFF C.B., & JOHNSON C. 1989. Structural adaptations of the femur and humerus to arborealand terrestrial environments in three species of macaque. American Journal of Physical Anthropology 79 :357-367.CAMPILLO D. & BARCEL J.A. 1985. Estudio morfometrico de la cara interna de la escama del huesooccipital. Paleontolgia i Evoluci 19: 69-129.CAMPILLO D. 1989. Study of the Orce man. Estudio del hombre de Orce. In J. Gibert, D. Campillo, & E.Garca-Olivares, Eds, Los restos humanos de Orce y Cueva Victoria, pp.187-223, Sabadell: Institut PaleontolgicDr M. Crusafont, Diputaci de Barcelona.CERDEO E. 1993. Rhinocerotidae (Mammalia, Perissodactyla) en la cuenca de Guadix- Baza. In M.T.Alberdi & EP. Bonadonna, Eds, Geologa y paleontologa de la cuenca Guadix-Baza, pp. 273-288, Madrid:Museo Nacional de Ciencias Naturales, Consejo Superior de Investigaciones Cientficas.DAVIS J. 1986. Statistics and data analysis in geology, New York: John Wiley.EHRLICH R., Pharr B. & Healy-Williams N. 1983. Comments on the validity of Fourier descriptors insystematics: a reply to Bookstein et al. Systematic Zoology 32: 202-206.

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GIBERT J., FERRNDEZ C., MARTNEZ B., CAPORICCI R. & JIMNEZ, C. 1992. Roturas antrpicasen los huesos de Venta Micena y Olduvai. Estudio comparativo. In J. GIBERT, D. CAMPILLO, E.GARCA-OLIVARES, A. MALGOSA, F. MARTNEZ, B. MARTNEZ, M. WALKER, P. PALMQVIST,F. SNCHEZ & A. ARRIBAS, Eds, Proyecto Orce-Cueva Victoria (1988-1992). Presencia humana en elPleistoceno inferior de Granada y Murcia, pp. 283-305, Orce: Museo de Prehistoria Jos Gibert.GIBERT J., IGLESIAS A., MAILLO A. & GIBERT L. 1992. Industrias lticas en el Pleistoceno de la reginde Orce. InJ. GIBERT, D. CAMPILLO, E. GARCA-OLIVARES, A. MALGOSA, E MARTINEZ, B.MARTNEZ, M. WALKER, P. PALMQVIST, E SNCHEZ & A. ARRIBAS, Eds, Proyecto Orce-CuevaVictoria (1988-1992). Presencia humana en el Pleistoceno inferior de Granada y Murcia, pp. 219-281, Orce:Museo de Prehistoria Jos Gibert.GIBERT J. & PALMQVIST P. 1992. Dimensin fractal de las suturas del crneo de Orce. Revista Espaola dePaleontologa 7: 1-11.GIBERT J., SNCHEZ E, MALGOSA A., MARTNEZ B., M. WALKER Y RIBOT E 1992. Nuevos restoshumanos en los yacimientos de Orce y Cueva Victoria. in Proyecto Orce-Cueva Victoria 1988-1992: Presencia

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humana en el Pleistoceno inferior de Granada y Murcia. Ed. Museo de Prehistoria "Jos Gibert". Orce,Granada.GIBERT J. & PALMQVIST P 1995. Fractal analisys of the Orce sutures. Journal of Human Evolution, 28 :487-493.GIBERT J., CAMPILLO D., ARQUS J.M., GARCA OLIVARES E., BORJA C., LOWENSTEIN G. 1998.Hominid status of the Orce cranial fragment reasserted. Journal of Human Evolution, 34: 203- 217.GONZLEZ J.M. & PALMQVIST P 1990. Sobre la caracterizacin biomtrica del crecimiento y la forma delos foraminferos planctnicos. Aplicacin de las series de Fourier al anlisis de la forma de las cmaras.Revista Espaola de Paleontologa 5: 81-90.KENNEDY G. E. 1973. The anatomy of Middle and Lower Pleistocene hominid femora. London, University ofLondon, Ph.D. thesis.KNUSSMANN R. 1967. Humerus, Ulna und Radius der Simiae.Vergleichende morphologische Untersuchungenmit Bercksichtigung ber die Funktion. Bibliotheca primatologica 5: 1-339.LOHMANN G.P. 1983. Eigenshape analysis of microfossils: a general morphometric procedure for describingchanges in shape. Mathematical geology 15: 659-672.LOHMANN G.P. & SCHWEITZER. P.N. 1990. On eigenshape analysis. In EJ. Rohlf & EL. Bookstein, Eds,

