Humeral fragments attributable to Horno sp.

from Lower
Pleistocene sites at Venta Micena (Orce, Granada, Spain)

Josep GIBERT 1 , Asumpció MALGOSA2,

Florentina SÁNCHEZ 1 , Francesc RIBOT 1 , Michael J. WALKER3
'Institut Paleontológic "Dr. M. Crusafont"
C/ de 1'Escola Industrial, 23, 08201 Sabadell, Spain
2Unitat d'Antropologia, Departament de Biologia Animal, Biologia Vegetal i d'Ecologia
Facultat de Ciencies (Edifici C)
Universitat Autónoma de Barcelona
08193 Bellaterra, Spain
'Área de Antropología Física, Departamento de Biología Animal, Facultad de Biología
Universidad de Murcia, 30100 Murcia Spain

ABSTRACT

Analyses are presented of long bone fragments attributable to the genus Horno, from the southeastern Spanish
Lower Pleistocene site of Venta Micena (Orce, Granada). Morphological and morphometrical analyses indicate
similarity to both modern human and fossil hominid humeri, whereas they demonstrate significant contrasts with
humeri of carnivores and cercopithecoid monkeys. Early Lower Pleistocene hominid presence in southwestern
Europe would represent a singularly important step in human evolution and a noteworthy contribution to the
present controversy over whether or not hominids were leaving Africa at that time. The two fragments,VM-1960
and VM-3691, were excavated in the same closed faunal assemblage (Table 1) and stratigraphical level in which
there was found the cranial fragment VM-0 which has been assigned to Horno sp. on both morphological and
palaeoimmunological grounds.The two fragments are considered from a wide range of complementary methodological
approaches which separate humeral fragments of Horno from humeri of corresponding size, not only of modern
carnivore and cercopithecoid species, but also of those fossil carnivore and cercopitheoid taxa identified in the
southeastern 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 Ribot
and M.Walker. The tomographies have been done in the UDIAT by Dr. Valls, following Dr. Gibert's indications.
With those results Drs. E Sánchez 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 and
Sánchez. The data for the morphometrical calculations were sent to Dr. P. Palmqvist and J. A. Pérez-Claros by
Dr. Gibert within the Research Project Spanish Guvernemant (DGICYT): "Estudio Paleontológico, Tafonómico
y Paleoantropológico de los yacimientos del Pleistoceno Inferior de la región de Orce y Cueva Victória" where
Dr. 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. Martínez's doctoral thesis, directed by Dr. J. Gibert and
modified by Martínez in 1992.
From 1982, sistematic excavation at Venta Micena (VM) ,have providet the following three potentially
hominid 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 has
been the subject of both bioimmune assays performed both at the University of Granada and by Professor Jerold
M. Lowenstein (n.d.) of the University of California at San Francisco, that indicate its human affinity (Borja et
al. 1992, 1995, 1997, 1998; García-Olivares et al. 1989, Lowentein 1995, Zihlman et al 1996), as well as of
detailed morphological studies (Campillo 1989; Gibert 1986, Gibert, Campillo et al. 1989, Gibert, Ribot,
Ferrández, Martínez, & Caporicci 1989, Gibert, Ribot, Ferrández, Martínez & Ruz 1989, Gibert & Palmqvist
1992, 1995, and cf. Campillo & Barceló 1989, Gibert et al. 1998) in contra to the opinion of Moyá-Soló &
Agustí (1989), Moyá & Kólher 1997, Palmqvist 1997.

87
 GIBERT, J., MALGOSA, A., SÁNCHEZ, E, RI130T, AND WALKER, M.J.

(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 demonstrable
absence of intercostal grooves, and the presence of clear-cut diaphyseal torsion inVM-1960 moreover, the section
of the diaphysis extremes and the spongious tissue distribution into them it's very different between VM-1960
and the greats felids ribs. Their morphological characteristics share more in common with humeri than with
other 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 struck
flakes of flint (Gibert et al. 1992), breakage patterns of long bones (Gibert ct al. 1992), and cut-marks on bones
(Gibert & Jiménez 1989).

ANATOMICAL DESCRIPTIONS OF THE DIAPHYSES

VM-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 the
supratrochlear crests are visible. That the bone belonged to a young child is suggested not only by its shaft-length
of only 185 mm, but also by the way the radiologically visible medullary cavity is replaced superiorly by
cancellous bone which fills the cross-sections at both ends of this short diaphysis, strongly suggesting metaphyseal
presence close by.The diaphyseal surfaces are quite well preserved.The diaphysis is slightly bowed anteroposteriorly
and 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 a
surgical neck which had undergone mediolateral compression probably caused by sedimentary pressure. The
medial and lateral lips of the bicipital groove are poorly marked and lie somwhat dorsally due to the aforementioned
post mortero deformation. A nutrient foramen is visible on the anterior surface of the medial border slightly
below 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 a
mediolaterally-elongated triangle. The distal half of the anterolateral surface shows damage, with loss of superficial
bony tissue perhaps caused by scavenging animals. The shaft ends at a break just aboye the superior margin of
the olecranon fossa. The medial border of the shaft begins to curve outwards towards the supratrochlear region
where the borders are well defined and clearly divergent.The medial and lateral surfaces twist in parallel together
laterally.
VM-3691 (Plate le, f). This seems to be a distal fragment, 122 mm long, of a left humeral shaft. The distal
break on the radial side occurred just aboye the superior margin of the olecranon fossa, lying 37 mm proximally
on the ulnar side. The shaft has three well-defined borders and surfaces. The borders are rounded, apart from
the 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 a
triangular shape with an anterior border which twists laterally in a more pronounced fashion than do the other
borders. The posterior surface of the proximal part of the fragment is smaller than the anteromedial or anterolateral
surfaces both of which are similar in size. Viewed laterally, the bone seems to be tilted forwards with respect to
the longitudinal diaphyseal axis. The lateral part of the dorsal surface presents numerous grooves and some loss
of superficial bony tissue; perhaps both owe to scavenging animals. At the proximal and distals breaks, a narrow
medullary cavity is visible, surrounded by a thick bony cortex that reaches maximal thickness of 9.7 mm at the
anterior border proximally and 8.8 mm distally.

ANALYTICAL METHODOLOGY

The foregoing anatomical descriptions are incompatible with any other long bone than the humerus. As
already mentioned, in order to distinguiste the correct large-mammal group to which the two humeri should
be assigned, it is necessary to bear in mind those carnivore and primate species, present in southeastern Spain
during the Lower Pleistocene, which had humeri of roughly comparable size and shape. The groups chosen for
comparison, therefore, are carnivores and large primates (cercopithecoid monkeys; Horno). Both modem and
fossil forms have been considered and, where feasible, different age-categories have been used.

