Escolar Documentos
Profissional Documentos
Cultura Documentos
reasserted
J. Gibert*, *Institut de Paleontología, Escola Industrial 23, 08201 Sabadell, Spain
D. Campillo†, †Museu Nacional d’Arqueología de Catalunya, S. Madrona 39,
08031 Barcelona, Spain
J. M. Arqués§, ‡Universidad de Granada, Facultad de Medicina, Av. de Madrid s.n.
E. Garcia-Olivares‡, 18012 Granada, Spain
C. Borja‡ & §Departament D’Informàtica, Fac. Ciències, Universitat Autònoma de
J. Lowenstein¶ Barcelona, Spain
¶University of California, School of Medicine, Department of Medicine,
San Francisco, U.S.A.
Figure 1. (a) Internal cranial surface of Equus sp. (Laboratory of Anatomy, Paris), 0–2 months. *Lambda.
T, Tentorial bony apophysis; SI, unobliterated internal lambdoid suture; C, unobliterated internal coronal
suture; SA, obliterated internal sagittal suture; B, obliterated internal interfrontal suture. (b) Frontal and
obelic region of a 4–5 month old equid. 1–1, sagittal suture. * Position of lambda comparable to that
on VM-0. A–A, lambdoid suture; B–B, interfrontal suture; C–C, coronal suture; SI, lambdoid suture;
OC, occipital; P, parietal; F, frontal.
(b)
206
.
ET AL.
Figure 2. (a) Internal surface of VM-0. 1–1, sagittal suture; 2–2, 3–3, 4, 5, 6, posthumous cranial fissures. *, Cortical scale.
OC, apical fragment of the occipital. C, occipital crest. *Lambda. It is possible to observe that the cortical scale 2(a)
corresponds to the presumed indentation of the presumed coronal suture of Agustí & Moyà (1987). Also the presumed
coronal suture coincides with part of the trace of the postmortem fracture. (b) Occipital bone from a current human specimen
showing a sagittal crest which reaches lambda. 1, superior sagittal crest; 2, right transversal groove; 3, left transversal groove;
4, inferior transversal groove; 5, Endinion.
207
Figure 3. Photograph of the VM-0 X-ray. The bone shows rounded zones with great density, that
correspond to the ruga cerebralia, whereas those that have less density correspond to the digitate markings.
The dotted area that is apparent in greater or lesser clarity, over the bony surface represents substantia
spongiosa indicating diploe of the cranial bone. 1–1, sagittal suture, in which the indentations show zones
of greater density formed by the substantia compacta that covers it; at the level of the obelionic region, a
small wormian bone (W) is present. Continuing anteriorly, the suture is straighter than in the obelic region
and anterior to the fissure 2–2, it is deformed posthumously and is inclined slightly toward the left side.
There is a greater density here on account of carbonatic concretions. It is important not to confuse the
small points of the parasagittal concretions (K) with the suture. OC corresponds to the small apical
fragment of the occipital that shapes lambda. The straight occipital crest (C) is clearly present and is
diverted towards the right. 2–2, a posthumous cranial fissure, that should not be confused with a supposed
coronal suture, because its direction is oblique and it does not meet the sagittal suture at a right angle.
Moreover, it is easy to verify that the substantia spongiosa reaches its edges and in no point is covered by
substantia compacta, as is apparent for the sutures, 3–3, 4, 5 and 6, also are postmortem fissures.
our opinion this interpretation is not correct. argued (Figure 4). Furthermore, Gibert
In VM-0 15 mm of crest on the occipital is et al. (1993) show that the kenyan hominid
preserved; it is small in relation to the large KNM-ER 3733 has an occipital crest which
equid tentorial process in this position. This reaches lambda. This means that VM-0
is sufficient to prove that the VM-0 crest is cannot be excluded from Homo by this
not an equid crest, as we have previously character.
208 . ET AL.