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TABLE 1Venta Micena Fauna.This revised minimal faunal list is based on Martnez (1991, 1992) whose

Megantereon sp. has subsequently been assigned to M. whitei (Palmqvist &Martnez, n.d.)Horno sp.Cans falconer MAJORCanis etruscus MAJORVulpes praeglacialis KORMOSHornotherium latidens OWENMegantereon whitei BROOMLynx sp.Pachycrocuta brevirostris AYMARDUrsus etruscus CUVIERcf. Meles sp.Hippopotamus amphibius antiquus DESMARESTPraeovibos sp.Soergelia mnor MOYA-SOLABubalus sp .Capra alba MOY-SOLPraemegaceros sollhacus ROBERTCervidae gen. et sp. indet.Dcerorhinus etruscus brachycephalus SCHROEDER(syn. Stephanorhinus etruscus, thus Cerdeo, 1993)Equus stenonis granatensis ALBERDI & RUIZ-BUSTOSMammuthus merdionalis NESTIProlagus calpensis MAJOROryctolagus cf. lacosti POMELDesmana sp .Allaphaomys pliocaenicus KORMOSCastillomys crusafonti subsp.Apodemus aff. mystacinus DANFORD & ALSTONEliomys intermedius FRIANTHystrix major GERVAISTestudo sp .Lacerta sp .Ophdia indet.Rana sp .Charadriiforme indet. (aff. Laridae)

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TABLE 2:Diaphyseal measurements (in mm) of humeri of Horno and other species taken between similar

anatomical landmarks. Modern human specimens are from the Mediaeval cemetery at Orce,Granada (CO) and iron-age burials at Illot de Porros, Menorca (IP); fossil specimens are

(VM-1960 and VM-3691) from Venta Micena (Orce, Granada)SPECIMEN Length

(mm)Max. diam.

(mm)MM. diana.

(mm)Ursus arctos (adult) 19 0 44 33Panthera leo (infant) 85 28 21Canis familiaris (adult) 126 23 15Acinonyx jubattts (adult) 16 5 24 16Panthera pardos (adult male) 16 5 30 19Panthera pardos (adult female) 11 5 20 15Papio hamadryas (adult) 12 8 20 12Papio harnadryas (young) 12 5 16 12Mandrillus sphynx (adult male) 18 0 22 17Mandrillus sphynx (adult female) 143 17 11CO-4 (adult) 310 23 18CO-6 (adult) 270 25 19CO-1 (infant) 10 0 11 9CO-2 (infant) 110 13 11IP C-32A (infant) 100 11 9IP (unnumbered) (infant) 80 10 8IP-2 (infant) 9 5 10 8IP-19 (infant) 15 5 14 10IP C32B (infant) 160 12 11IP SW-2 (infant) 200 18 13VM-1960 185 14 11VM-3691 - 20 14

3 4 5

Plate 1: 1, VM-1960. Humeral fragment from Venta Micena (Orce, Granada, Spain), anterior view; 2: VM-3691.Humeral fragment from Venta Micena (Orce, Granada, Spain) anterior view. Scale in centimetres.

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TABLE 3:Humeral diaphyseal indices at different cross-sectional levels. F= fossil specimens: WT= KNM-WT-15000, 1960= VM-1960. Ch=

children, J= juvenile modern humans, adult modern humans: C-32= IP C-32 (iron-age burials at Illot de Porros, Menorca), CO-1, CO-2, CO-3, CO-5, CO-6= (Mediaeval cemetery at Orce, Granada), T-30, T-32, T-35, T-93= Roncan cemetery at Tarragona, N-1,N-2= Roman cemetery at Tarragona, MZ= cemetery at Mequinenza, Zaragoza, M-1= Cercopithecoid monkeys: 68.2= adult maleMandrillus sphynx, 108.1= young Papio hantadryas, 6974= , 35.3= Macaca sylvana, 1690= adult Papio hamadryas, 68.1= adult female

Mandrillus sphynx. Carnivores: 23.10, 131.5, 109.1, CANIS= Canis familiaris, LYNX = Lynx (Felis) lynx.

