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Shape dierences in the pelvis of the rabbit, Oryctolagus
cuniculus (L.), and their genetic associations
John Watson, Simon Davis
To cite this version:
John Watson, Simon Davis. Shape dierences in the pelvis of the rabbit, Oryctolagus cuniculus (L.),
and their genetic associations. 2019. <hal-01918838v2>
Shape differences in the pelvis of the rabbit, Oryctolagus cuniculus
(L.), and their genetic associations
J.P.N. Watson1
S.J.M. Davis2
1Chercheur associé, ASM - Archéologie des Sociétés Méditerranéennes, UMR 5140, Université Paul-Valéry, CNRS,
MCC, F-34000, Montpellier, France. Labex ARCHIMEDE programme IA- ANR-11-LABX-0032-01.
2Laboratório de Arqueociências (LARC), Direcção Geral do Património Cultural, Calçada do Mirante à Ajuda 10A,
1300-418 Lisbon, Portugal; e-mail: simonjmdavis@gmail.com
Abstract
Six measurements were taken on the pelvis of modern and archaeological rabbits from France and
the Iberian Peninsula and a difference in shape was noticed between the French and Iberian
populations. A discriminant function was calculated using the geographically most extreme samples
and it appeared that the shape difference was confined to samples from the mitochondrial DNA
(mtDNA) lineage B. Since many of the modern rabbits had been previously analysed genetically, it
was possible to relate the pelvis morphology to the mtDNA type, and it was found that there was an
association between the two. A re-evaluation of the phylogeny using all available mtDNA control
region sequences suggests that the main groups of haplotypes all branched off at much the same
point, presumably as a result of the end of the ice age and the consequent expansion of rabbit
populations both within the Peninsula and out of the Peninsula into France. Two groups (B9 and
B10) are exclusive to the Peninsula and one (B4) is virtually exclusive to France. Another group
(B1) is associated with domestic rabbits. Finally, two subgroups of the mtDNA type B3 are now
seen to be distinct groups, one of them (B3A) being associated with domestic rabbits while the
other (B3C) is only found in French wild rabbits. Previous work in which B3 rabbits were treated as
a single entity therefore needs to be re-evaluated. Whether the association between pelvis shape and
mtDNA type stands up to further investigation or not, their study in archaeological material could
help to elucidate the pattern of dispersal of rabbits after the ice age and the process of their
domestication.
1
Introduction
When the junior author (SJMD) announced that he was going to take measurements on the
long-bones of a large number of rabbits from different parts of western Europe with the aim of
documenting in detail the relationship between winter temperatures and body size first pointed out
by Bergmann (1847) (Davis, in prep.), the senior author (JPNW) persuaded him to take at the same
time a series of six measurements on the innominate bone of the pelvis, or coxal bone (os coxae), in
the hope of being able to discriminate between the sexes. In the pelvis of rabbits, in contrast to other
species of mammals, there are no obvious morphological differences between the sexes because the
young are so small when born.
The analysis of the difference between the sexes was disconcertingly inconclusive, but on the other
hand, to our great surprise, there was a marked difference between rabbits from different
geographical areas. Three large samples from France, two from the north and one from the south,
stood out as being different from those from the Iberian Peninsula.
Materials and Methods
The modern rabbit specimens studied come from the collections of the "Muséum national d'Histoire
naturelle" in Paris (MNHN) and belong to the same animals from France and the Iberian Peninsula
on which the initial genetic analyses on mitochondrial DNA (mtDNA) were carried out (Biju-Duval
et al., 1991; Hardy et al., 1994; Monnerot et al., 1994; Mougel, 1997; Callou, 2003). The material
from the Tunisian island of Zembra was also measured, even though Zembra is not of direct
relevance to the issues considered here. A metrical analysis of many of these animals, especially
their skulls, has already been made by Callou (2003).
The specimens are listed in Appendix 2 and both the number in the MNHN register and the number
allocated by Callou are given, together with the values of the measurements taken and any
information about the sex and age of the animal. Only part of the material had been determined to
sex.
For the present analysis only the localities with more than 10 adult specimens have been used,
although specimens from many other localities from which only a few rabbits had been obtained
were also measured. Exceptions to this are those adult specimens from small samples from France
or Navarre for which genetic information was available. The main samples used are thus those from
Roissy, Versailles, Tour du Valat, Peralta, Tudela, Las Lomas, Badajoz, Vila Viçosa and Santarém.
The smaller samples from Navarre, those from Arróniz, Caparroso, Castejón, Falces, Miranda de
Arga, Ribaforada and Sesma, are all from the south, not far from Peralta and Tudela.
Since the rabbits were wild, precise ages were not available. However, the relative ages of the
innominate bones could be estimated from the state of fusion of the other bones of each individual,
in particular the proximal humerus, proximal tibia and calcaneum. In the main analyses only
specimens in which the proximal humerus was fully fused were included, since that is usually the
last of the three epiphyses mentioned to fuse.
In addition, material from three archaeological sites was studied. The six measurements used had
been devised during the study of the Magdalenian site of Cova Matutano (Watson, 1999; Olària,
2
1999) and so the better preserved specimens from that site were included for comparison; in some
layers many bones were burnt to a greater or lesser degree and so any bone that was discoloured in
a way that might have been due to burning was excluded from the study. Material from two sites in
southern Portugal studied by SJMD was also measured: Silves dates from the 12th-13th century AD
and Beja from the 15th century AD (Davis et al., 2008; Martins et al., 2010).
The measurements taken are described in detail in Appendix 1 (see also Figures 1 and 2). Two
measurements were taken on the shaft of the ischium (H and I) and three on the shaft of the ilium
(E, F and F1). The remaining measurement (J) is between the rim of the acetabulum and the lateral
border of the ischium. These are not measurements that have generally been taken previously, with
the exception of E, which appears in practice to be equivalent to the KB (or SB) of von den Driesch,
and possibly F, which may or may not be equivalent in practice to her KH (or SH) (von den
Driesch, 1976a; von den Driesch, 1976b). They were chosen for their unambiguous definitions and
the precision with which they can be taken, together with their relatively low susceptibility to
damage in archaeological material; as long as the ilium and ischium are not broken apart at the
acetabulum and the bone is not heavily gnawed the measurements can usually all still be taken.
In the modern samples only the left innominate bone was measured. In the archaeological samples
both left and right bones were measured.
Analysis and Results
A preliminary investigation plotting the various measurements against each other showed no
obvious sexual dimorphism. On the other hand, the three French samples (Roissy, Versailles and
Tour du Valat) showed a certain separation from the Iberian samples when F1 or I was plotted
against H (Figure 3).
To extract the maximum information from the measurements a multivariate analysis was then
carried out in two stages, using only the specimens from individuals in which the humerus was fully
fused. First of all, the principal components for the 6 measurements were calculated. The first
principal component was then treated as representing the variation in size. The other 5 principal
components can then be regarded as essentially independent of size, so that any variation in them is
variation in shape (Jolicoeur & Mosimann, 1960). This procedure has been criticised for not taking
into account allometric changes (Sundberg, 1989), and many recent studies have allowed for
allometry (e.g. Callou, 2003), but for the kind of exploratory analysis with small samples attempted
here allometry is irrelevant, especially since the study is based on the adult specimens from a single
species. In the field of zooarchaeology, size and shape components have previously been
distinguished by Kuşatman (1991), Davis (1996, 605) and Burke et al. (2003, 462).
To bring out the shape-difference between the French and Iberian rabbits we therefore calculated a
linear discriminant function based on the 2nd to 6th principal components, using the Roissy and
Versailles samples as one group and the Las Lomas sample as the other group, since they were the
geographically most extreme samples. The first principal component was then plotted against the
discriminant score to give a graph in which the vertical axis represents size and the horizontal axis
represents the shape-difference (Figure 4). The discriminant score in terms of the original variables
was – 31.65 F1 + 9.14 F – 3.81 E – 1.94 J + 35.86 H – 33.57 I + 96.583 and the first principal
component was 0.925 F1 + 0.903 F + 1.015 E + 0.861 J + 0.849 H + 1.180 I - 32.482.
3
Although the two groups are almost completely separated, the separation is based mainly on
size. Nevertheless, there is a considerable difference in mean discriminant score between the two
groups. The coefficients resulting from the calculations were then applied to other samples to see
how they fitted in.