Figure 4. (a) Comparison of transverse and sagittal sections from the obelionic region of two equid crania
(0–1 month old and 7 months old) and of VM-0. 1, Transverse section at lambda; 2, transverse section
10 mm posterior to lambda. (b) VM-0 superimposed on a sagittal section of an equid skull (4–5 months
old). The sections are superimposed with lambda aligned. (c) Comparison of the size and sagittal
curvature of VM-0 and an equid (7 months old). 1, Sagittal sections at lambda; 2, sagittal sections 10 mm
posterior to lambda.
and it is not obliterated (degree of obliter- deviation of 13) (Campillo, 1989). This is
ation of Broca=0). On both the internal clearly visible in the internal face of VM-0
and external surface, the suture is similar and in Figure 5.
to that of humans in both morphology Agustí & Moyà (1989, Figure 1 in that
and fractal dimension. The fractal dimen- paper: p. 449), Moyà & Köhler [1997,
sion values are: sagittal external (infant Figure 1(a)] drew the sagittal and coronal
human)=1·138–1·201, VM-0=1·106; sagit- suture over the original drawing of Bergante
tal internal (infant humans)=1·167–1·051, [in Campillo (1989), Figures 7 and 8:
VM-0=1·128 (Arqués & Gibert, 1998). The p. 194–195]. The drawing of the internal
lambdoid suture on both the external and sagittal suture made by these authors is not
internal surfaces is also simple and not precise and it is not possible to detect the
obliterated. The right segment is 22 mm deviation of 13). As a result, the presumed
long and the left 23 mm [Figures 2(a), 5(a, coronal suture, following partly posthumous
b) and 6(a, b)]. The pattern of these sutures fractures [Figures 2(a) and 3], points 2–2;
on the internal face is different from the Gibert et al., 1989a–c), seems to be sym-
external pattern, as it is normal in humans metric with and perpendicular to the sagittal
(Testut & Latarjet, 1975; Masset, 1982). suture [Figure 5(d)].
In the correct drawing of Bergante and
also in the X-ray it is possible to observe how
False coronal suture
the sagittal suture deviates [Figure 5(b)].
Agustí & Moyà (1987: p. 536), Moyà & If we consider that the correct trace of
Agustí (1989: p. 448, lam. 1) and Moyà & the sagittal suture on the external surface
Köhler [1997; Figure 1(a)] suggest the followed the points 1–2 in Figure 5(a, b),
existence of a presumed coronal suture. the coronal suture would have to move
They indicate the trace of the presumed anteriorly [as is indicated in Figure 5(a)] in
internal coronal suture on photographs of order to be perpendicular and symmetric
the internal face of the right parietal. Only with the correct interpretation of the sagittal
Moyà & Köhler (1997) indicate presumed suture. Neither direct observation, nor the
indentations of the presumed coronal X-ray, or the drawing of Bergante shows any
suture, on the external face of the right indication of coronal suture in this area.
parietal. All of the points indicated by these Agustí & Moyà (1987) and Moyà &
authors on both faces of the right parietal Agustí (1989) invoke obliteration to justify
correspond to cortical scales or fractures the existence of a coronal suture. In our
(Gibert et al., 1989a–c; Figure 6). opinion they make serious errors in obser-
In reply, we argue that: (1) the coronal vation and method because they have not
suture in hominids, equids and other studied the sequence of obliteration in
mammals is always symmetric and perpen- equids. In the equids, we have observed the
dicular to the sagittal suture; (2) in VM-0 following. (1) The endocranial aspect of
it is essential before drawing the coronal the sagittal suture and the internal inter-
suture to study the X-ray data, which permit frontal suture are the first sutures that are
sutures to be distinguished from fractures obliterated in equids, and this occurs at
(Figure 3); (3) it is important to note that on 2–3 months of age [Figure 1(a)]. (2) The
the internal and external faces, cortical external sagittal suture is the second suture
scales are present [Figures 2(a), 6]; and (4) that is obliterated. This process begins
the anterior portion of the parietals has not distally and continues progressively towards
been correctly restored. This is the reason the coronal suture (three examples at the
why the sagittal suture in this zone has a VUAB, four examples at the MZB, 951160,
212 . ET AL.
950410, 941229, 941159, one example at and similar to that of modern human
the IPS). The formation of the external infants, and it is also similar to the suture
sagittal crest coincides in equids with this drawn by us. The values of the fractal
obliteration. (3) The third suture that is dimensions [Agustí & Moyà (1987), Df=
obliterated is the external interfrontal 1·128; Moyà & Köhler (1997), Df=1·162]
suture. This process begins in equids before can be included within the modern human
the external sagittal suture is totally obliter- infant range of variation [1·078–1·167;
ated (an example at the VUAB, two at the Arqués & Gibert (1998)].