88
 HUMERAL FRAGMENTS ATTRIBUTABLE TO HOMO SP. FROM POWER PLEISTOCENE SITES ATVENTA

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. Cercopithecoid
monkey humeri were chosen from collections at the Zoological Gardens at Barcelona and Hamburg: Papio
hamadryas (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 the
iron-age cemetery at Illot de Porros on Majorca (IP), the Mediaeval cemetery at Orce (CO), and the early
Christian 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 a
prehistoric Holocene H. sapiens series from Sepúlveda (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 and
published accounts and illustrations in standard works of reference (Pales & Lambert 1971; Pomba 1950; Sisson
1947).
(2) Comparison of diaphyseal cross-sectional descriptive morphology. Cross-sections were taken, both using
a "Microtecnica" comparator and by computer-assisted X-ray tomography, in both cases at 15 mm intervals
numbered upwards from the apex of the olecranon fossa to the surgical neck. Humeral orientation was kept
constant when cross-sections were taken, by keeping horizontal that plane formed at right-angles to the longitudinal
axis of the shaft which bisects the apex of the olecranon fossa. Because VM-3691 is broken aboye the olecranon
fossa, tomography commenced at the distal end, since divergence of the margins indicated their proximity to
the olecranon fossa. The number of cross-sections feasible varied with diaphyseal length which, in turn, varíes
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 also
to humeral diaphyses found in some adult and immature carnivores and cercopithecoid monkeys (mammalian
groups represented in the southeastern Spanish Lower Pleistocene). This was done by comparing its humeral
cross-sections with those of 32 humeral diaphyses selected with a view to taking into account the range of
variation and sexual dimorphism in humeral diaphyseal shape in those taxa (Cercopithecoidea n = 10; Carnivora
n = 6; Homo n = 16). The cross-sections were digitalized for reproduction by means of computerized axial
tomography (CAT) at 15 mm intervals from the proximal margin of the olecranon fossa. Classical parameters
were 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 for
carnivores, cercopithecoid monkeys, Horno, and the Venta Micena humeri, and used for graphical displays as well
as for regression analysis.Values for cercopithecoid monkeys, Horno andVM-1960 were graphed for the diaphyseal
índex (y-axis) against values of the measured height of each cross-section aboye the epiphysis (x-axis). Polynomial
regression analysis was undertaken of the diaphyseal índex (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 + cx2 (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 for
this study were Grain Shape Index (GSI), Circularity 2 and 3 (C2, C3), and Form Ratio (FR). Where A and
P are area and circumference of the cross-section, and L the length between the two most widely separated points
on its circumference, then
GSI = P/L
C2 = 4A/P2
C3 = 4A/LP
FR = A/L2

89
 GIBERT, J., MALcíosA, A., SÁNCHEZ, E, RIBOT, E AND WAI KER,

The Radial Variation (SR2) was calculated from radial lengths from the centroid of the section to the
circumference, as follows:

SR2 = ( R,_ r R.
i=1 irl
where Ri is that radial length which joins the centroid to the ith point of the n circumferential points
digitalized, and Rnt is the arithmetical mean of the radial lengths.
Size-free multivariate morphometrical descriptors have been derived which overcome difficulties caused by
possible incommensurability of measurements and indices, particularly those which could be due to comparison
in (2) of non-homologous cross-sectional levels where diaphyses were of different length.The size-free descriptors
used here, on which between-taxa comparisons and contrasta were then made, characterize the cross-sectional
geometry of proximal and distal diaphyseal regions in terms of successive harmonic amplitudes of the Fourier
series fitted to diaphyseal cross-sectional outlines.
Different modalities of Fourier series analysis can be applied to outlines (whether open or closed) of
biological 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). Fourier
analysis of closed outlines is used here, following the polar radii approach (Ehrlich & Weinberg, 1970), being
one of the most widely used techniques in morphometrical analysis (González & 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 gravity
intercept, once only, the periphery of that figure. The shape-outline is determined from the following equation
which defines the expansion of a radius, running from the centre of gravity of the outline to its periphery, as
a 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 + A cos (nO) + B n sin (nO) ),

an equation which is normally transformed to:

RO = Ro (1 + C cos (n0 - P) )
n=1

with C . = (A.2 + 13;2) 1 and Pn arctan (B,/A„),

where O is the polar angle formed by the radius R(0) with a horizontal referente-line which crosses the
centre of mass for the outline, Ro is the radius of a circumference whose arc is equivalent to that of the figure
analyzed, n is the harmonic order number, C n is the harmonic amplitude of the nth-order harmonic, and P is
its 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 coordinates
initially taken to determine its periphery and the number of harmonics used in fitting the series to it. As a broad
rule, there have to be taken at least twice as many peripheral points as the highest harmonic number to be
sought. The analysis allows the figure to be split into its geometrical components, regardless of its size or a need
to have homologous points.
Characterization of the outline can be as detailed as is desired; amplitudes of low-order harmonics will
account for overall geometrical aspects of the outline, whereas higher ones take increasing account of fine-scaled
sculpture. Take, for instante, a figure-of-eight: here, the amplitude of the second harmonic contributes to that
aspect of the outline which is elongation, whereas the third harmonic shows how far it approaches a clover-leaf
shape and thereby measures the triangularity of the outline. By and large, the amplitude of the nth-order
harmonic measures how far a shape approaches an n-leaved clover-leaf. The amplitude of the first harmonic can
be regarded as measuring die error produced in fitting the Fourier series to an outline. Phase angles, divided
by their corresponding harmonic oder, locate the whereabouts in the figure of the influence of harmonics. These
parameters may be employed as multivariate descriptors (Younker Ehrlich, 1977).
Any proposed system of shape-analysis will be regarded differently by its detractors and its defenders. Fourier
analysis 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
 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 homologous
point 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 some
cases 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 function
appear clearly in such periodic or radially symmetrical forms as echinoderm exoskeletons, but less clearly in more
commonly aperiodic outlines.
Figure 1 illustrates how the analysis is applied to a computer-assisted simulation of a humeral cross-sectional
outlines, by incorporation of successive harmonics into the Fourier series fitted to the original outline. The
lowest 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 well
for the second, third, ad fourth harmonics to be useful as morphometrical variables for carrying out discriminant
analysis of the cross-sectional outlines.
The relative size of each cross-section was normalized with respect to all other cross-sections of the same
bone as follows:

I"' A
Ar. —

1= 1

A. iisthe arca of the j-th cross-section and m the number of cross-sections analyzed (1 = < j = <
ni). In order to overcome incommensurability between diaphyseal cross-sections of any one humerus with those
of any other, arithmetical mean were calculated for each variable for the proximal and distal regions of each
humeral shaft, thereby ensuring definition of the variables for comparable anatomical regions and enablig
comparison between different humeri. The regions in question were described in terms of presence of absence
of markings for muscle insertions. The extrernities of the bones and apophyses were, of course, excluded froni
consideration. Because markings of muscular insertions are feebly developed in children before about the age of
9 years old, their comparison has been based on taking the rnost proximal 3-5 cross-sections of the hurneralshaft
as being equivalent, broadly speaking, to those of the proximal part of the diaphysis in carmivores and cercopithecoid
m o nkeys
Moreover, because short shafts, on which fewer tomographs could be taken than on long ones, led to a
distorted view of the relative size of their cross-sections, correction was 1-nade for the differing influence of the
available number of cross-sections on variables which reflect relative cross-sectional size. This was done by
normalizing them in terms of the ratio between mean relative size of distal to proximal cross-sections; this is far
preferable to simply taking mean proximal or distal values, because it affords an equivalent variable for all
specimens of all taxa.
A matrix was then generated, with values of the variables forming columns corresponding to the morphometrical
descriptors of proximal and distal diaphyseal regions, respectively, and with each humeral specimen occupying
a separate row. Discriminant analysis using canonical variables was then performed between the taxa (cf. R.eyment
et al., 1984) in a two-step approach by which analysis leads on to classification (note 2).
(5) Comparison of the angle of diaphyseal torsion. For determining the angle of diaphyseal torsion, the
traditional osteometrical method (Broca, 1981; Olivier, 1951; Martins, 1968) is unreliable where the humeral
head is missing. A different, new approach has therefore been devised for the diaphyseal fossil fragments, namely,
computer-assisted X-ray tomographical projection of the most proximal diaphyseal cross-section on a distal one
near the apex of the olecranon fossa (Figure 2a). This enables measurement of that angle which is subtended
between the distal diaphyseal diameter, taken across the medial and lateral borders, and the greatest proximal
diaphyseal diameter, taken from the lateral lip of the bicipital groove.Values so determined are slightly lower than
those obtained using classical osteometry (Knussmann, 1967) which was also carried out here, on the longer fossil
diaphyseal fragments, by projecting the major axis of the proximal extremity of the shaft onto a line parallel to
the distal dorsal surface.
(6) Comparison of ratios of cortical thickness to cross-sectional diameters. Absolute and relative cortical
thicknesses were determined froni six cross-sectional measurements, namely, anterior, posterior, medial, and
lateral cortical thicknesses, and anteroposterior and mediolateral diaphyseal diameters (cf. Kennedy, 1973) (Figure
2b). Mid-shaft cross-sections are highlight important differences in size of the medullary canal; it is useful to
compare those percentages of anteroposterior and mediolateral diaphyseal diameters which correspond to cortical
bone thickness (ibid.).