The difference between males and females for Roissy alone is shown in Figure 5, Roissy being the
sample that shows the clearest sexual dimorphism. It can be seen that although there is a size
difference, the males tending to be larger than the females, there is little difference in shape. Figure
6 shows the mean size plotted against the mean discriminant score for the males and females at each
site and it can be seen that the other sites do not follow the Roissy pattern. Not only are the mean
sizes for the females slightly larger than those for the males at some sites, but also the shape differs
in one way at some sites and in the opposite way at other sites, as measured by the discriminant
score.
To investigate the influence of age, the specimens for which the corresponding proximal humerus
was fully fused were compared with those for which it was unfused or incompletely fused. The
proximal humerus was generally the last epiphysis to fuse in the bones that were studied, since the
femur was excluded. Only one animal, which came from Badajoz, had a proximal tibia with the
fusion plane still visible after the proximal humerus had become fully fused. Figure 7 shows the
results of the comparison for the individual specimens from Roissy. It can be seen that in general
there is some difference in size but not a great difference in shape. Figure 8 shows the mean size
plotted against the mean discriminant score for the specimens with fully fused and incompletely
fused humeri from each site. Once again it can be seen that the direction of the difference in shape
varies from site to site but that the difference is not large. Only sites where there were a
considerable number of young animals have been shown in Figure 8, so that several bones have
been left out although they are included in the lists in Appendix 2 for the sake of completeness.
Figure 9 shows, for all the main sites, the mean size plotted against the mean discriminant score for
the specimens for which the corresponding proximal humerus was fully fused. The way in which
the points are distributed suggests that the differences in discriminant score cannot be accounted for
entirely by sex or age differences, particularly since age should not be an important influence in
Figure 9, given that further change with age once the humerus is fully fused must be rather limited.
There is a general trend from south-west Iberian sites on the left to French sites on the right, but two
sites stand out. One is Tour du Valat, which has an even higher score than Roissy or Versailles in
spite of being in a less extreme position geographically. The other is Tudela, which in spite of being
geographically very close to Peralta is even more "Iberian" than Las Lomas.
Figure 10 shows the individual scores for Tour du Valat and Las Lomas, the two southernmost sites
in France and the Iberian Peninsula respectively, with bones of comparable size. It shows that
although there is still a good deal of overlap the general difference in shape is quite marked.
At this point the specimens from the three lowest faunal phases at Cova Matutano were added to
Figure 9, in the expectation that they might fall somewhere above the Peralta or Tudela samples. In
fact they fall even further to the left than Tudela (Figure 11).
It was therefore decided to calculate a new discriminant function using Roissy and the earliest phase
at Matutano as the groups to be discriminated, especially since the sizes of the bones from the two
sites were similar. For good measure, the two other archaeological sites, Beja and Silves, were also
4
added to the data used in the calculation of the principal components. The new discriminant score
was – 91.57 F1 + 24.62 F + 73.93 E – 46.01 J + 90.33 H – 59.91 I + 223.993 and the first principal
component in terms of the raw measurements was 0.755 F1 + 0.885 F + 1.025 E + 0.799 J
+ 0.868 H + 1.173 I - 30.956.
Figure 12 shows the result when the samples used in Figure 11 are plotted using the new
discriminant scores. The improvement in the separation between the French and Iberian sites is
surprising, and Tour du Valat is now more aligned with Roissy and Versailles than before. On the
other hand Tudela is even further to the left and away from Peralta. However, on reflection it occurs
that there may be a good reason for these differences. The initial discriminant function was based
on the comparison of one sample of rabbits belonging to mtDNA lineage A (Las Lomas) with two
samples belonging to mtDNA lineage B (Roissy and Versailles). The new discriminant function is
based on Roissy, belonging to lineage B, and Matutano, a sample that from its geographical location
might be presumed to also belong to lineage B.
The present-day transition zone between lineage B and lineage A lies well to the SW of Matutano
(Figure 13) and any movement of the zone since the end of the last glaciation might be expected to
be towards the north rather than the south. Further analyses have discovered many other genetic
differences between the rabbits on either side of the transition zone and have shown that lineage A
corresponds to the subspecies Oryctolagus cuniculus algirus and lineage B to the subspecies
Oryctolagus cuniculus cuniculus. Although there is strong gene flow and hybridisation within the
zone there are also strong genetic barriers to reproduction that mean that the zone must have existed
for a long time and that a process of speciation has begun (Carneiro et al., 2013; Carneiro et al.,
2014).
Figure 14 shows the individual specimens from Roissy and Matutano at the same scale as in Figure
12. There is some overlap, but not a great deal. Figure 15 compares the individual specimens from
Peralta and Tudela. The Tudela samples are widely scattered and so might represent a mixture of
two types, one like Peralta and another with a more “Iberian” shape.
Figure 16 is the same as Figure 12 but with with Beja, Silves and Zembra added. The samples from
Beja and Silves, which presumably belong to lineage A, fit in well with the modern members of that
group. The sample from Zembra also fits in well, but is known to belong to lineage B. However,
when considered as merely rather small members of lineage B, their position below Peralta fits in
well with the origin postulated for the animals with which the island of Zembra was originally
populated. They seem most likely to come from somewhere on the east coast of the Iberian
Peninsula where their mtDNA type B6 is also found at the present day (Branco et al., 2000, Table
2). Analyses on archaeological material implied that the modern rabbits from Zembra were
descendants of those there 1400 years ago (Hardy et al., 1995), with no sign of any subsequent
introduction of new stock.
Finally, Figure 17 shows the effect of adding a few specimens of two breeds of domestic rabbits
from the MNHN collection. Since they are so large compared with the wild rabbits, the vertical
scale has had to be compressed so that they would fit, but otherwise the rest of the figure is identical
to Figure 12, except that the range shown on the horizontal axis has been reduced, so that the mean
discriminant scores appear more spread out than in Figure 12. What is interesting is that the
domestic rabbits fall on the right of the graph, above Roissy and Versailles. This fits perfectly with
the postulated French origin for all breeds of domestic rabbit, which mainly show the mtDNA type
5
B1 that is common in northern French wild rabbit populations, which derive from from the
Mediaeval expansion of the range of the species when the keeping of rabbits in warrens became
popular (Queney et al., 2002; Hardy et al., 1995).
Genetic Associations
The reasons for the difference in shape would appear to be genetic. Other possible explanations
seem unlikely; it is clearly not related to temperature or other climatic factors and there is no
obvious relationship to other features of the environment, such as the terrain.
As far as the results of the discriminant analysis are concerned, it seems that the large differences in
shape that have been found affect only lineage B, that is to say only the subspecies Oryctolagus
cuniculus cuniculus. Although shape differentiates lineage A from some of the lineage B samples,
the variation in shape is essentially a phenomenon confined to lineage B.
The lineage A samples are remarkably similar to each other in spite of the differences in their
chronological and geographical origins and are in fact clustered more closely together in the new
analysis than in the original one. On this basis, the Beja and Silves samples, in spite of their date,
are likely to be just local wild rabbits belonging to lineage A and not domesticated animals
belonging to lineage B, with the possible exception of one specimen from Beja.
Given that lineage A and lineage B have been separate for such a long time, it is not surprising that
there should be differences that affect one subspecies but not the other. What is surprising is the
lack of correlation between obvious physical differences and major genetic differences. The two
subspecies are so similar morphologically that it was only recently that the identity between the
subspecies and the genetic lineages was firmly established (Geraldes et al., 2006). Size had long
been used as a criterion, but it turned out that some of the smallest rabbits, those from Zembra, were
from lineage B (Hardy et al., 1994; Callou, 2003, 23) Now it seems that in the same way a
morphological difference in the pelvis has nothing to do with the difference between subspecies.
There is no reason, of course, to suppose that it is differences in the mtDNA itself that are
responsible for the differences in shape, but at present there is very little other genetic information
that can be related to them. Other kinds of genetic material have been studied and the types that
were found have been related to their geographical origins, but although in some cases samples
were from the same localities as the animals studied here, no direct link can be made between pelvis
shape and genetic type. A link can only be made indirectly by association with the same
geographical area, but even when two samples come from the same locality their composition may
differ. In general the material studied has not been sufficient to show clear differences except
between the A and B zones (Geraldes et al., 2006; Geraldes et al., 2008; Carneiro et al., 2010;
Carneiro et al., 2014). The modern rabbit material from the MNHN was collected as part of a
project that studied the variation both in the mtDNA and in microsatellites (Queney et al., 2001).