MZB, 941229, 950410, one example at In equids the internal sagittal suture is
the IPS). (4) The last sutures that are very simple from 0 to 2 months of age
obliterated in equids are the coronal and the (Gibert et al., 1989; Figure 5) and after this
lambdoid. age it is obliterated, as can be observed in
Therefore, if VM-0 was an equid, it could the example from the LAP (Figure 1).
not have an obliterated coronal suture at These arguments imply that the suture
the same time as having an unobliterated drawn by these authors can not belong to an
external and internal sagittal suture. This equid older than 2 months.
finding contradicts Agustí & Moyà (1987) Palmqvist (1996: p. 19) described and
and Moyà & Agustí (1989), and invalidates drew the external sagittal and lambdoid
the claim that a coronal suture is present suture of VM-0 based on Moyà’s interpret-
in VM-0. Furthermore this characteristic ation of the sutures drawn for detailed
permits VM-0 to be included in Homo. photographs. The internal sagittal suture is
The X-ray (Figure 3) shows the following not very difficult to follow, but it is necessary
features. (1) The sagittal and lambdoid to take care when the external sagittal suture
sutures do not show any sign of synostosis. is drawn, because it is eroded (as is normal
(2) Several posthumous fractures are in the Venta Micena fossils) and the cortical
apparent. The most important of these is the scales, small fissures and carbonatic precipi-
fracture 2–2, which Agustí & Moyà (1987), tations must be discriminated from the trace
Moyà & Agustí (1989) and Moyà & Köhler of the real suture. This makes direct obser-
(1997) confuse with a suture without notic- vation of the fossil necessary, in order to
ing the following that the cancellous bone interpret correctly anatomical and tapho-
reaches the fracture line, which annuls the nomical factors. It is not possible to trace the
suture interpretation because the edges of correct suture over a photograph (Figure 6).
the sutures are always covered by compact Because we feel that the suture trace of
tissue (it is well known that the cancellous Palmqvist (1996) is not correct, his fractal
bone is never visible in normal bone); and dimension value (Df=1·335) is not relevant
that at the edges of sagittal and lambdoid to the taxonomic interpretation of VM-0.
sutures, compact tissue can be seen. (3) The Palmqvist’s (1996) fractal dimension value
digitate impressions are apparent and are (Df=1·335) suggests to him that VM-0
normal for the age of the subject. belonged to an equid of 4–5 months of age
[Figure 1(b)]. At this age, however, the
equids have an obliterated internal sagittal
Complexity of the VM-0 sagittal and
suture [Figure 1(a)], which is not the case in
lambdoid sutures according to Moyà &
VM-0 (Agustí & Moyà, 1987; Moyà &
Köhler and Palmqvist
Agustí, 1989; Moyà & Köhler, 1997). In
The endocranial sagittal suture described by relation to the detail of the posterior part of
Agustí & Moyà (1987), Moyà & Agustí the sagittal suture and the parieto-occipital
(1989) and Moyà & Köhler (1997) is simple suture of VM-0, Moyà & Köhler (1997) say
213
that the complexity of the sutures is clearly within the range of variation of juvenile large
visible. As we have mentioned earlier, the mammals. According to them
distal part of the sagittal ectocranial suture
of VM-0 is the most complex, but the rest of ‘‘the degree of the parietal curvature (sagittally
and transversally), the general size of the speci-
the suture becomes simpler, as can be
men, and the slenderness of the bone suggest
observed in Figure 6(a–c) and Figure 2 of that it belongs to an infant or juvenile large
Moyà & Köhler (1997). Therefore, in our mammal. Curvatures of surfaces of the size of
opinion, the interpretation of these authors the Orce fragment (75 mm transversal diam-
is not correct. eter) can easily be found in other mammals
such as young equids, rhinos etc.’’
of modern colts. The comparison with The drawing of the sagittal suture presented
VM-0 does not allow any doubt. VM-0 is by Palmqvist (1996, 1997) is not correct.
different in sagittal curvature, in the length (4) The coronal suture does not occur in
of the sagittal arc and in the angle of the VM-0, whether the obliterated zones are
lambdoid sutures. For more details see considered (Agustí & Moyà, 1987; Moyà &
Gibert et al. (1989a–c). Agustí, 1989) or not (Moyà & Köhler,
1997). (5) The digitate impressions of
VM-0 are similar to those present in modern
Anatomical conclusions
humans. (6) VM-0 is very different from
VM-0 presents three important anatomical the infant E. granatensis parietal found at
characters, which point to a large cranial Venta Micena and other infant equids and
capacity, the main characteristic of Homo. nonhuman mammals.