91
 GIBERT, J., MALGOSA, A., SÁNCHEZ, E, RIBOT, F. ANO WALKER, M.J.

VM-1960: PINCIPAL FINDINGS

(1) Anatomical comparison with humeri of known species. Notwithstanding its poorly developed anatomical
features, there is broad similarity in length of VM-1960 to humeri of such medium-sized carnivores as Acinonyx,
Panthera or Ursus (Table 2), although the maximum and minimum diaphyseal diameters ofVM-1960 are noticeably
less (11 and 14, see Table 2) even with respect to those of a young female Panthera, which was the most gracile
specimen considered.
Whereas Acínonyx and Panthera humeral shafts are anteroposteriorly broad and radioulnarly narrow throughout
their 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 hurnans
or VM-1960 (Figure 3a). Furthermore, the medial and lateral lips of the bicipital groove of both mature and
immature cercopithecoid monkeys are prominent and sharply edged. However, in children they are poorly
developed and rounded. Those of VM-1960 are barely perceptible and, therefore, resemble those of children.
The border of the deltoid impression in Mandríllus, Macaca, and Papio is V-shaped with a strongly-marked
rim, 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 and
immature individuals alike. However, in both children and VM-1960 the ulnar border is straight throughout the
middle third, coming to lie obliquely to it both distally and proximally, but never presenting a smooth, continuous
curve throughout its entire length. This difference is fundamental.
VM-1960 has both a diaphyseal length and maximum and minimum mid-shaft diameters which are similar
to those of 7 year-old children (cf. Table 2). The anteroposterior flattening of the distal third of VM-1960 and
mediolateral 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 have
the humeral cross-sectional form of carnivores, which is oval or triangular and has its maximum diameter lying
anteroposteriorly throughout the upper two-thirds of the shaft (Figure 3a, b). Unlike hurnans, carnivores often
have 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 taken
on children and VM-1960 on the one hand, and those taken on mature or immature cercopithecoid monkeys
on 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 third
of the humeral shaft presents a more "compressed" radioulnar diameter and cross-sections are oval with rounded
borders, 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-á-vis the longitudinal diaphyseal axis
reflects 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 to
circular. That of a Roman child (specimen N-1) from the Tarragona cemetery (Figure 3b) mirrors well the
situation ni VM-1960.
There is a significant difference in the distal third of the humeral shaft between cross-sections taken on
children or VM-1960 on the one hand (Figure 3b), and those taken on mature or immature cercopithecoid
monkeys 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, whereas
in the latter they present an anteroposteriorly narrow radial border, not unlike that of tree-dwelling carnivores
such as Panthera pardus (Figure 3a).
(3) Comparison of measurements and indices. In carnivores, diaphyseal indices show values significantly
different from that ofVM-1960 (Table 3). Figure 4a highlights the similarity between the three carnivore species
chosen for comparison, especially as regards the proximal third of the humeral shaft. Their curves stand in marked
contrast to those for VM-1960 or human children, shown in Figure 4c. Curves for cercopithecoid monkeys differ
from both, as can be seen in Figure 4b. However, there are points of similarity between them and carnivores
as regards the more rounded cross-sections of the proximal third of the shaft (probably related to the quadrupedalism
of both), which distinguish them from the narrower shafts of humans. The graph for VM-1960 differs in form
from graphs for carnivores and cercopithecoid monkeys.
The curves for children show a common pattern. Those for 7-to-8 year-old children closely resemble the
graph for VM-1960, (Figure 4c) although the greater length of VM-1960 may imply an older child. Its curve
is very similar to that of the Tarragona Roman child N-1 mentioned aboye, although Holocene children have
more rounded humeral cross-sections.

92
 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 close
to 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 and
adult 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 is
16, in the case of VM-1960 (Table 3). To sum up, radioulnar "compression" of the most proximal 40 millimetres
or so of the humeral shaft of humans produces a numerical deviation in the diaphyseal index with respect to
values for comparable mammal species. Compression is normal in children's humeri, and is found in VM-1960
which presents noteworthy values at its uppermost extremity. The values of the index from the corresponding
region of KNM-WT-15000 resemble more those of modern human or hominid children than they do those of
modern juveniles or adults. That is consistent with the slight or nonexistent development both of the crests at
the 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) gave
divergent values for Horno, cercopithecoid monkeys and VM-1960, as follows (where n is the total number of
sections 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)x2
r = 0.647; F = 34.239; p <0.001;
cercopithecoid monkeys (n = 81)
y = 52.608 + 0.613(s.e. = ±0.077)x - 0.0029(s.e.= ±0.00055)x2
r = 0.762; F = 54.123; p <0.001;
VM-1960 (n = 11)
y = 51.126 + 0.795(s.e. = ± 0.07)x - 0.0052(s.e. = ±0.00045)x2
r = 0.971; F = 66.878; p <0.001.

As can be seen from those values, the term c (i.e. the slope of x2 , responsible for inflexion of the curve
after its initial upswing) took on lower values in hominids (-0.0038) andVM-1960 (-0.0052) than in cercopithecoid
monkeys (-0.0029). The interval between the 95% confidente limits (Figure 5) for that parameter of the equation
in VM-1960 (from -0.00621 to -0.00414) partly overlaps that in Homo (from -0.00477 to -0.00281), but not
that of the monkeys (from -0.00403 to 0.00183). These data suggest hominid assignation for VM-1960.
(4) Canonical discriminant function analysis of Fourier series harmonic descriptors of diaphyseal cross-
sectional outlines. Table 4 shows the mean values for the morphomeffical variables determined on the distal and
proximal cross-sections, respectively, of each diaphysis analyzed. Canonical discriminant functions were extracted
from the data of the modero humeri in Table 4, after excluding the variables of radial variance and first -and
second- order harmonic amplitudes which failed to show significant differences between the groups under
review.
The importante of a particular variable in discriminating between groups cannot be evaluated directly from
the coefficients standardized on the discriminant vectors because the values of coefficients show intercorrelation
and depend on values of other variables included in the function. Nevertheless, analyzing together the standardized
coefficients for the variables and their correlations with the discriminant functions (Table 4a, b ) suggests that
the most relevant variables for discriminating between Horno and cercopithecoid monkeys (first discriminant
function) are, on the one hand, those which are correlated with the triangularity and squareness of proximal
diaphyseal cross-sections and with their relative size with respect to distal ores (variables which give higher values
for cercopithecoid monkeys) and, on the other, those which are correlated with distal diaphyseal circularity
(variables which highlight the greater circularity in carnivores and humans than in the monkeys) (Figure 6). The
second discriminant function separates carnivores from humans and cercopithecoid monkeys and is correlated
with elongation and triangularity of distal cross-sections, being greater in the human and cercopithecoid diaphyses
considered than in those of carnivores (Figure 6).
Table 4 shows some of the parameters associated with the func-tions. The canonical correlations describe
the proportion of the total vari-ance accounted for by the between-group differences for each function, and their