Although the microsatellite results show differences between broad regions, such as south-western
and south-eastern France, there is for example no difference between Tudela and Peralta in their
microsatellite alleles.
The disadvantage of mtDNA is that since it is essentially transmitted only through the maternal line
it may not be at all representative of the population to which an animal belongs and may have little
6
relation to the rest of its genetic makeup and thus to the factors affecting pelvis shape, supposing
that they are not determined by the mtDNA.
In spite of this, the mtDNA is still likely to be the most informative subject for analysis. The first
reason is that in rabbits, as in many species, the females tend to move around less than the males, so
that on the whole the mtDNA is likely to be representative of the population in which it is found.
The second reason is that in this particular case the limitations imposed by the climate and thus by
the barriers created by mountain ranges, especially the Pyrenees, are important in determining the
pattern of expansion of rabbit populations during the last stages of the glaciation. Under natural
conditions the area of distribution of the rabbit seems to be limited by the need for the mean January
temperature to be above 1 to 2 degrees Centigrade. The females are likely to remain within this
zone. The males may well wander further afield in warmer months and thereby cross a mountain
range, but if they then breed with females from another population their mtDNA will not be
transmitted. This means that the Pyrenees are an effective barrier to the spread of mtDNA at the
present day except at the western and eastern ends.
In rabbits it is the young females for whom there is no room at the parental warren that move away
to found new colonies (Lockley, 1965, 134-135). The mtDNA is thus the best indicator of the
pattern of evolution and spread of rabbit populations in the short term, although in the longer term
older-established populations that suffer a setback, for example as a result of a colder climate, will
tend to be replaced by populations with more recently evolved mtDNA types and the previous
pattern will become obscured.
Through the kindness of Florence Mougel in giving us data from her thesis (Mougel, 1997) and
from an unpublished paper (Mougel et al., unpublished; Mougel, pers. comm.), we have been able
to relate many of the specimens measured to their mtDNA types.
In the initial studies of mtDNA polymorphism in rabbits (Biju-Duval et al., 1991; Monnerot et al.,
1994), three specimens from Versailles were classified as being of type B1, as were the two
specimens from Donzère and three of the specimens from Arzujanx. Unfortunately only one of the
measured specimens from Versailles can be related to its type, which is indeed B1, but on the other
hand the Roissy specimens can be identified to type and it is not surprising to find that most of them
are of type B1. One adult specimen from Roissy belongs to type B3 and two more are either B3 or
B4. The young specimens from Roissy are also all B1 or B3-B4. The remaining specimens from
Arzujanx belong to type B3.
The Tour du Valat sample was established by Monnerot et al. (1994) as being all of type B4, except
for one that was B5, and it is now clear that all the measured specimens are B4. The French rabbits
would thus all be B1, B3 or B4 and the “French” morphology would seem to be associated with
these types.
However, in Navarre three other types were found by Biju-Duval et al. (1991), coming from Sesma,
Arróniz and Tudela. Monnerot et al. (1994) subsequently named them B8, B9 and B10 respectively,
also adding a new type, B11, that they identified from Castejón. The “Iberian” morphology might
therefore be expected to be associated with these types. The problem was that 24 specimens from
Tudela had been measured but only 14 had been typed by Monnerot et al. (1994), to types B9 and
B10, while no information was available about the types present in the other large sample from
Navarre, the one from Peralta.
7
Thus the marked difference in average shape between the specimens from Tudela and Peralta
(Figure 15) could not easily be explained. A subsequent study by Branco et al. (2000) of different
material from Tudela and Peralta gave a rather different pattern of types. Types B8, B9 and B10
could not be distinguished from each other but accounted for only about half the types from Peralta
and three-quarters of the ones from Tudela, the remaining types including B1, B3 and B4 as well as
some new ones.
Fortunately, Mougel's data provide the answer. All the more extreme Tudela specimens are of type
B10 (Figure 18), while B10 is absent from Peralta and the other samples measured from Navarre,
which are mainly of types B9 and B3.
Examination of the scatter of points for the adult specimens for which the mtDNA type has been
determined shows that there is no noticeable shape difference between the B1 and B4 rabbit pelves.
Nor is there any noticeable difference between B3 and B9. Thus there are essentially three groups,
B1 and B4 on the right, B3 and B9 in the centre and B10 on the left; B6, represented by the sample
from Zembra, can be added to the central group (Figures 19 & 20). Unfortunately, B11 was not
found in any of the specimens measured. For what it is worth, the single B8 specimen measured, the
type-specimen from Sesma, falls quite far to the left, nearer to the centre of the B10 cluster than to
that of any other (Figure 20). Figure 20 shows that there is not a great deal of overlap between B1
and B4 on the right and B10 on the left. (The extreme outlier from Roissy shown in Figures 4, 5, 7
and 14 does not appear since it was not identified to type and so may not be B1, or even B3 or B4.)
The differences in discriminant score between the various types were tested with the Mann-Whitney
test (two-tailed). The difference between B10 and all the other types was significant at the 98%
level or more. With the exception of B3, each member of the central group was found to be
different from each member of the group on the right at a significance level of 95% or more. Type
B3 was probably not distinguishable from the members of the group on the right because the sample
was so small. Table 1 shows the probability levels for the differences between the various types.
The original types defined by Monnerot et al. (1994) were based on RFLP (restriction fragment
length polymorphism) analysis of the complete mtDNA genome. That is to say, restriction enzymes
that cut DNA in specific places are used to find differences between the DNA sequences of two
individuals. The differences are manifested as differences in the lengths of the DNA fragments
produced. Monnerot et al. (1994) tested for differences at 110 sites along the length of the genome,
but only 17 of these showed variation within the B-lineage and so provided the basis for the B
mtDNA types.
However, Mougel (1997) made a more detailed analysis and defined types on the basis of
sequencing part of the control region of the mtDNA and so being able to compare every nucleotide
site in all the individuals studied. Thus the possible number of differences was much higher and is
reflected in the large number of types identified. Within the B-lineage there were four times as
many sites that varied as there were in the RFLP analysis by Monnerot et al. (1994). At the same
time, many of these control region types could be related to the types defined previously since the
same individuals had been studied. Thus, for example, the sequence found in those specimens
classified by RFLP as B9 was called B9zsa and the three different sequences found in the
specimens classified as B3 were called B3Zsa, B3Zsb and B3Zsc respectively. The ten control
region types that could not be related to an established RFLP type were given numbers prefixed
with “Bzs”and were thus called Bzs1 to Bzs10.
8
In Mougel's thesis, a "z" was used to indicate a sequence with a length of around 200 nucleotides
and a "Z" to indicate one with a length of around 400. Most of the specimens were subsequently
re-sequenced and lengths of more than 300 obtained, so that the difference is no longer relevant. In
general a "z" has since been used for sequences of all lengths, both in publications (Branco et al.,
2002; Letty et al., 2007 ) and in the NCBI database (http://www.ncbi.nlm.nih.gov), but some types
in the NCBI database are still written with a "Z". In the present paper "z" will be used throughout
from now on.
When a longer sequence was obtained as a result of resequencing a specimen, in some cases further
differences were then found, so that the specimen had to be reclassified; for example, specimen RI5
(or Ri05) from Ribaforada was originally classified as B9zsa but later assigned to a new type,
Bzs53 (Mougel et al., unpublished; Mougel, pers. comm.).
As far as relating the measurements on the pelvis to these other types is concerned, single
specimens of various other types from Navarre (Bzs3, Bzs5, Bzs7 and Bzs10) plot out close to the
B8zsa specimen, while 3 specimens of a further type, Bzs4, which is very close to B9zsa, are
scattered over the central area amongst the B3 and B9 individuals.
The biggest limitation to associating the shape differences with the mtDNA types is that several of
the types are only recorded from one locality or almost entirely from one locality. B10zsa is only
recorded from Tudela, although 3 other animals assigned to B10 through their cytochrome-b were
found at the nearby locality of Caparroso. B4zsa is almost entirely from Tour du Valat. As far as the
measured pelves are concerned, the only other specimens recorded as belonging to a B4 type are
two B4zsa from Miranda de Arga and one from Caparroso, both in Navarre. However, the
specimens from Miranda de Arga were subsequently reclassified as Bzs55 (Mougel et al.,
unpublished; Mougel, pers. comm.), which is considerably closer to B9zsa than to B4zsa.