These characters also exclude VM-0 from
the equids and other mammals. These three
Biomolecules
characters are as follows: length of the
sagittal arch—longer in VM-0 (60 mm) Palmqvist (1997) maintains that the
than in the young equids (maximum 45, meanders of the sagittal suture of the Orce
minimum 15, average 32; Gibert et al., skull (VM-0) are a more reliable method of
1989a–c; large size of the parietal curves of species identification than the immunology
VM-0 [Figure 4(a, c)]; and large angle of macromolecules. DNA encodes proteins
between the lamboid sutures (105)). that determine morphology, but molecules
Moyá & Köhler (1997) asked ‘‘But where of DNA and proteins have highly species-
are the evident features diagnostic (autopo- specific sequences, whereas morphological
morphic) for humans?’’ We have a clear criteria such as skull shape and suture lines
answer, VM-0 has a large brain size. It is have an unavoidably subjective component,
also important to note that VM-0 is larger especially when the specimen is frag-
than are the skulls of 0–2 month-old equids; mentary, as is the case here. It would be
there is nonobliteration of the internal ideal if DNA and proteins persisted in
sagittal suture in VM-0, which is found in fossils in their original form, but of course
equids older than 2–3 months; and there is only a small proportion survive the chemical
nonobliteration of the external sagittal vicissitudes of weathering and diagenesis,
suture in VM-0, which is found in equids and those that do survive are frag-
older than 7 months. In our opinion this mentary and to some extent altered by
means that VM-0 can not be an equid. substitutions, racemization and other such
We have demonstrated that (1) the inter- processes.
nal occipital crest of VM-0 lies within the Nevertheless, under favourable circum-
range of human variability. It is not possible stances, it has been possible using radio-
to equate this crest with the tentorial bony immunoassay (RIA) to obtain useful
process of equids (Figure 4). (2) The phylogenetic information from extinct
internal sagittal suture has simple trace species such as mammoths and mastodons,
(Campillo, 1989), which is similar to that the South African quagga (a zebra-like
described by Agustí & Moyà (1987), Moyà equid), the Tasmanian marsupial wolf,
& Agustí (1989) and Moyà & Köhler Steller’s sea cow, 8-Ma-old fossil apes from
(1997). It can be included within the range Pakistan, and human remains that include
of the human infants by virtue of its fractal an Egyptian mummy and Homo erectus
dimension. The external sagittal suture also (Lowenstein, 1981, 1983; Lowenstein &
has a simple trace, similar to humans. (3) Scheuenstuhl, 1991).
215
A valid criticism of immunological tests detected was indeed integrated in the min-
on fossils is that there have not been eral structure of the bone, and was not
enough independent analyses of the same superficial, as would be expected if this
specimens, which would either validate or protein had originated from exogenous
refute the reliability of the various method- contamination.
ologies (Downs & Lowenstein, 1995). In the Another source Palmqvist (1997) cites in
present instance, however, independent support of his criticism of our results with
tests were carried out by two groups, one fossil bones is a review article reporting the
working at the University of Granada in discrepant results obtained in immunologi-
Spain (Borja & Garcia-Olivares, 1995) and cal studies of blood residues of prehistoric
the other at the University of California stone tools (Fiedel, 1996). Although
in San Francisco (UCSF), U.S.A. Palmqvist claims and erroneously reported
(Lowenstein, 1995). Different methods that Fiedel’s paper deals with ‘‘studies of
were employed (enzyme-linked immuno- proteins recovered from fossil bones and
sorbent assay and RIA), yet the results were blood residues from prehistoric artefacts’’,
nearly identical (Borja et al., 1997). the paper deals only with prehistoric arte-
Both detected human albumin in Venta facts, and not with fossil bones.