I T Ch Ch Ch Ch Ch

1 1 1 1 1 4 1 N 0 I D 8

Ch Chh Ch J J 3 A A A o CSRCOPITIMCOID XONUYS e4 casetrycetes c./)oNN ? 1960 C- 3 2 C O 1 00 13 5 T32 130 5-1 M-2 M 2 CO3 9-1 19 3 C0 4 CO3 C06 68.2 108.1 6974 35.3 1690 68.1 23.10 131.5 109.1 c2512 LYNX 11172 33 7 1 51 6 3 72 00 73 64 78 4 0 16 90 6 0 91 07 17 95 7 6 75 70 73 75 se 62 G O 73 6 612 64 01 73 73 12 17 el. 77 45 4 6 94 90 5 4 78 17 es 93 75 75 9 4 00 10 72 67 56 73 78 u9111793

71 .75131 1 51313

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11II77

099393

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92941 7345419

71146 6

537575127661

160065

7 17371777377

9 41717756675

4176es

7071606974eo

SI6551647074

717012o01

5577 4310 MI100 7547 927 3 5741 33 1545 5555 9649 0079 5411 0076 03839390 7175 1077 11el es1 1 1 6369 $177 15e0 0265 8349 6065 4774 4200 es90 9zzz12 75 95 4 4 59 59 80 91 69 1 1 3 05 78 17 90 43 47 54 33 73 89 43 60 72 1453 66 II 71 51 79 94 39 89 70 69 17 76 44 37 44 53 69 87 O 4990 46 15 55 62 73 69 35 69 51 35 ti 63 45 73 8056 48 52 39 44 94 87 3 3 42 13 32 5 6 7367 45 79 76 70 39 53 5 389 71 31 1 5380 52 44 1057 4033

O9u

uo


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HUMERAL FRAGMENTS ATTRIBUTABLE TO HOMO SP. FROM LOWER PLEISTOCENE SITES ATVENTA

TABLE 4Standardized canonical coefficients.for discriminant functions 1 and 2 Correlations of variates with canonicaldiscriminant functions 1 and 2Function 1 Function 2 Function 1 Function 2

GSI p 0.7531 -1.5045 -0.1832 -0.4006C1p -7.1643 1.2175 0.0704 0.3859C2 p 5.6241 -2.6971 0.0418 0.5024FRp -0.3849 2.2037 -0.3043 0.6214SR2 p -0.4402 -0.3764 0.2118 -0.6169H2 p 0.5679 1.8039 -0.1203 -0.6740H3 p 1.0767 0.3959 0.8932 0.2473H4p 0.3732 0.8200 0.6185 -0.0050GSI d 0.3124 -0.2636 -0.4835 -0.2479Ci d 3.8898 1.0693 0.4643 0.3022C2, -1.4236 -0.3772 0.4497 0.3225FRd -0.3627 0.8108 -0.1699 0.2905H3 d -0.5423 0.9414 -0.1074 0.6005Areap/Aread 0.2815 -0.8337 0,5120 -0,6224

(Table 4a): GSI = Grain Shape Index; C1 and C2= Circularity Indices 1 and 2; FR= Form Ratio; SR2= radialvariance (Davis 1986). H2, H3 and H4= harmonic amplitudes of order 2, 3 and 4. Area p /Areaa = mean proximalarea divided by mean distal area. Subscript p = proximal. Subscript d = distal.

Function 1 Function 2Eigenvalus" 10,57 5,42% of explained variance 66,12 33,88canonical correlations 0,956 0,919Wilk's lambda 0,0135* 0,1559chi-square test 96,915* 41,82degrees of freedom 28* 13significante level 0,00001* 0,0001

(Table 4b): Parameters associated with canonical discriminant functions (* = Function 1 and 2 jointly)TABLE 5: Angle of diaphyseal torsion.