Unfortunately all three specimens are from young animals.
In order to understand better the relationships between the various control region types and the way
in which they might have evolved, a tentative evolutionary tree has been constructed on the basis of
the similarity of the types to each other. Initially it was thought more appropriate to compare
populations (Mougel, 1997; Queney et al., 2001), but it has subsequently become evident that the
probable introduction of rabbits of possible domestic ancestry brought from elsewhere into some
populations confuses the results obtained. Grouping the rabbits by type and then studying the
distribution of those types amongst the various populations gives better results.
In this particular case an evolutionary tree seems appropriate for various reasons. Firstly, mtDNA is
simpler because it is single-locus and essentially non-recombining. Secondly, the types being
compared are much more closely related than in most analyses, belonging not only to the same
species but also to the same subspecies. Thirdly, the time period involved is short, most of the types
presumably having diverged since the last glacial maximum some 25000 to 30000 years ago.
Fourthly, since the control region, or at least large parts of it, does not have to code for amino acids,
it is able to mutate freely, so that in practice the evolution seems to have followed a branching
pattern rather than creating a network. The various branches generally unite sequences of similar
geographical origins. Only in one case does it seem necessary to invoke convergent evolution.
The other part of the mtDNA genome for which much information is available is the cytochrome-b
region. However, the ability of the this region to mutate freely is restricted by its need to
9
maintain functionality. The sequences, presumably as a consequence, tend to be related in a
network with side-branches rather than a freely branching tree.
The sequences from the control region and the cytochrome-b are available from the NCBI database
at http://www.ncbi.nlm.nih.gov and come from various sources (Van der Loo et al., 1997; Bolet et
al., 2000; Letty et al., unpublished; Mougel et al., unpublished; Queney et al., 2002; Queney et al.,
unpublished; Long et al., 2003; Zenger & Richardson, unpublished; Seixas et al., 2014; Emam et
al., unpublished). Four additional control region sequences from Table 1 in Branco et al. (2002)
were added to the analysis and although they are rather shorter sequences than the others they fit in
well and make good sense, one of them being a possible intermediate.
The fact that the control region sequences give much more information about the genetic variability
than the RFLP analysis of Monnerot et al. (1994) means that the original classification is now
outdated as a guide to the phylogeny of the rabbit and indeed it is not even mentioned by Seixas et
al. (2014). However, the broad outline remains similar.
Monnerot et al. (1994) showed B8, B9 and B10 as the closest types to the A-lineage, with two main
branches subsequently forming, one with B3 and B4 and the other with B1, B6 and B11. The new
phylogenetic tree for the control region (Figure 21) shows three main branches forming at about the
same time, one to B3 and B4, one to B8, B9 and B10 and one to B1, B6 and B11. The network
shown by the cytochrome-b types is more like the original scheme, showing a branch to B8 and B9
at an early stage, then B3 and B4 close together in a network and finally a long branch to B1.
In the evolution of the control region shown in Figure 21, the main line coming from the A-lineage
shows two early side-branches, to Bzs43 and Bzs30, and then a third branch to Bzs3 and Bzs29,
before reaching an area in which it splits into three main lines. One of the lines leads to B8zsa,
B9zsa, B10zsa and other similar types. Another line leads to B3zsa, B3zsc, B4zsa and related types.
The third line leads to B1zsc, B6zsa and B11zsa. The most economical interpretation has the branch
to B1, B6 and B11 going off before the branch to B8, B9 and B10 separates from the branch going
to B3 and B4. It thus differs slightly from the pattern shown by the RFLP and the cytochrome-b
sequences.
Of the Bzs types found in the specimens measured, types Bzs4 and Bzs53 are close to B9zsa in this
new evolutionary tree. Bzs5 and Bzs55 are fairly close to B8zsa and B9zsa, while Bzs7 may have
branched off slightly earlier. Bsz10 is very close to B11zsa.
The evolutionary tree in Figure 21 has been constructed using trial-and-error and common sense,
following the principle of Occam's Razor, since the control region has its own patterns of evolution
(Meyer et al., 1999; Howell et al., 2007; Endicott et al., 2009; Mignotte et al., 1990) and the
commonly used computer programs make assumptions that are not necessarily valid. In particular,
insertions and deletions are treated in the same way as substitutions, with the result that in this case
the trees produced by computer programs often showed an insertion that was almost immediately
deleted again, a highly implausible event. In any case, the methodology employed is itself very
much open to debate (Farris, 1983; Kluge 1997; Haber, 2005). In the construction of the tree shown
in Figure 21, biogeographical considerations have been taken into account as well as purely
phylogenetic ones. In addition it has been possible to use a limited amount of information from the
cytochrome-b sequences, since occasionally the two kinds of sequencing were carried out on the
same specimen. This has allowed a possible convergent evolution of Bzs27 and Bzs29 to be
detected (see Note 1).
10
For comparison, various parsimony analyses were carried out using version 3.696 of the program
dnapars from the PHYLIP phylogeny inference package (Felsenstein, 1989;
http://evolution.gs.washington.edu/phylip.html). Only the 314 sites shared by the majority of the
control region sequences published in the NCBI database were used, corresponding to positions
15516 to 15827 in the complete mtDNA sequence published by Gissi et al. (1998), and so 111
distinct sequences were analysed. A large number of equally parsimonious trees were produced, one
of which is shown in Figure 22. The overall parsimony score (including the outgroup) is in fact only
slightly better than that for our own tree using the same sites, being 282 as opposed to 287.
The pattern shown by the main branches is much the same in all the trees and the large number of
trees produced is the consequence of uncertainty in the positions at which they branch off. This in
turn is the result of a concentration of the mutations in only a small number of sites. In the complete
tree for the B-lineage from which Figure 21 is taken, nearly a third of the changes (77 out of 240)
take place at only 10 of the sites. In the parsimony analyses the concentration shown was much the
same, the tree shown in Figure 22 having 75 of the 232 changes occurring at only 10 sites.
The principal sites are always the same, so it seems that there are a number of genuine mutational
hot-spots (Stoneking, 2000; Galtier et al., 2006). These hot-spots appear to be related to the
secondary structure of the DNA (Pereira et al., 2008; Damas et al., 2012). No less than four of these
hot spots are involved in the main branching area, so that both independent identical mutations at
the same site and reverse mutations are not improbable. Consequently in that area the most
economical tree is not necessarily the correct one. If one or two mutations occurred on more than
one occasion or happened to be subsequently reversed, the branching pattern could easily come into
line with that produced by the RFLP and cytochrome-b analyses.
Figure 23 shows a modified version of Figure 21 in which two mutations are duplicated in such a
way as to bring the branching pattern into line with those shown by the RFLP and cytochrome-b
analyses. The new parsimony score is 289.
The main branches in Figures 21, 22 and 23 are marked with the names of the corresponding RFLP
types. However, it is evident that B3 includes three different groups, which have been labelled B3A,
B3B and B3C in Figure 22, and that B4 is likely to correspond to at least two groups, since a
comparison of the geographical distributions of the cytochrome-b and control region types in the
unpublished study by Mougel et al. shows that at least some of the types in the group that includes
Bsz40 must also belong to B4 (Mougel et al., unpublished; Mougel, pers. comm.).
Many of the types in the group that includes B3zsa are found in domestic rabbits, as are all the types
in the B3zsb group. On the other hand, none of the types in the group that includes B3zsc are
definitely domestic, although they are strongly associated with the spread of rabbits across northern
France, which suggests that they might reflect the Mediaeval expansion of rabbits kept in warrens
without being deliberately domesticated. It is interesting that the only B3 specimens recorded from
the Peninsula are of type B3zsa (Mougel, 1997; Branco, 2002), which suggests that they are likely
to be descended from introduced domestic stock.