Micena fossils VM-0 and VM-1960. The Discrepant results from archaeological
UCSF team also detected human collagen stones cannot be extrapolated to fossil
and transferrin in VM-0. The Granada bones, in contrast to what Palmqvist appears
team detected human immunoglobulin in to be implying. Downs & Lowenstein
VM-1960 and in VM-3691. Both found (1995) found that archaeological blood
equine albumin in several fossils un- residues generally could not be identified
equivocally identified as equid. The UCSF reliably. Nevertheless, whereas proteins can-
team found bison albumin in two bovine not be clearly identified on artefacts only a
fossils. Neither team obtained albumin reac- thousand years old, these biomolecules can
tions in Cueva Victoria putative hominids survive under the right conditions and be
CV-1 or CV-2. The UCSF team also characterized in much more ancient fossil
detected anomalous human-like albumin in bones (Wykoff, 1972; Lowenstein, 1981;
one equid fossil. This has been used by Lowenstein, 1983; Rowley et al., 1986;
Palmqvist (1997) as the basis for his Ulrich et al., 1987). These reports clearly
criticism of the rest of the concordant results show that proteins from some fossil bones,
of both groups. although fragmentary, can persist and retain
Although great care is taken to avoid enough of their immunological properties
contamination from human hands or hair, to provide usable genetic information for
we have occasionally detected human periods of millions of years.
albumin on nonhuman fossils and stone Preservation in bones, unlike stones, is
tools, a well-known potential problem in all favoured by the proteins being embedded in
molecular evolutionary research. It seems the mineral phase that provides considerable
extremely unlikely, however, that the suite protection from the chemical environment.
of reactions described above are invalidated We are still learning which conditions of
by a single discrepancy. The overall findings fossilization are most likely to result in
of the two teams are concordant with each long-term stability of DNA and protein
other and include appropriate equine and molecules. Nevertheless, some authors
bovine controls. We also included strict con- found that there is no strict relation between
trols, especially with fossils assigned to the biomolecule content (proteins or DNA)
hominids, to demonstrate that the protein and the age of the fossils or ancient bones. In
216 . ET AL.
many instances, older bones contain more findings are complementary to those of the
biomolecules than younger ones (Ulrich palaeontologist, physical anthropologist and
et al., 1987; Poinar et al., 1996). Fossils archaeologist. At Orce, several lithic assem-
therefore cannot be judged solely by their blages testify to the presence of hominids in
age. Other factors such as average tempera- the lower Pleistocene. Three fragmentary
ture, presence or absence of water and pH bones from Venta Micena (out of the 15,000
all appear to play more important roles. The excavated) are thought by some physical
fossils at Venta Micena were apparently anthropologists to be hominid, by others to
accumulations around hyaena dens, pre- be equid or indeterminate. Our immuno-
served in a lake bed, in a chalky white, logical findings provide molecular evidence
probably alkaline soil. Nevertheless, the that these fossils are probably hominid
‘‘high’’ concentrations of protein recovered remains.
from some of them, referred to by
Palmqvist, are still less than one-hundredth
Acknowledgements
of 1% of the amount present in a fresh bone.
Moreover, the amounts of protein detected This paper has been supported by the
in the fossils from Venta Micena are within project DGICYT PB 94-1222-01 of the
the range of those proteins detected in even Spanish Ministry of Education and Science.
older fossils (Ulrich et al., 1987). We thank Prof. Dr Philip V. Tobias and Drs
The last part of Palmqvist’s (1997) F. Ribot, F. Sanchez, A. Malgosa and M.
criticism is based on a study by Cattaneo Walker for helping us in the discussion of
et al. (1993). These authors reported that the article, and Dr M. Uribe and Dr Valls
proteins did not survive in a bone buried for from UDIAT for performing the tomogra-
a few months, leading Palmqvist to wonder phy. We thank Rosa Campos and K.
how proteins can survive for 1·6 Ma. Kupczik for assistance of the translation.
A fossil bone is a singular specimen that Finally, we thank L. Gibert, who drew
becomes a fossil only under very specific the figures, translated and discussed the
conditions. These conditions are responsible text.
not only for the existence of the fossil, but
also, probably, for the preservation of bio-
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