Genus Sample size Mean RangeHorno (adult) 15 0 145.1 134.0-156.2Macaca 10 2 98.8 92.1-105.5Papio 10 96.0 82.2-109.8Ccrcocebus 8 99.1 92.6-105.6Cercopithecus 16 97.0 88.5-105.5

Table 5a: after Senut 1983, who based calculations on data froni Knussmann 1967.10 3


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GIBERT, J., MALGOSA, A., SNCHEZ, E, RIBOT, E AND WALKER, M.J.

Specimen Valu eChild MZ-124-2 128Fossil VM-1960 131Child MZ-87 137Child IP-C-32 139Child MZ (unnumbered) 139Child IP-C-19 140

Table 5b: values for angle of torsion obtained by measurements taken here from computer-assisted tomographs.MZ= cemetery at Mequinenza (Zaragoza); IP= iron-age cemetery at Illot de Porros (Menorca).

o

Figure 1: Computer-assisted simulationsof a humeral cross-section obtained fromFourier series fitted to the outline.Numbers indicate the harmonic orderused in each simulation. Amplitudevalues of the second, third and fourthharmonics are shown, accompanied bythe corresponding simulations obtainedusing only these harmonics.econd h.arm clichird harmonicp litu de 0.2073 Amp litu de 0.1061 Fnurth harrrtonieAmplitude 0.0801

104


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1

e


Figure 2a: Computer-assisted tomographicallysuperimposed images of proximal and distal cross-sections of humeral shafts, from which the angle ofhumeral torsion was determined. MM= medial margin.ML= lateral margin. CT= margin between greatertuberosity and deltoid tuberosity of the proximal cross-section; the other cross-section is in the supratrochlearregion at the apex of the olecranon fossa: the angleof humeral torsion lies between the two mediolateraldimensions.

Figure 2b: Measurements of cortical thickness taken oncross-sections of humeral shaft (after Kennedy, 1973).a= anterior cortical thickness. b = posterior corticalthickness. c= anteroposterior dimension. d= lateralthickness. e= medial thickness. f= mediolateral dimension.

A BC FGHZ

PP .

' r

-

G e %% ti5C es

J Q Z G6 5 a c ,tha l'9 e e z c ) ,z0 0 D a o . D

C c3 cpc (1 ) 0

0 c

Figura 3a: Comparison of humeraldiaphyseal cross-sections of carnivores,cercopithecoid monkeys, and VentaMicena fossils. Scale in centimetres.A= adult Acinonyx juba tus. B= adultmale Panthera pardus. C= adult maleMandrillus sphynx. D= Macaca sylvana.E= young Papio hamadryas. F= Macacasylvana. G= VM-1960 fossil fromVenta Micena. H= VM-3691 fossilfrom Venta Micena.

10 5


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GIBER.T, J., MALGOSA, A., SNCHEZ, E, RIBOT, E AND WALKER,

A 3 C D F G H Io o o o z ) % 4 1 ,

1

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o c o . o o z) z ) too c i ) o O i, soQ e o e c , o OooooDe CD00(C)n5 C O (3 CZN e0O G 1 acz C*G o z ,n c po o z , c _ - , ga c _ ) 9z b t ; , , , j

Ali

Figure 3b: Comparison between humeral cross-sections of infant Horno sapiens humeri (A, B, C, D, E and F),VM-1960 (G), and of an adult Horno sapiens humerus (H) and VM-3691 (I).

10 6


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PR 0.X.

DI ST.

10 cm

HUMERAL FRAGMENTS ATTRII3UTABLE TO HOMO SP. FROM LOWER PLEISTOCENE SITES AT VENTA

A CD Enoo'oO %o SO 9

0 %O1 c lOO O C 9 )D oiL C1'_ '

Figure 3c: Comparison of humeral cross-sections from Venta Micena with fossil hominid humeri. A: KNM-ER-1504, B: KNM-ER-1808, C: KNM-WT-15000, D: VM-3691 and E: VM-1960.10 7


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10 090807060 -

401-30

AB'