The archaeological evidence from early sites in southern France that date from the Azilian to the
Neolithic (Hardy et al. 1995) shows only cytochrome-b type B4sa, apart from a single example of
type B4sb. The type could possibly be B3sa rather than B4sa, since footnote “c” to Table 2 says
“mtDNA types B3sa and B4sa are distinguishable by RFLP in modern samples but not in the
11
cytochrome-b gene sequence used for ancient samples”; however, as we have shown, the
corresponding control region B3zsa group is associated with modern domestic animals, so that the
type is much more likely to be B4sa. Later sites that date to the Bronze Age and Gallo-Roman
periods show almost entirely cytochrome-b type B3sc (equivalent to the control region B3zsc
group). This suggests that the two B4 groups of control region types and the group including
B3zsc might be the original groups that dominated in southern France prior to domestication (see
also Callou, 2003, 243).
The precise area of the refugium, or perhaps refugia, for the B-lineage rabbits during the last
glaciation is not at all clear. Although there is archaeological evidence for rabbits from quite far
back into the glaciation, it is not until the final stages of the main glaciation that there is a rapid
expansion of their range and clear evidence for their exploitation by humans, as for example at
Cova Matutano, where a high proportion of the bones show cut-marks. However, even after that,
the Younger Dryas cold event would have been a severe setback for rabbits and have affected the
evolution of their populations in the recently colonised areas.
The Pyrenees would have remained a barrier during this initial expansion and essentially still are
one at the present day. At the eastern end, the Cerdagne is the westernmost place on the crest of the
range at which rabbits are found. At the western end, the range is continuous with the Cantabrian
mountains and it would seem from the archaeological evidence that rabbits did not reach the north
coast of the Peninsula until well into the Holocene.
Thus any genetic differences between the rabbits that spread across France and those that spread up
the Ebro valley to Navarre would be perpetuated and become established over wide areas. On the
one hand, B9zsa, B10zsa and types close to them are widespread on the south side of the Pyrenees
but have not been recorded from France. On the other hand, B4zsa, Bzs40 and types close to them
are widespread in southern France but there is only one specimen recorded from south of the
Pyrenees. Similarly, B3zsc and types close it to have only been found north of the Pyrenees.
The distribution of the B1 types and also of B3zsa and types close to it cannot be relied upon as a
guide to the origin of those types since they are types found in domestic rabbits and are likely to
have been spread by human activity.
Discussion and Conclusions
It has been shown that there are differences in shape in the pelvis of rabbits from France and the
Iberian Peninsula that are not merely due to differences in sex and age. These differences are
associated with differences in the type of mtDNA. Unfortunately the association of type with shape
is sometimes confined to only one or two populations and so may be due to chance, at least in part.
However, if the associations are found to hold up when further samples are examined, measurement
of the shape difference could be a useful way of tracing the evolution of archaeological populations.
It could also be useful for detecting the presence of animals introduced into France from the Iberian
Peninsula and vice versa. Such introductions would in turn presumably imply domestication or the
processes leading to it.
More research is therefore urgently needed into the morphology of the pelvis of both ancient and
modern rabbits in southern France and the Iberian Peninsula and then into their genetic affinities.
12
The eastern end of the Pyrenees and adjacent areas are of particular interest, since it is there that the
divergence of different evolutionary branches must have occurred towards the end of the last
glaciation. Such modern material as we have been able to examine from the south side of the
eastern end of the Pyrenees shows the same extreme "Iberian" pelvis shape as the B10zsa
specimens from Tudela or the samples from Matutano. The nature and position of the transition
zone to the "French" shape remains to be established.
The extreme "French" shape of the B4zsa rabbits from Tour du Valat suggests that the shape
difference might not be just a consequence of the spread of rabbits across France in Mediaeval
times or of the processes leading to domestication, but may be a characteristic of populations from
south-eastern France.
It would be particularly interesting to see whether a consistent shape difference exists in France,
both in modern and in archaeological material, between individuals belonging to the B4 and B3zsc
groups of types, those being apparently the types occurring in wild French rabbits prior to
domestication. Unfortunately only two of the specimens measured can be assigned to B3zsc and
although one of them plots out amongst the B4zsa specimens the other plots out far out on the left.
The B1 types, apart from B1zsc, and those in the B3zsa and B3zsb groups appear to be associated
with deliberate breeding of domestic rabbits, so that individuals possessing them seem unlikely to
be of entirely wild ancestry, at least in most cases. The origin of these types is therefore of great
interest. B1zsc has only been recorded from England and since it is the earliest B1 type may well
have arrived there in Mediaeval times with the spread of the fashion for creating warrens (Veale,
1957).
There have been suggestions that the B1 group may have originated in the Iberian Peninsula
(Loreille et al., 1997; Callou, 2003), but unfortunately the published documentation is sparse and
the analyses were of the cytochrome-b rather than the control region. In retrospect it seems unlikely
that B1 should be found at a prehistoric site in the south of the Peninsula that is well within the zone
of the A lineage, so the suspicion must be that at least in this case there was an intrusion of more
recent material of domestic origin.
Finally, the results of the analysis made in the present study show that there is still room in
zooarchaeology for measurements of the traditional kind. In recent years, geometric morphometrics
has become fashionable, but conventional measurement has stagnated because of the almost
universal adoption of the measurements proposed by von den Driesch (1976a, 1976b) and the
associated obsession with orientation in certain fixed directions. This insistence on orientation was
derived from Duerst (1926), who was concerned with comparative anatomy, but zooarchaeology
has very different aims from comparative anatomy. Apart from giving a general idea of size,
measurements in zooarchaeology serve to find small differences between closely related taxa or
between different sexes. Orientation is irrelevant; precision and repeatability are the essentials.
In zooarchaeology, multivariate analysis has been little used except on the skull because there are
usually too few well-defined measurements, and the skull suffers from the drawback that it is rarely
found sufficiently intact in sufficient numbers on archaeological sites. Measurements on the
postcranial skeleton have generally been too few in number, as well as too poorly defined, to
provide enough precision.
13
Geometric morphometrics has played a positive part in so far as it has disrupted the status quo and
stimulated an interest in a multivariate approach, but it is not without its own difficulties and
limitations. Above all, precise and repeatable positioning of the landmarks is essential and not
always easy in archaeological material. It would be as well to exhaust the possibilities of traditional
measurements first.
There is thus a good deal of scope for the application of the methods presented here, both in
archaeological material and in modern wild and domestic rabbits. Although genetics will inevitably
have the last word in tracing the evolution and spread of the species and of the various domestic
breeds, the role of this traditional metrical approach in combination with multivariate analysis could
be that of finding the most profitable material for investigation.
14
Note 1
It is possible to associate some of the cytochrome-b types with control region types since in some
cases both kinds of sequencing were carried out on the same specimen. The main consequence of
this, as far as the present argument is concerned, is that a group of closely related control region
types seem to belong to two different lines that have converged, the only instance in which
convergent evolution needs to be invoked. One of these types, Bzs27, is from a specimen that has
cytochrome-b type Bpbs14, which in turn can be classified as B3-4 from the way it is affected by
the restriction enzyme AluI. Another type, Bzs29, is from a specimen that has cytochrome-b type
Bpbs15, which in the same way can be classified as not belonging to B3-4. However, the restriction
enzyme EcoR1 classifies both specimens as not being B8-10. This leaves the second specimen with
the possibility of belonging to B1 or B11, but neither the cytochrome-b nor the control region
sequence resembles the ones for these types. The conclusion must be that it belongs to an unknown
RFLP type, presumably earlier in the evolutionary tree, and the control region sequence Bzs29 does
indeed fit in well as part of an early side branch. Bzs3 is close to Bzs29 and since it is only found in
Navarre is more likely to belong with it than to B3-4. The remaining types, Bzs18 and Bzs28, are
on the same side-branch as Bzs27 and so have been provisionally treated as most likely to belong to
B3-4.
15
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Van der Loo, W., Mougel, F., Sánchez, M. S., Bouton C., Castien E., Soriguer, R., Hamers, R. &
Monnerot, M. (1997). Evolutionary patterns at the antibody constant region in rabbit (Oryctolagus
cuniculus L.): Characterisation of endemic b-locus allotypes and their frequency correlations with
major mitochondrial gene types in Spain. Gibier Faune Sauvage 14 (3), 427-449.
Veale, E.M. (1957). The Rabbit in England. Agricultural History Review 5, 85-90.
Watson, J.P.N. (1999). Análisis Preliminar de los Mamíferos del Sector 3 de Cova Matutano
(Vilafamés, Castellón). Sant Cugat del Vallès: Arpali.
Zenger, K.R. & Richardson, B.J. (unpublished). Genetic variation and evolution of a colonizing
species: the mitochondrial DNA control region in the European rabbit (Oryctolagus cuniculus) in
Australia.