93
 GIBERT, J., MALGOSA, A., SÁNCHEZ, E. RIBOT, E AND WALKER, M.J.

tendency towards the value of unity is largely responsible for a non-homogeneous spatial distribution of the
discriminant variables with respect to the 3 pre-established mammalian groups.This is corroborated by low values
for Wilk's lambda statistic (which compares between-group variance with total variance) for both functions (both
together and separately), which, when transformed into values of chisquare, give a probability of p < 0.00001
for the null hypothesis that there are no significant differences between the multivariate centroids of the three
groups.
All the same, despite the significance of the value for Wilk's lambda, this does not necessarily imply that
there is effective discrimination, because there could nevertheless be partial overlap between values of the
functions for the various groups.That is why the usefulness of canonical functions for between-group discrimination
has to be demonstrated by checking whether their use results in correct reassignment of those individuals which
were originally chosen to define the pre-established groups. Bayes' rule was used to determine the probability
that each individual belongs to its correct group, of the three under review, in terms of the values taken by the
discriminant functions. In this step, a probability of 1 was found that each of the individuals belonged to that
group which it had originally been chosen to establish; hence reclassification was 100% correct. How do fossil
diaphyses fare? The procedure classified as belonging to Horno humeri with a probability of one, not only the
KNM- WT-15000 humerus, but also the VM-1960 and VM-3691 diaphyseal fragments. Figure 6 shows the values
of the discriminant functions for these examples and shows that they do not overlap with those for humeri of
cercopithecoid 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 here
to 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° within
a range of 130°-137°: the value for VM-1960 of 128.5° stands far closer to the hominid range than to the ranges
for either of the other two groups. Froni the standpoint of those values obtained by traditional osteometry (Table
5a: values calculated by Senut, 1983, from data in Knussmann, 1967) the new methodolo-gy used here gave
comparable results, although numerical values are consistently somewhat lower for all species measured, due to
lack of the humeral head (Table 5b). Children's values are below adult human ones (the angle being inversely
correlated with humeral length: Olivier, 1951) but, nonetheless, are far aboye those of adult monkeys. The value
obtained 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 the
reliability of the new method.
(6) Comparison of ratios of cortical thickness to cross-sectional diameters. Cortical thickness of VM-1960
was measured at mid-shaft prior to refitting the diaphyseal fragments. The medullary canal was found to be very
narrow (Figure 7, 8). This is a well-knowm feature of fossil hominids, such as those from Atapuerca (Arsuaga
et al., 1991), East Turkana, and elsewhere (Tattersall et al., 1988). It is also sometimes found in recent human
skeletons, such as the prehistoric Holocene ones from Sepúlveda (in Segovia, Spain). By contrast, at mid-shaft
it 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 broken
ends of the shaft, they nevertheless exceed the corresponding values from Atapuerca or Sepúlveda and those
found in carnivores or cercopithecoid monkeys. Values for the robusticity índex of VM-1960 follow those for
children, not carnivores or monkeys.

VM-3691: PRINCIPAL FINDINGS

For the fragrnent VM-3691, interspecific comparison is only possible for regions of the humeral shaft below
the deltoid insertion, where it could be regarded as similar to humeri of cercopithecoid monkeys although their
lateral border is sharper in the distal third (Figure 3b, c). The cross-section of VM-3691 changes from triangular
at its distal end to a mediolaterally compressed oval shape, with rounded borders, higher up. This morphology
is 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.

94
 HUMERAL FRAGMENTS ATTRII3UTABLE TO Homo SP. FROM LOWER PLEISTOC:ENE SITES ATVENTA

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 contrasts
not only with respect to carnivore and cercopithecoid humeri from European Pleistocene deposits, but also with
respect to modern human and fossil hominid humeri.
From comparative analysis, it is inferred that VM-1960 belonged to a hominid child, some differences from
modem children's humeri notwithstanding.Those differences included both proximal diaphyseal flattening (possibly
augmented by post mortero taphonomical agents) and the slight nature of some anatomical features that are more
pronounced in modem children's humeri of comparable size, which might hint at a difference in rates of growth
between 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 cercopithecoid
monkeys, the only ones with which it can be compared given the particularities of morphology and biological
age.
It is proposed that that VM-1960 and VM-3691 have much more in common with humeri of fossil and
modern Horno than with those of other mammalian contenders. Together with the Horno sp. neurocranial
fragment VM-0, not to mention the associated Venta Micena chopping-tool assemblage, these new finds from
the sealed lithostratigraphical lacustrine deposit point unequivocally to hominid presence in southern Spain
during the Lower Pleistocene between 1.5 and 1.2 m.y.a. in a faunal context characterized by both African and
Eurasian mammals, long before the onset of the Middle Pleistocene at 0.73 m.y.a.. These singularly important
findings demand reconsideration both of a much-repeated opinion that hominids did not reach Europe (or, for
that matter, South-East Asia) before 1 m.y.a., and of a conjectural Eurasian evolutionary trajectory of the genus
Horno outside Africa which is usually couched in terms of mammalian-community ecology, palaeolithic responses
and hominid palaeontology, all in a fundamentally Middle, rather than Lower, Pleistocene context, and almost
never in a Eurasian early Lower Pleistocene context.

ACKNOWLEDGEMENTS

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

NOTES

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

(Yi +1 +Y j )

1 2

95


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

Then, the first total moments about the X-axis and the Y-axis are estimated by:

2
(Y ,2 ,-1-YJ+1 j + Y ) ( X
j -X i +1 )
3
6

2 2
(X j +1 +Y +1 X +X j ) (Y - Yj )
j +1
MX-E
i=1 6

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

MX MY
X,- Y, -
A A

Each 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:

Ri = ( (Yi - Y c.) 2 + - X,) )l/2

(Y-Y)
e =arctan 3
(X -X )
3 c

and bearing in mind, once more, that

RTH-1 °LIT - el

The next step is to calculate the mean radius of the figure (R . ), by means of the following equation:

II
(R i fi +R i ) ( e ,-e
4
Ro=
6
in which it is to be noted that, if Oi+ , be located in the first quadrant (O° to 90°) and O. in the fourth (270°
to 360°), then must 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;
= Ri /R 0 ), which renders the analysis independent of size, and the tercos A, and of the harmonic series are
then calculated by:

) (cos -cos (ne o )) (R,--1'cos (nej +1) - cos (ne j ) )

—2_, ( -
n 1 =1 (e j+1_ -ej )n2

(R.' - ) (sin (nej+1) -sin (ne.) )

j +1 -Rj - (R j' +1 cos (ne ) -R 1 cos (n e o))
Ea =_E
1
1 (
ni
_11
n
3 )

(e 1+1 -0 )n2

(2) The analytical stage involves establishing two matrices: [B] (of dimensions mxm, where m is the number
of variables) for the estimates of between-group variance of the variables, and [W] (of identical dimensions) for
their within-group variance. The analysis provides that series of vectors [A] of linear coefficients which weights
the variables such that they maximize the ratio of between-group variance to within-group variance, according
to 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: their
ability to discriminate between pre-established groups is in proportion to their associated eigenvalues and,

96
 HUMERAL FRAGMENTS ATTRIBUTABLE TO Homo SP. FROM LOWER PLEISTOCENE SITES ATVENTA

generally speaking, the number of eigenvalues greater than zero is equal to the number of groups less one (or
number of variables if this is less than the number of groups). By definition, eigenvectors are orthogonal, hence
the discriminant functions are uncorrelated with one other and, consequently, account for independent aspects
of the variante. A fundamental condition for calculating these functions is, of course,-that these specimens used
for this purpose must already belong to groups which have been established by independent criteria. It is only
in 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 , where
SC.. represents the value due to the j-th observation of the discriminante function [A] i , and [X]i is a vector with
the values which that observation imposes on the several variables. The Stepwise Discriminant Analysis program
of SPSS 4.0 (Norusis, 1988) has been used on the VAX computer of Málaga University's Central Computing
Service.