VM-1960 35-3 - 19-- 68-2 66-1 K. 106-1 1693

10 09 0 - 1807 0 -6050

GIBER.T, j., MALGOSA,ANCHEZ, F, RIBOT, F. AND WALKER, M.J.100

1 1 - /0 '70n ,igure 4: Comparison of values for thediaphyseal index (y-axis) at cross-sections 15mm apart (x-axis: 1= most distal, at apex ofolecranon fossa; number of cross-sections isdetermined by length of shaft and biologicalage).VM-1960= Venta Micena fossil diaphysis.Figure 4a: the carnivores are as follows: 109-1= adult Acinonyx jubatus, 23-10= adult male

Panthera pardus, Canis= adult Canes familiares.

60504 030

A

VIVI-1960 Cariis - 109-1 23-10 I

Figure 4b: the cercopithecoid monkeys are asfollows: 68-2= adult male Mandrillus spliynx,35-3= Macaca sylvana, 1690= adult Papiohamadryas, 106-1=Papiohamadryas, 68-1=Macaca sylvana.

Figure 4c: the Horno specimens are as follows:CO-2= child from Mediaeval cemetery at Orce,Granada, N-1 and N-2-= children from Romancemetery at Tarragona, IP C-32= iron-age childfrom Illot de Porros, Menorca.4 030

V M - 196 0 -- N -1 N -2' 3 -P-32 C O - 2

10 8


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CERCOPI THECOI DMONKEYS


\14".Yr .*4 VM-1960100 _

Figure 5: Comparison of fossil VM-1960 with values of the diaphyseal ndex(y-axis) in cercopithecoid monkeys(Figure 5a) and Homo (Figure 5b), takenat successive heights (x-axis) aboye theapex of the olecranon fossa. Brokenlines represent the 95% confidencelimits.

100 _

80 -60 -

40 -

20 -3 P

HOMO

0 610 90 12 0 1 15080 (mm) ....

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-s Seconddhscrimjnnt function(33..1O31,i, nl varinnre

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Fi rst d isdiminant fu ndion --I-(66,12% ni vaaiaore r-planccli- -2 CERCOP

O fromo C,r,orilh5cid:A earnivercs*Grnup centrnida

w-m Gona/ A

A

CARNIVORES

Figure 6: Brivariate plot of the first two canonical discriminant functions obtained from the humeri of carnivores,cercopithecoid monkeys, and fossil specimens KNM-WT-15000 andVM-1960.Dotted unes represent the p


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30 4011 3

1 3

% 10 201

AP

MLx

50 60 70 80 90AT- 93

2j4 5 i 6AT- 217 AT- 25

1808M VM1960 CV 1 VM 3691

4 6 2 5y yAT 217 AT -25

VM1960 ** CV1VM 3691

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Figure 7: Comparisons of cortical thickness in fossil specimens and living species (from Arsuaga et al. 1991 modified). Carnivores: 1 =ynx (Pchs) lynx, 2=Panthera pardus. Cercopithecoidea: 3= Mandrillus sphynx, 4 and 5 = Macaca, 6= Papio hamadryas. Middle Pleistocene Homo: AT- 25, AT-93 and AT-217=Atapuerca, Burgos. Lower Pleistocene Horno: KNM-ER-1808, CV-1= Cueva Victoria, Murcia, VM-1960 and VM-3691 = Venta Micena. Solid bar= prehistoricHolocene Horno sapiens from Seplveda, Segovia. AP= anteroposterior index. ML= mediolateral index.


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HUMERAL. EFRAGMENTS ATTRIBUTABLE TO Homo SP. FROM LOWER PLEISTOCENE SITES ATVENTA

a

d

e

VFigure 8: Comparison of medullary cavity-size in mid-shaft humeral cross-sections. a: CV-2 fossil humeros fromCueva Victoria; b: VM-1960 fossil humeros from Venta Micena, c: VM-3691 fossil humerus from Venta Micena,d: Horno sapiens humerus; e: Mandrillus sphinx humerus; f: Papio hamadras humerus; g: Hornotherium latdens fossilhumerus from Venta Micena, and h: male Panthera pardus humerus. (x 1.25)

gibert et al 1999 - humeral fragments attributable to horno sp. from lower pleistocene sites at...

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