20
Figures:
Figure 1. Drawing showing the approximate positions in which the measurements H, J and F1 are taken.
Figure 2. Drawing showing the approximate position in which the measurement F is taken.
21
Figure 3. The result of plotting F1 against H. The grey circles represent Las Lomas, Badajoz, Vila Viçosa and
Santarém, the sites from the south-west of the Iberian Peninsula. The black crosses represent Peralta and Tudela, the
sites from Navarre. The black triangles represent the French sites of Roissy, Versailles and Tour du Valat.
Figure 4. The first principal component plotted against the discriminant score obtained by using Roissy and Versailles
as one of the groups in the discriminant analysis and Las Lomas as the other group. The individual specimens for the
three sites are plotted, those from Las Lomas as grey circles, those from Roissy as black triangles and those from
Versailles as black inverted triangles.
22
Figure 5. The first principal component plotted against the discriminant score for the Roissy specimens that had been
identified to sex. Males are represented by black triangles and females by grey ones. It can be seen that although there is
a certain difference between the sexes, it is mainly a difference in size. The difference in discriminant score is small
compared to the difference between the scores for the Roissy specimens as a whole and those from Las Lomas. The
isolated male on the extreme left has had its measurements retaken and is a genuine outlier.
Figure 6. The means of the first principal components plotted against the means of the discriminant scores for the
males and females for each site.
23
Figure 7. The first principal component plotted against the discriminant score for Roissy specimens of different ages.
The black triangles represent those specimens for which the corresponding proximal humerus is fully fused, the dark
grey circles represent those for which its fusion plane is still visible and the light grey squares represent those for which
it is still unfused.
Figure 8. The means of the first principal components plotted against the means of the discriminant scores for the
specimens for which the corresponding proximal humerus is fully fused (in black) and those for which it is
incompletely fused or unfused (in grey).
24
Figure 9. The means of the first principal components plotted against the means of the discriminant scores for the
adults from the same 9 sites as in Figure 3.
Figure 10. The first principal component plotted against the discriminant score for the individual adult specimens from
Tour du Valat (black inverted triangles) and Las Lomas (grey circles).
25
Figure 11. The means of the first principal components plotted against the means of the discriminant scores for the
adults from the 9 modern samples, as in Figure 9, together with those for the specimens from faunal phases A, B and C
at Cova Matutano.
Figure 12. The means of the first principal components plotted against the means of the discriminant scores for the same
specimens as in Figure 11, but with a new discriminant function obtained by using Roissy as one of the groups in the
discriminant analysis and Matutano A as the other. In addition, the specimens from Matutano, Beja and Silves are now
included in the calculation of the principal components.
26
Figure 13. A map showing the locations of the sites, together with the transition zone (in grey) between the lineages A
and B (after Branco et al., 2002 and Geraldes et al., 2008).
Figure 14. The first principal component plotted against the discriminant score for the individual specimens from
Matutano A, B and C (grey squares) and the adult specimens from Roissy (black triangles), using the new principal
components and discriminant function.
27
Figure 15. The first principal component plotted against the discriminant score for the individual adult specimens from
Tudela (grey crosses) and Peralta (black crosses), using the new principal components and discriminant function.
Figure 16. The means of the first principal components plotted against the means of the discriminant scores for the same
specimens as in Figure 12, but with the archaeological sites of Beja and Silves added, together with the adult specimens
from a modern sample from the Tunisian island of Zembra.
28
Figure 17. The means of the first principal components plotted against the means of the discriminant scores for the same
specimens as in Figure 12, with the addition of the means for a few domestic specimens of the Gris de Vienne and
Papillon breeds. The range on the horizontal axis has been reduced, so that the mean discriminant scores appear more
spread out than in Figure 12.
Figure 18. Tudela and Peralta as in Figure 15, but with the type B10 specimens from Tudela indicated by the addition
of a circle.
29
Figure 19. Specimens with mtDNA types B3 (black triangles), B6 (grey inverted triangles) and B9 (black crosses).
Figure 20. Specimens with mtDNA types B1 (black circles), B4 (black inverted triangles) and B10 (grey crosses within
a circle). The B4 specimens are all from Tour du Valat and the B10 ones are all from Tudela. The single B8 specimen is
also shown and is indicated by a black square.
30
Figure 21. Proposed phylogeny of the main mtDNA control region types discussed. In order to keep the diagram as
simple as possible, most types that are not referred to in the text, or that are not intermediate types on branches to types
referred to in the text, have been excluded. Postulated intermediate types that have not so far been recorded are marked
by black dots. A bar across a black dot indicates the existence of a side-branch at that point. The evolutionary
development begins on the left, where there are 55 differences between the closest member of the A lineage and the
start of the branch leading to Bzs43. The symbols in large print are the corresponding RFLP types of Monnerot et al.
(1994). The number given to each link is the position of the difference between the two sequences, relative to the
beginning of the longest sequence used in the matching. Since the complete mtDNA sequence of Gissi et al. (1998)
differs from the sequences compared here in an insertion and up to three deletions, the following values need to be
added in order to obtain the positions in that sequence: 15395 to positions 1 to 126; 15394 to positions 128 to 282;
15392 to positions 285 to 296; 15393 to positions 297 to 571. The positions 283 and 284 shown here do not exist in the
sequence of Gissi et al. (1998) and the links constitute insertions. The order of the numbers in a series of links between
branching points is of course arbitrary. All detectable mutations between positions 1 and 571 have been shown, but
since the sequences are of different lengths there may be other mutations that could not be detected; thus it is possible
that a type might be on a short side-branch from the position it is shown at, rather than at the position itself.
31
Figure 22. A tree produced by the PHYLIP program dnapars (Felsenstein, 1989) from the 111 sequences in the NCBI
database that share positions 15516 to 15827 in the complete mtDNA sequence published by Gissi et al. (1998). The
haplotype A02zsa from Lineage A has been used as the outgroup. On the right the names of the RFLP types are shown
opposite the branches that include the corresponding haplotype, except that B3 is split into subtypes B3A, B3B and
B3C since the haplotypes B03zsa, B03zsb and B03zsc are distributed across three different branches. The tree has been
plotted with SeaView (Gouy et al. 2010). 32
Figure 23. A modified version of Figure 21 that brings the main branching pattern into line with that produced by the
RFLP and cytochrome-b analyses. The changes at positions 165 and 404 are duplicated in such a way that the the main
branch to B8, B9 and B10 goes off before the B3 and B4 branch separates from the B1, B6 and B11 branch. The
reversal of the change at position 404 in the branch to Bzs7 is no longer needed and the anomalous situation of B3zsb is
eliminated with the result that all the B3 and B4 types are now on the same main branch.
33
B3 B4 B6 B9 B10 n
B1 >0.1 >0.1 <0.05 <0.02 <0.002 21
B3 - >0.1 >0.1 >0.1 <0.02 8
B4 - - 0.02 <0.02 <0.002 18
B6 - - - >0.1 0.002 12
B9 - - - - <0.02 14
B10 - - - - - 13
n 8 18 12 14 13
Table 1. Probabilities of obtaining differences between types as large as those found, according to the Mann-Whitney
U test, on the assumption that there had been in fact no difference between the populations. The tables used are in
Siegel (1956) and Mann & Whitney (1947).
34
Appendix 1:
Definitions of the measurements taken.
Figures 1 and 2 show the approximate positions in which the measurements are taken. All the
measurements are taken with a vernier gauge with "knife-edge" blades (i.e. they should have
reasonably narrow edges, although that is only really important for F1).
The formal definitions are as follows:
H = minimax diameter of ischial shaft, between the acetabulum and the spina ischiadica.
I = absolute minimum diameter of ischial shaft (usually in the same place as H, but sometimes
much more caudal).
J = minimum height of the lateral side of the acetabulum, i.e. the minimax. This is only just possible
in lagomorphs, one of the blades of the vernier gauge lying almost flat against the back of the
acetabulum (i.e. the medial side). It also needs a certain delicacy of touch because the edge of the
acetabulum is easily damaged. In addition the acetabulum needs to be fully fused.
E = absolute minimum diameter of the shaft of the ilium. This is very oblique in rabbit, about 45
degrees to the axis of the ilium. It should not enter the area of fusion with the sacrum (which it very
occasionally tends to do).