REFERENCES

ANADÓN P., JULIA R., DE DEKKER P., ROSSO J-C., & SOULIÉ-MÁRSCHE I. 1987. Contribución a
la paleolimnología del Pleistoceno inferior de la cuenca de Baza (sector Orce-Venta Micena). In S. Moyá-
Solá, J. Agustí, J. Gibert & J.A. Vera, Eds, Paleontológia i Evolució. Memoria especial núm. 1. Geología
y paleontología del Pleistoceno inferior de Venta Micena, pp. 35-72, Sabadell: Institut Paleontológic Dr. M.
Crusafont, Diputació de Barcelona.
ARSUAGA J.L., CARRETERO J.M., MARTÍNEZ I., & GRACIA A. 1991. Cranial remains and long bones
from 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 the
history of Eurasia. Palaeogeography, Palaeoclimatology and Palaeoecology 47: 117-139.
AllAROLI A., DE GIULI C., FICCARELLI C., & TORRE D. 1987. Late Pliocene to early mid-Pleistocene
mammals in Eurasia: faunal succession and dispersal events. Palaeogeography, Palaeoclimatology and Palaeoecology
66: 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., GARCÍA-PACHECO J.M., RAMÍREZ-LÓPEZ J.P. & GARCÍA-OLIVARES, E., 1992. Cuantificación
y caracterización de la albúmina fósil del cráneo 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., GARCÍA OLIVARES E. 1995. Detection and characterization of proteinas in fossils from Venta
Micena and Cueva Victoria by inmunonological methods.Orce Conference. Abstracts. Orce, 1995
BORJA C., GARCÍA-PACHECO M., GARCÍA OLIVARES E., SCHEUENSTUHL G., LOWENSTEIN, J.
1997. Immunoespecificity of Albumin Detected in 1.6 Million-Year-Old Fossils from Venta Micena in
Orce, Granada, Spain. American Journal of Physical Anthropology, 103: 443-441.
BORJA C. 1998. Detección y caracterización de proteinas fósiles mediante técnicas inmunes: The hominids and
their environment in the middle and lower Pleistocene of Europe and Asia. (In Gibert et al eds.) Museo de
Prehistoria y Paleontología "J.Gibert" (Orce, Granada).
BROCA P. 1881. La torsion de 1' humérus et le tropométre. Rédigé par L. Manouvrier d'aprés les notes de
Broca aprés sa mort. Revue Anthropologíque 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 arboreal
and terrestrial environments in three species of macaque. American Journal of Physical Anthropology 79:
357-367.
CAMPILLO D. & BARCELÓ J.A. 1985. Estudio morfometrico de la cara interna de la escama del hueso
occipital. Paleontológia i Evolució 19: 69-129.
CAMPILLO D. 1989. Study of the Orce man. Estudio del hombre de Orce. In J. Gibert, D. Campillo, & E.
García-Olivares, Eds, Los restos humanos de Orce y Cueva Victoria, pp.187-223, Sabadell: Institut Paleontológic
Dr M. Crusafont, Diputació de Barcelona.
CERDEÑO E. 1993. Rhinocerotidae (Mammalia, Perissodactyla) en la cuenca de Guadix- Baza. In M.T.
Alberdi & EP. Bonadonna, Eds, Geología y paleontología de la cuenca Guadix-Baza, pp. 273-288, Madrid:
Museo Nacional de Ciencias Naturales, Consejo Superior de Investigaciones Científicas.
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 in
systematics: a reply to Bookstein et al. Systematic Zoology 32: 202-206.

97
 GIBERT, J., MALGOSA, A., SÁNCHEZ, E, RIBOT, E AND WALKER,

GARCÍA-OLIVARES E. GALLARDO J.M., MARTÍNEZ E, BORJA C. & GARCÍA-OLIVARES D. 1989.

Detección y caracterización de proteínas fósiles en el cráneo de Orce (resultados preliminares). In J-.-
Gibert, D. Campillo & E. García-Olivares, Eds, Los restos humanos de Orce y Cueva Victoria, pp. 225-228,
Sabadell: Institut Paleontológic Dr M. Crusafont, Diputació de Barcelona.
GIBERT J. & PONS-MOYA J. 1984. Estudio morfológico de la falange del género Horno de Cueva Victoria
(Cartagena, Murcia). Paleontología i Evolució 18: 49-55.
GIBERT J., PONS-MOYA J. & RUZ C. 1985. Comparación métrica y morfológica de la falange del género
Homo de Cueva Victoria (Cartagena, Murcia) con las de primates y úrsidos. Paleontología i Evolució 19:
147-154.
GIBERT J. 1986. El yacimiento de Venta Micena (Orce, Granada): su importancia, acción antropológica y
caracteristicas paleoantropológicas del fragmento de cráneo de Horno sp. In O. Arteaga, Ed, Homenaje a
Luis Siret (1934 a 1984). Actas del Congreso: Homenaje a Luis Siret, Cuevas de Almanzora, Junio 1984, pp.
37-49. Madrid: Dirección General de Bellas Artes de la Consejería de Cultura de la Junta de Andalucía.
GIBERT J., CAMPILLO D., CAPORICCI R., RIBOT F., FERRÁNDEZ C. & MARTÍNEZ B. 1989.
Anatomical study: comparison of the hominid cranial fragment froni Venta Micena (Orce, Spain) with
fossil and extant mammals. Human Evolution 4 (4): 283-305.
GIBERT J., PONS-MOYA J. & RUZ, C. 1989. Estudio del resto humano encontrado en el yacimiento cárstico
del Pleistoceno inferior de Cueva Victoria (Cartagena, Murcia). In J. Gibert, D. Campillo & E. García-
Olivares, Eds, Los restos humanos de Orce y Cueva Victoria, pp. 395-405, Sabadell: Institut Paleontológic Dr.
M. Crusafont, Diputació de Barcelona.
GIBERT J., RIBOT E, FERRÁNDEZ C., MARTÍNEZ B. & CAPORICCI R. 1989. Características diferen-
ciales entre el fragmento del cráneo de Horno sp. de Venta Micena (Orce, Granada) y los équidos. Estudios
Geológicos 45: 121-138.
GIBERT J., RIBOT E, FERRÁNDEZ C., MARTÍNEZ B. & RUZ C. 1989. Diagnosis diferencial del frag-
mento del cráneo de Horno sp. del yacimiento de Venta Micena (Orce, Granada). In J. Gibert, D.,
Campillo, & E. García-Olivares, Eds, Los restos humanos de Orce y Cueva Victoria, pp. 31-108, Sabadell:
Institut Paleontológic Dr M. Crusafont, Diputació de Barcelona.
GIBERT J. & JIMÉNEZ C. 1989. Investigations into cut-marks on fossil bones of Lower Pleistocene age from
Venta Micena (Orce, Granada, Spain). Human Evolution 4,
GIBERT J., MARTÍNEZ B., CAPORICCI R., JIMÉNEZ C., FERRÁNDEZ C. RIBOT E, SORIA E J.,
PÉREZ-CUADRADO J.L., ARRIBAS A., CANALS J., GARCÍA-TARGA J.M., IGLESIAS A. &
ROMERO R. 1989. Resumen de las investigaciones paleoantropológicas y arqueológicas de Orce (Gra-
nada) y Cueva Victoria (Cartagéna). COL-PA 42, 11-60.
GIBERT J., CAMPILLO D., MARTÍNEZ B., SÁNCHEZ E, CAPORICCI R., JIMÉNEZ C., FERRÁNDEZ
C. & RIBOT E 1991. Nouveaux restes d' hominides dans les gisements d'Orce et de Cueva Victoria
(Espagne). In E. BONIFAY & B. VANDERMEERSCH, Eds, Les premiers européens. Actes du 114éme
Congrés National des Societés Savantes (Paris 3-9 avril 1989), pp. 273-282. Paris: Comité des Travaux
Historiques et Sociales.
GIBERT J., ARRIBAS A., MARTÍNEZ B., ALBALADEJO S., GAETE R., GIBERT L., PEÑAS C. &
TORRICO R. 1992. Síntesis cronoestratigráfica del Pleistoceno inferior de la región de Orce. In J.
GIBERT, D. CAMPILLO, E. GARCÍA-OLIVARES, A. MALGOSA, E MARTÍNEZ, B. MARTÍNEZ,
M. WALKER, P. PALMQVIST, E SÁNCHEZ & A. ARRIBAS, Eds, Proyecto Orce-Cueva Victoria (1988-
1992). Presencia humana en el Pleistoceno inferior de Granada y Murcia, pp. 107-114, Orce: Museo de
Prehistoria José Gibert.
GIBERT J., FERRÁNDEZ C., MARTÍNEZ B., CAPORICCI R. & JIMÉNEZ, C. 1992. Roturas antrópicas
en los huesos de Venta Micena y Olduvai. Estudio comparativo. In J. GIBERT, D. CAMPILLO, E.
GARCÍA-OLIVARES, A. MALGOSA, F. MARTÍNEZ, B. MARTÍNEZ, M. WALKER, P. PALMQVIST,
F. SÁNCHEZ & A. ARRIBAS, Eds, Proyecto Orce-Cueva Victoria (1988-1992). Presencia humana en el
Pleistoceno inferior de Granada y Murcia, pp. 283-305, Orce: Museo de Prehistoria José Gibert.
GIBERT J., IGLESIAS A., MAILLO A. & GIBERT L. 1992. Industrias líticas en el Pleistoceno de la región
de Orce. In J. GIBERT, D. CAMPILLO, E. GARCÍA-OLIVARES, A. MALGOSA, E MARTINEZ, B.
MARTÍNEZ, M. WALKER, P. PALMQVIST, E SÁNCHEZ & A. ARRIBAS, Eds, Proyecto Orce-Cueva
Victoria (1988-1992). Presencia humana en el Pleistoceno inferior de Granada y Murcia, pp. 219-281, Orce:
Museo de Prehistoria José Gibert.
GIBERT J. & PALMQVIST P. 1992. Dimensión fractal de las suturas del cráneo de Orce. Revista Española de
Paleontología 7: 1-11.
GIBERT J., SÁNCHEZ E, MALGOSA A., MARTÍNEZ B., M. WALKER Y RIBOT E 1992. Nuevos restos
humanos en los yacimientos de Orce y Cueva Victoria. in Proyecto Orce-Cueva Victoria 1988-1992: Presencia