F = minimax diameter of the shaft of the ilium, usually the absolute minimax but sometimes a local
minimax. The measurement is near to or equivalent to the dorso-ventral height in most cases (the
KH or SH of von den Driesch), but taken as a minimax. However, in some cases, when the spina
iliaca dorsalis caudalis extends a long way caudally, this is only a local minimax and the absolute
minimax is further round, more or less parallel to F1. Sometimes it is difficult to take because the
depression on the ventral side of the spina iliaca ventralis caudalis is very shallow and the bone
then has to be placed right between the tips of the blades of the vernier gauge in order to find the
minimax.
F1 = minimax diameter of the shaft of the ilium with one blade of the vernier gauge placed between
the edge of the acetabulum and the spina iliaca ventralis caudalis. One needs to be careful to avoid
damaging the spina.
For detailed drawings and nomenclature see Barone et al. (1973, 24).
35
Appendix 2:
List of measurements, by site.
Column 1 shows SJMD's reference number, except in the case of Cova Matutano, where the
number is that used by JPNW in the study of the site.
For the modern material, the information given in the remaining columns is as follows:
Column 2 shows the number given in the register of the Muséum national d'Histoire naturelle
(MNHN).
Column 3 shows the reference number used by Callou (2003).
Column 4 shows the sex, when determined; M = male; F = female.
Column 5 shows the state of fusion of the proximal humerus: f = fully fused; fv = fusion plane still
visible; u = unfused.
Columns 6 to 11 show the measurements in millimetres.
For Cova Matutano, Column 2 shows the old level name and the depth in centimetres, while
Column 3 shows the faunal phase defined in Watson (1999).
Arjuzanx:
No. MNHN ref. Callou ref. Sex Fusion F1 F E J H I
342 2001.136 AX4 - f 8.3 7.5 4.4 8.2 6.9 3.3
346 2001.135 AX3 - f 7.9 7.6 4.2 7.7 6.4 3.0
348 2001.134 AX2 - f 7.9 6.8 3.8 7.7 6.0 3.0
Arróniz:
No. MNHN ref. Callou ref. Sex Fusion F1 F E J H I
278 2001.130 AZ - f 7.1 6.2 3.6 7.4 5.7 2.7
Badajoz:
No. MNHN ref. Callou ref. Sex Fusion F1 F E J H I
118 2001.177 BD6 M f 8.0 7.0 3.7 7.8 6.0 3.3
119 2001.181 BD10 F f 6.9 6.0 3.5 6.2 5.2 2.9
120 2001.187 BD16 F - 7.6 7.0 3.7 7.6 6.4 3.3
121 2001.174 BD3 F f 8.0 7.2 4.6 7.2 6.0 3.4
122 2001.186 BD15 F - 7.4 6.5 4.4 7.1 6.0 3.4
123 2001.185 BD14 M f 7.5 6.5 3.8 6.9 5.7 3.1
124 2001.180 BD9 M f 7.5 6.7 4.0 6.9 5.5 2.9
125 2001.173 BD2 M f 7.1 6.6 3.8 6.7 5.6 3.4
126 2001.176 BD5 M f 7.6 6.5 3.8 7.3 6.0 3.3
36
127 2001.175 BD4 F f 7.0 6.3 4.2 6.9 5.8 3.1
128 2001.182 BD11 F f 7.4 6.4 3.6 7.2 5.9 2.7
129 2001.172 BD1 M f 7.4 6.7 4.3 7.1 6.1 3.1
130 2001.179 BD8 M f 7.5 6.4 3.8 7.0 5.9 3.4
131 2001.178 BD7 M f 7.3 7.2 4.2 7.4 6.3 3.5
132 2001.184 BD13 F f 7.6 6.0 3.4 6.6 5.4 3.2
133 2001.183 BD12 M f 7.2 6.3 3.9 6.9 5.5 2.8
Caparroso:
No. MNHN ref. Callou ref. Sex Fusion F1 F E J H I
257 2001.127 CAP4 - f 6.9 6.6 3.5 6.9 6.0 2.7
730 2001.129 CAP6 M f 8.3 7.3 4.1 7.3 6.2 3.3
Castejón:
No. MNHN ref. Callou ref. Sex Fusion F1 F E J H I
277 2001.131 CN1 - f 7.9 6.7 4.4 7.9 5.9 3.5
Donzère:
No. MNHN ref. Callou ref. Sex Fusion F1 F E J H I
146 2001.53 DO1 - f 7.6 6.7 3.8 7.5 5.8 3.3
147 2001.54 DO2 - f 7.7 6.8 4.2 7.6 6.2 3.1
Falces:
No. MNHN ref. Callou ref. Sex Fusion F1 F E J H I
248 2001.82 FA1 F f 7.4 6.4 4.0 7.5 5.8 3.4
249 2001.83 FA2 F f 6.8 6.5 3.6 7.4 5.5 3.1
Las Lomas:
No. MNHN ref. Callou ref. Sex Fusion F1 F E J H I
271 1991.57 R34 F f 7.1 6.1 4.1 6.9 5.2 3.4
272 1991.34 R4 M f 7.3 7.0 3.9 7.0 5.9 3.2
273 1991.56 R33 F f 8.0 6.5 3.7 7.3 5.9 2.9
274 1991.53 R30 M f 7.6 6.5 4.3 7.1 5.9 3.3
275 2001.157 LL86 M u 6.6 5.6 3.3 6.8 5.5 2.8
276 2001.154 LL83 F f 6.9 6.3 4.2 7.2 5.6 2.9
280 2001.166 LL95 F fv 6.5 5.5 3.6 6.1 5.1 3.1
281 2001.161 LL90 F f 7.4 6.6 3.8 6.8 5.6 3.3
282 2001.156 LL85 F fv 7.1 6.3 3.5 6.6 5.2 2.7
37
283 2001.167 LL96 M f 7.6 6.6 3.7 7.7 6.0 3.3
284 2001.152 LL81 F f 7.1 6.4 3.9 7.1 5.8 3.3
285 2001.162 LL91 F f 7.6 6.4 4.2 7.9 5.6 2.9
286 2001.160 LL89 F f 7.6 6.4 4.0 7.0 5.8 3.5
287 2001.163 LL92 M f 7.3 6.6 4.2 7.1 5.9 3.3
288 2001.164 LL93 F f 7.4 6.7 4.1 7.2 6.2 2.8
289 2001.153 LL82 M f 7.6 6.1 4.1 7.2 6.0 3.1
290 2001.147 LL76 M fv 7.7 6.8 3.8 7.4 5.7 3.8
291 2001.159 LL88 M f 7.4 6.3 3.6 7.1 6.2 3.0
292 2001.141 LL70 M f 7.4 6.3 3.6 6.7 5.5 3.0
293 1991.60 R39 F f 7.4 6.2 3.9 6.7 5.4 3.2
294 1991.47 R22 F f 7.5 7.0 3.5 7.1 5.7 3.0
295 1991.59 R38 M f 7.8 6.8 4.1 7.3 6.0 3.8
296 1991.54 R31 M f 7.4 6.4 3.9 7.2 6.0 3.4
297 1991.40 R14 F f 6.9 6.2 3.5 6.9 5.4 3.0