98
 HUMERAL FRAGMENTS ATTRIBUTABLE TO Homo SP. FROM LOWER PLEISTOCENE SITES ATVENTA

humana en el Pleistoceno inferior de Granada y Murcia. Ed. Museo de Prehistoria "José 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., ARQUÉS J.M., GARCÍA OLIVARES E., BORJA C., LOWENSTEIN G. 1998.
Hominid status of the Orce cranial fragment reasserted. Journal of Human Evolution, 34: 203- 217.
GONZÁLEZ J.M. & PALMQVIST P 1990. Sobre la caracterización biométrica del crecimiento y la forma de
los foraminíferos planctónicos. Aplicación de las series de Fourier al análisis de la forma de las cámaras.
Revista Española de Paleontología 5: 81-90.
KENNEDY G. E. 1973. The anatomy of Middle and Lower Pleistocene hominid femora. London, University of
London, Ph.D. thesis.
KNUSSMANN R. 1967. Humerus, Ulna und Radius der Simiae.Vergleichende morphologische Untersuchungen
mit Berücksichtigung über die Funktion. Bibliotheca primatologica 5: 1-339.
LOHMANN G.P. 1983. Eigenshape analysis of microfossils: a general morphometric procedure for describing
changes in shape. Mathematical geology 15: 659-672.
LOHMANN G.P. & SCHWEITZER. P.N. 1990. On eigenshape analysis. In EJ. Rohlf & EL. Bookstein, Eds,
Proceedings of the Michigan Morphometrics Workshop, pp. 147-166, Ann Arbor, University of Michigan Museum
of Zoology, Special publication number 2.
LOWENSTEIN G. 1995. Immunological reactions on fossil bones from Orce. Orce. Orce Conference. Abstracts.
Orce 1995.
MARTÍNEZ B. 1991. Revision sistemática y estudio cuantitativo de la fauna de macromamíferos del yacimiento de Venta
Micena (Orce, Granada). Barcelona: Universitat Autónoma de Barcelona, Ph.D. thesis.
MARTÍNEZ B. 1992. Revisión sistemática de la fauna de macromamíferos del yacimiento de Venta Micena
(Orce, Granada, España). In J. GIBERT, D. CAMPILLO, E. GARCÍA-OLIVARES, A. MALGOSA, F.
MARTÍNEZ, B. MARTÍNEZ, M. WALKER, P. PALMQVIST, E SÁNCHEZ & A. ARRIBAS, Eds,
Proyecto Orce-Cueva Victoria (1988-1992). Presencia humana en el Pleistoceno inferior de Granada y Murcia, pp.
21-85, Orce: Museo de Prehistoria José Gibert.
MARTINS C. 1868. Sur la torsion de l'humérus. Bulletin de la Société Anthropologique de Paris 3: 320-328.
MENDOZA M., PALMQVIST P., GUERRERO S., MARTÍNEZ B., PÉREZ-CLAROS J.A., GIBERT J. &
EISENMANN V. 1993. Consideraciones tafonómicas, paleoecológicas y paleoetológicas sobre la comu-
nidad de macromamíferos de Venta Micena (Orce, Granada). In Comunicaciones. IX Jornadas de Paleontología,
pp. 73-78, Málaga, Universidad Málaga, Sociedad Española de Paleontología.
MOYA-SOLÁ S. & AGUSTÍ J. 1989. Una reinterpretación de fragmento craneal de Orce: Equus stenonis. In
J. GIBERT, D. CAMPILLO & E. GARCÍA-OLIVARES, Eds, Los restos humanos de Orce y Cueva Victoria,
pp. 447-451, Sabadell: Institut Paleontológic Dr. M. Crusafont, Diputació de Barcelona.
MOYA S. & KÓLHER M 1997. Anatomy of a mistake. Journal of Human Evolution, 33: 911-97.
NORUSIS M.J. 1988. SPSS-X Advanced statistics guide. Chicago: SPSS Inc.
OLIVIER G. 1951. Pratique anthropologique. Paris: Vigor Fréres.
PALES L. & LAMBERT C. 1971. Atlas ostéologique pour servir á Pidentification des manuniféres du Quaternaire. Vol.
1, Paris: Centre National de la Recherche Scientifique.
PALMQVIST P., GIBERT J. & MARTÍNEZ B. 1992. Sobre la relación tamaño/abundancia en los macrornarníferos
de Venta Micena y sus implicaciones tafonómicas. Revista Española de Paleontología 7: 174-180.
PALMQVIST P. 1997.A critical re-evaluation of the evidence for the presence of hominids in lower pleistocene
times at Venta Micena, soutthern Spain. Journal of Human Evolution, 33: 83-89.
PÉREZ-PÉREZ A. 1989. La falange de Cueva Victoria: análisis discriminante y afiliación taxonómica. In J.
GIBERT, D. CAMPILLO & E. GARCÍA-OLIVARES, Eds, Los restos humanos de Orce y Cueva Victoria,
pp. 407-413, Sabadell: Institut Paleontológic Dr. M. Crusafont, Diputació de Barcelona.
POMBA B. 1950. Clave para la descripción de mamíferos de la U.R.S.S. a partir del esqueleto. Moscow: Boletín
de la Comisión del Pleistoceno.
PONS-MOYA J. 1985. Nota preliminar sobre el hallazgo de Horno sp. en los rellenos cársticos de Cueva Victoria
(Murcia, España). Endins 10-11: 47-50.
REYMENT R.A., Blackith R.E. & Campbell N.A. 1984. Multívariate morphométrics. London, Academie Press.
ROHLF F.J. 1990. Fitting curves to outlines. In F.J. Rohlf & EL. Bookstein, Eds, Proceedings of the Michigan
Morphometrics Workshop, pp. 167-177, Ami Arbor, University of Michigan Museum of Zoology, Special
publication number 2.
SENUT B. 1983. Quelques remarques á propos d'un humérus d'hominoide pliocéne provenant de Chemeron
(bassin du lac Baringo, Kenya). Folia Primatologica 41: 267-276.

99
 GIBERT, J., MALGOSA, A., SÁNCHEZ, E, RIBOT, E AND WALKER, M.J.

SISSON S. 1947. The anatomy of the domestíc animals. Philadelphia and London: W.B. Saunders.
SORIA EJ. 1986. El Neógeno-Cuaternario en el sector de Orce (Depresión de Guadix- Baza). Granada: Universidad
de Granada, Ph.D. thesis.
SORIA EJ., LÓPEZ-GARRIDO A.C. & VERA J.A. 1987. Análisis estratigráfico y sedimentológico de los
depósitos neógeno-cuaternarios en el sector de Orce (depresión de Guadix-Baza). In S. MOYÁ-SOLA,
J. AGUSTÍ, J. GIBERT & J.A. VERA, Eds, Paleontológía i Evolució. Memoria especial núm. 1. Geología
y paleontología del Pleistoceno inferior de Venta Micena, pp. 11-34, Sabadell: Institut Paleontológic Dr. M.
Crusafont, Diputació de Barcelona.
STRANEY D.O. 1990. Median axis methods in morphometrics. In EJ. Rohlf & EL. Bookstein, Eds, Proceedings
of the Michigan Morphometrics Workshop, pp. 179-200, Ann Arbor, University of Michigan Museum of
Zoology, Special publication number 2.
TATTERSALL I., DELSON E. & VAN COUVERING J. 1988. Encyclopaedia of human evolution. New York:
Garland Publishing.
VERA J.A., FERNÁNDEZ J., LÓPEZ-GARRIDO A.C. & RODRÍGUEZ-FERNÁNDEZ J. 1984. Geología
y estratigrafia de los materiales plio-pleistocenos del sector de Orce-Venta Micena (prov. Granada). Paleontológia
i Evolució 18: 3-11.
YOUNKER J.L. & EHRLICH R. 1977. Fourier hiometrics: harmonic amplitudes as multivariate shape descriptors.
Systematic Zoology 26: 336-342.
ZIHILMAN A., LOWENSTEIN G. 1996. A Spanish Olduvai?. Current Anthropology, Volume 37, Number 4,
August-October, 1996

TABLE 1
Venta Micena Fauna.This revised minimal faunal list is based on Martínez (1991, 1992) whose
Megantereon sp. has subsequently been assigned to M. whitei (Palmqvist & Martínez, n.d.)

Horno sp.
Canís falconerí MAJOR
Canis etruscus MAJOR
Vulpes praeglacialis KORMOS
Hornotherium latidens OWEN
Megantereon whitei BROOM
Lynx sp.
Pachycrocuta brevirostris AYMARD
Ursus etruscus CUVIER
cf. Meles sp.
Hippopotamus amphibius antiquus DESMAREST
Praeovibos sp.
Soergelia mínor MOYA-SOLA
Bubalus sp.
Capra alba MOYÁ-SOLÁ
Praemegaceros solílhacus ROBERT
Cervidae gen. et sp. indet.
Dícerorhinus etruscus brachycephalus SCHROEDER
(syn. Stephanorhinus etruscus, thus Cerdeño, 1993)
Equus stenonis granatensis ALBERDI & RUIZ-BUSTOS
Mammuthus merídionalis NESTI
Prolagus calpensis MAJOR
Oryctolagus cf. lacosti POMEL
Desmana sp.
Allaphaíomys pliocaenicus KORMOS
Castillomys crusafonti subsp.
Apodemus aff. mystacinus DANFORD & ALSTON
Eliomys intermedius FRIANT
Hystrix major GERVAIS
Testudo sp.
Lacerta sp.
Ophídia indet.
Rana sp.
Charadriiforme indet. (aff. Laridae)

100
 HUMERAL FRAGMENTS ATTRIBUTABLE TO Homo SP. FROM LOWER PLEISTOCENE SITES ATVENTA

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 Max. diam. MM. diana.

(mm) (mm) (mm)

Ursus arctos (adult) 190 44 33

Panthera leo (infant) 85 28 21
Canis familiaris (adult) 126 23 15
Acinonyx jubattts (adult) 165 24 16
Panthera pardos (adult male) 165 30 19
Panthera pardos (adult female) 115 20 15
Papio hamadryas (adult) 128 20 12
Papio harnadryas (young) 125 16 12
Mandrillus sphynx (adult male) 180 22 17
Mandrillus sphynx (adult female) 143 17 11
CO-4 (adult) 310 23 18
CO-6 (adult) 270 25 19
CO-1 (infant) 100 11 9
CO-2 (infant) 110 13 11
IP C-32A (infant) 100 11 9
IP (unnumbered) (infant) 80 10 8
IP-2 (infant) 95 10 8
IP-19 (infant) 155 14 10
IP C32B (infant) 160 12 11
IP SW-2 (infant) 200 18 13
VM-1960 185 14 11
VM-3691 - 20 14

3 4 5 1

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.

101
 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- O
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, u9
N-2= Roman cemetery at Tarragona, MZ= cemetery at Mequinenza, Zaragoza, M-1= Cercopithecoid monkeys: 68.2= adult male
Mandrillus 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.
u
o
o e
1111141N0ID8 CSRCOPITIMCOID XONUYS casetrycetes
c./)

Ch Ch J J 3 A A A
o
I T Ch Ch Ch Ch Ch Ch Ch 4
ó N
N? 1960 C-32 CO1 00 135 T32 130 5-1 M-2 M2 CO3 9-1 193 C04 CO3 C06 68.2 108.1 6974 35.3 1690 68.1 23.10 131.5 109.1 c2512 LYNX
111
33 71 51 63 72 00 73 64 78 40 16 90 60 91 07 17 95 76 75 70 73 75 se 62 GO 73 66
72
el. 77 45 46 94 90 54 78 17 es 93 75 75 94 00 10 72 67 56 73 78
12 64 01 73 73 12 17 u
91 71. 115 el 79 52 90 45 11 09 114 14 92 34 71 53 12 16 71 77 94 75 41 70 69 SI 64 71 o
90 92 93 el 59 94 54 14 75 76 00 73 73 17 66 76 71 74 65 70 70
117 75 13 77 71I 90 II
93 13 13 90 II II 52 96 77 93 15 67 17 19 66 75 61 65 71 77 17 75 es 60 eo 51 74 12 01
57 55 96 00 54 00 03 93 71 10 11 es 63 $1 15 02 83 60 47 42 es 9
55 43 MI 75 92 33 15
77 10 100 47 73 41 45 55 49 79 11 76 83 90 75 77 el 111 69 77 e0 65 49 65 74 00 90 zzz

44 59 59 80 91 69 113 05 78 17 90 43 47 54 33 73 89 43 60 72 14
12 75 95
53 66 II 71 51 79 94 39 89 70 69 17 76 44 37 44 53 69 87 O 49
90 46 15 55 62 73 69 35 69 51 35 ti 63 45 73 80
56 48 52 39 44 94 87 33 42 13 32 56 73
67 45 79 76 70 39 53 53
89 71 311 53
80 52 44 10
57 40
33
 HUMERAL FRAGMENTS ATTRIBUTABLE TO HOMO SP. FROM LOWER PLEISTOCENE SITES ATVENTA

TABLE 4

Standardized canonical coefficients. Correlations of variates with canonical

for discriminant functions 1 and 2 discriminant functions 1 and 2

Function 1 Function 2 Function 1 Function 2

GSIp 0.7531 -1.5045 -0.1832 -0.4006

C1 p -7.1643 1.2175 0.0704 0.3859
C2p 5.6241 -2.6971 0.0418 0.5024
FRp -0.3849 2.2037 -0.3043 0.6214
SR2p -0.4402 -0.3764 0.2118 -0.6169

H2p 0.5679 1.8039 -0.1203 -0.6740

H3p 1.0767 0.3959 0.8932 0.2473
H4p 0.3732 0.8200 0.6185 -0.0050

GSId 0.3124 -0.2636 -0.4835 -0.2479

Ci d 3.8898 1.0693 0.4643 0.3022
C2, -1.4236 -0.3772 0.4497 0.3225
FRd -0.3627 0.8108 -0.1699 0.2905

H3 d -0.5423 0.9414 -0.1074 0.6005

Areap/
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= radial
variance (Davis 1986). H2, H3 and H4= harmonic amplitudes of order 2, 3 and 4. Area p/Areaa = mean proximal
area divided by mean distal area. Subscript p = proximal. Subscript d = distal.

Function 1 Function 2

Eigenvalués" 10,57 5,42

% of explained variance 66,12 33,88
canonical correlations 0,956 0,919
Wilk's lambda 0,0135* 0,1559
chi-square test 96,915* 41,82
degrees of freedom 28* 13
significante 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 Range

Horno (adult) 150 145.1° 134.0°-156.2°

Macaca 102 98.8° 92.1°-105.5°
Papio 10 96.0° 82.2°-109.8°
Ccrcocebus 8 99.1° 92.6°-105.6°
Cercopithecus 16 97.0° 88.5°-105.5°

Table 5a: after Senut 1983, who based calculations on data froni Knussmann 1967.

103
 GIBERT, J., MALGOSA, A., SÁNCHEZ, E, RIBOT, E AND WALKER, M.J.

Specimen Valu e

Child MZ-124-2 128°

Fossil VM-1960 131°
Child MZ-87 137°
Child IP-C-32 139°
Child MZ (unnumbered) 139°
Child 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).

Figure 1: Computer-assisted simulations

of a humeral cross-section obtained from
Fourier series fitted to the outline.
Numbers indicate the harmonic order
used in each simulation. Amplitude
values of the second, third and fourth
harmonics are shown, accompanied by
Fnurth harrrtonie
the corresponding simulations obtained
Second h.arm clic Third harmonic
p litu de •• 0.2073 Amp litu de 0.1061 Amplitude 0.0801 using only these harmonics.

104
 HUMERAL FRAGMENTS ATTRIBUTABLE TO Homo SP. FROM LOWER PLEISTOCENE SITES ATVENTA

1
Figure 2b: Measurements of cortical thickness taken on
Figure 2a: Computer-assisted tomographically cross-sections of humeral shaft (after Kennedy, 1973).
superimposed images of proximal and distal cross- a= anterior cortical thickness. b = posterior cortical
sections of humeral shafts, from which the angle of thickness. c= anteroposterior dimension. d= lateral
humeral torsion was determined. MM= medial margin. thickness. e= medial thickness. f= mediolateral dimension.
ML= lateral margin. CT= margin between greater
tuberosity and deltoid tuberosity of the proximal cross-
section; the other cross-section is in the supratrochlear
region at the apex of the olecranon fossa: the angle
of humeral torsion lies between the two mediolateral
dimensions.

A BC 0 F GH

G e %
•
% es Ç)
ti 5 'C
Z r

á
'

J Q Z G

6 5 a c, PP.

a l'
t © h °'

9 e e z c) ,z -

0 0 ¿D a o. D Figura 3a: Comparison of humeral

diaphyseal cross-sections of carnivores,

C c-3 cp cercopithecoid monkeys, and Venta

Micena fossils. Scale in centimetres.
c (1) 0 A= adult Acinonyx juba tus. B= adult
male Panthera pardus. C= adult male

0 ®4 c Mandrillus sphynx. D= Macaca sylvana.

E= young Papio hamadryas. F= Macaca
sylvana. G= VM-1960 fossil from
Venta Micena. H= VM-3691 fossil
from Venta Micena.

105
 GIBER.T, J., MALGOSA, A., SÁNCHEZ, E, RIBOT, E AND WALKER,

A 3 C D F G H I

o o o o z) % 41,
o co. o o z) z Ç) to
o ci) ® o O i, so Ex' 'I-
Q e o e c, o O
oolooDs
e..
e CID 0-0,(C.)
1

n 15 C) O (3 CZN e 0 DiSt

O G 1 acz, Cr*

o o z ,c_-,
G o z ,n, cp
D)jc_ 9
zb
ga ---

át. ;,,,

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).

106
 HUMERAL FRAGMENTS ATTRII3UTABLE TO HOMO SP. FROM LOWER PLEISTOCENE SITES AT VENTA

n
A BCD E

o
o'
o
O
%
PR 0.X.

o S
DI ST.

O 9
0 %
O(11 cl 10 cm

O
O O C9)
D o

iL C1'
__•'
3

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.

107


GIBER.T, j., MALGOSA, A SÁNCHEZ, F, RIBOT, F. AND WALKER, M.J.

100
911-/
80 Figure 4: Comparison of values for the
' diaphyseal index (y-axis) at cross-sections 15
70 n ,
mm apart (x-axis: 1= most distal, at apex of
60 A
olecranon fossa; number of cross-sections is
determined by length of shaft and biological
50 age).VM-1960= Venta Micena fossil diaphysis.
Figure 4a: the carnivores are as follows: 109-
40 1= adult Acinonyx jubatus, 23-10= adult male
30 Panthera pardus, Canis= adult Canes familiares.

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

100
AB'
90
80
70 Figure 4b: the cercopithecoid monkeys are as
follows: 68-2= adult male Mandrillus spliynx,
60- 35-3= Macaca sylvana, 1690= adult Papio
hamadryas, 106-1= Papio hamadryas, 68-1=
Macaca sylvana.
401-
30

VM-1960 35-3 - 19-- 68-2

° 66-1 K. 106-1 1693

100
90-1
80
70- Figure 4c: the Horno specimens are as follows:
CO-2= child from Mediaeval cemetery at Orce,
60 Granada, N-1 and N-2-= children from Roman
cemetery at Tarragona, IP C-32= iron-age child
50 from Illot de Porros, Menorca.
40
30

VM-1960 -- N-1 N-2

'3- IP-32 CO-2

108
 HUMERAL FRAGMENTS ATTRIBUTABLE TO Homo SP. FROM LOWER PLEISTOCENE SITES ATVENTA

C ERCOPI THE COI D Figure 5: Comparison of fossil VM-

MONKEYS 1960 with values of the diaphyseal índex
100 _
(y-axis) in cercopithecoid monkeys
(Figure 5a) and Homo (Figure 5b), taken
80 - at successive heights (x-axis) aboye the
apex of the olecranon fossa. Broken
60 - lines represent the 95% confidence
limits.
40 - —3 P
\14".
Yr.
20 - * 4 VM-1960

HOMO
100 _

0 610 90 120 1 50
1 180 (mm)

-s Second dhscrimjnánt function

(33..1O31,i, nl varinnre

-4

-3
...„.
■ r^ 11/`,
d'- 2
° O O '■ p
O
,
•
1
O '\
CRF/44) 1
® 0
•
% t
1
. O,
wt

-. i ‘ -",4 0 1 : 1 L 1 I
-1 -a .-3.^ -2 .1 ..; 2 t 3 4 5 . 1 7
`----__,- e w6
Fi rst d isdiminant fu ndion --I
-(66,12% ni vaaiaore r-plaínccli

- -2 CERCOP

O fromo
A
• C,r,orilh5cid:
A earnivercs
*Grnup centrnida
■
/A
w-m Gona
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<0.05
confidence ellipses around group centroids.

109
 % 10 20 30 40 50 60 70 80 90
1 1
AT- 93
1808M VM1960 CV 1
AP • •

1 3
2j4 5i 6 VM 3691
AT- 217 AT- 25

x
o 46 2 5 VM1960
y y
ML • ** CV1
VM 3691
1 3
AT 217 AT -25

Figure 7: Comparisons of cortical thickness in fossil specimens and living species (from Arsuaga et al. 1991 modified). Carnivores: 1 = Lynx (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 = prehistoric
Holocene Horno sapiens from Sepúlveda, Segovia. AP= anteroposterior index. ML= mediolateral index.
 HUMERAL. EFRAGMENTS ATTRIBUTABLE TO Homo SP. FROM LOWER PLEISTOCENE SITES ATVENTA

V
Figure 8: Comparison of medullary cavity-size in mid-shaft humeral cross-sections. a: CV-2 fossil humeros from
Cueva 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 hamadrías humerus; g: Hornotherium latídens fossil
humerus from Venta Micena, and h: male Panthera pardus humerus. (x 1.25)

111