298 1991.58 R36 M f 7.8 7.2 4.0 7.4 6.1 3.3
299 1991.33 R1 F fv 7.0 6.3 4.2 6.8 6.1 3.1
300 1991.46 R20 F f 7.2 6.3 3.7 6.8 5.3 2.9
301 1991.62 R41 F fv 6.9 6.6 3.9 7.3 5.5 3.3
302 1991.48 R23 M f 7.3 6.2 4.1 6.5 5.2 3.1
303 1991.61 R40 F f 7.5 6.9 3.9 6.8 5.8 3.4
304 1991.41 R15 F fv 7.3 6.5 4.2 6.5 5.8 3.3
305 1991.42 R16 M f 7.3 6.4 4.2 7.0 5.8 3.5
306 1991.65 R44 F f 7.2 6.7 3.9 7.3 5.7 3.8
307 1991.45 R19 F f 7.9 6.7 3.8 7.0 5.8 3.1
308 2001.151 LL80 F f 7.8 7.1 4.3 7.2 6.0 2.8
309 2001.148 LL77 F f 7.4 6.3 4.1 7.3 5.9 3.6
310 2001.146 LL75 M f 8.0 6.8 4.1 7.5 6.3 3.7
311 2001.145 LL74 M f 7.4 6.2 4.0 7.0 5.6 3.4
312 2001.171 LL100 F f 6.9 5.9 3.7 6.9 5.3 2.9
313 2001.165 LL94 M f 7.9 7.1 4.4 7.3 6.7 3.4
314 2001.150 LL79 F f 7.5 6.4 3.7 6.8 5.9 2.9
315 2001.168 LL97 F f 8.3 7.0 4.7 7.8 6.3 3.9
316 2001.170 LL99 F f 7.9 7.4 4.4 7.3 5.9 3.7
317 2001.169 LL98 M f 7.3 6.6 3.8 6.9 6.4 3.2
318 2001.158 LL87 M fv 7.5 6.2 3.5 6.4 5.3 3.2
319 2001.149 LL78 F f 7.7 6.5 4.1 6.8 6.0 3.4
320 2001.144 LL73 M f 7.1 6.5 3.9 6.9 5.7 3.3
321 2001.143 LL72 M f 7.5 6.3 3.8 7.2 5.9 3.7
322 2001.142 LL71 M f 7.1 6.6 3.8 7.0 5.9 3.1
323 1991.68 R47 F fv 7.3 6.5 3.7 7.0 5.4 3.1
324 1991.50 R25 F f 7.6 6.4 3.6 6.7 5.6 3.6
325 1991.37 R8 M fv 7.4 6.3 3.5 6.8 5.9 3.5
326 1991.64 R43 F f 7.3 6.4 3.8 7.2 5.4 3.4
327 1991.44 R18 F f 7.6 6.3 3.9 7.1 5.8 3.1
328 1991.35 R5 M fv 7.4 6.4 4.1 7.2 5.8 3.3
329 1991.63 R42 M f 7.8 6.6 4.2 7.6 6.2 3.7
330 1991.55 R32 F fv 7.3 6.5 4.1 7.1 5.6 3.3
38
331 1991.66 R45 F fv 6.6 5.9 3.8 6.8 5.4 2.8
332 1991.43 R17 F fv 7.6 7.1 3.9 7.2 6.2 2.9
333 1991.49 R24 M fv 6.9 5.7 3.6 6.5 5.3 3.0
334 1991.36 R7 M fv 7.6 7.2 3.9 6.6 5.6 3.6
335 1991.39 R12 M f 7.5 6.5 3.8 6.9 5.6 3.1
336 1991.38 R10 M fv 7.7 6.9 4.3 7.5 6.0 3.7
337 1991.52 R29 F f 7.3 6.4 3.7 6.6 5.9 3.0
338 1991.51 R27 M f 7.3 6.2 3.8 6.5 5.5 3.0
339 1991.67 R46 M f 7.7 6.8 3.8 7.4 6.1 3.4
340 2001.155 LL84 M f 7.2 6.4 3.8 7.3 6.0 3.0
Miranda de Arga:
No. MNHN ref. Callou ref. Sex Fusion F1 F E J H I
262 2001.91 MI4 M f 8.2 7.0 4.1 7.4 6.5 3.2
Peralta:
No. MNHN ref. Callou ref. Sex Fusion F1 F E J H I
221 2001.58 PE4 M u 7.2 6.4 4.4 7.0 5.6 3.0
222 2001.64 PE10 F fv 7.3 6.5 4.2 7.3 6.3 3.2
223 2001.66 PE12 M f 7.4 7.1 3.9 7.1 6.0 3.5
224 2001.77 PE23 F f 7.7 6.9 4.0 7.2 6.3 3.2
225 2001.55 PE1 F fv 7.6 6.5 3.6 7.1 6.1 3.0
226 2001.59 PE5 F fv 7.4 6.9 4.2 7.7 6.2 2.9
227 2001.75 PE21/27 F f 7.5 7.3 4.5 7.6 6.2 3.3
228 2001.67 PE13 M f 7.9 7.0 3.9 7.7 6.6 3.1
229 2001.68 PE14 M f 8.0 7.2 4.1 8.1 6.7 3.5
230 2001.81 PE27/21 M f 7.7 6.9 4.0 7.3 6.6 3.3
231 2001.74 PE20 M f 7.2 6.9 3.8 7.5 6.2 2.9
232 2001.72 PE18 F u 7.1 5.9 3.7 7.2 5.8 3.1
233 2001.60 PE6 M f 8.1 7.3 4.5 7.6 6.7 3.8
234 2001.80 PE26 M u 7.9 7.2 4.1 7.8 6.6 3.3
235 2001.63 PE9 F u 6.9 6.2 3.5 7.0 5.7 3.1
236 2001.61 PE7 F fv 7.7 6.8 3.9 7.3 6.2 3.3
237 2001.78 PE24 F u 7.3 7.0 3.9 7.7 6.6 3.2
238 2001.71 PE17 M fv 7.9 7.4 3.8 7.7 6.7 3.2
239 2001.65 PE11 F u 6.9 5.9 3.2 6.8 5.6 2.5
240 2001.70 PE16 F f 8.0 6.9 4.2 8.1 6.7 3.7
242 2001.69 PE15 M f 7.2 6.6 4.0 7.1 6.0 3.2
243 2001.57 PE3 M fv 7.9 6.7 4.3 7.6 6.2 3.4
244 2001.56 PE2 M f 8.4 7.5 4.3 7.8 6.5 3.4
245 2001.62 PE8 F f 8.4 7.5 4.0 7.7 6.8 3.7
246 2001.73 PE19 M fv 7.8 7.1 4.5 7.5 6.5 3.0
247 2001.76 PE22 M u 7.2 6.3 3.6 6.7 6.1 3.1
39
Ribaforada:
No. MNHN ref. Callou ref. Sex Fusion F1 F E J H I
263 2001.118 RI1 M f 7.7 7.4 3.8 7.1 6.6 3.0
265 2001.120 RI3 M f 7.7 7.2 4.2 8.3 6.7 3.8
267 2001.122 RI5 F f 7.2 6.9 4.0 7.7 6.5 3.3
Roissy:
No. MNHN ref. Callou ref. Sex Fusion F1 F E J H I
82 2001.33 RO32 M u 7.4 6.8 4.3 7.6 6.5 3.6
83 2001.50 RO52 M f 7.7 7.2 4.2 8.3 6.7 3.4
84 2001.40 RO42 M f 8.0 7.6 4.3 8.3 6.7 3.5
85 2001.39 RO41 M f 7.9 7.6 5.1 8.6 7.2 3.8
86 2001.45 RO47 M f 8.2 7.4 4.3 7.8 7.2 3.7
87 2001.23 RO22 M u 8.4 7.5 4.6 7.9 7.0 3.8
88 2001.32 RO31 M u 8.2 7.5 4.5 7.5 6.7 3.6
89 2001.47 RO49 M f 8.1 7.4 5.1 8.1 7.3 3.7
90 2001.49 RO51 M f 8.1 8.0 5.0 8.0 7.0 3.5
179 2001.3 RO2 - f 7.7 8.1 4.8 7.6 6.5 3.5
180 2001.4 RO3 - u 8.0 7.1 4.8 7.8 7.0 3.4
181 2001.5 RO4 - u 7.5 7.3 4.7 8.0 6.8 3.5
182 2001.6 RO5 - f 8.0 8.0 4.9 7.9 7.3 4.0
183 2001.8 RO7 - u 8.4 8.5 5.2 8.1 7.8 3.9
184 2001.9 RO8 - f 8.3 8.2 4.4 7.3 6.9 3.3
185 2001.10 RO9 - f 8.4 7.4 4.4 7.8 6.8 3.3
186 2001.11 RO10 - u 8.1 7.6 5.0 8.0 7.5 3.4
187 2001.12 RO11 - u 7.9 7.2 4.8 7.9 7.0 3.8
188 2001.13 RO12 - f 7.5 7.2 4.5 8.0 6.6 3.1
189 2001.14 RO13 - f 8.7 7.4 4.5 8.0 7.0 3.3
190 2001.15 RO14 - u 7.7 6.9 4.2 7.6 6.8 3.3
191 2001.16 RO15 M f 7.6 7.4 4.3 7.8 7.1 3.8
192 2001.17 RO16 M f 8.4 7.7 4.9 7.8 7.5 3.6
193 2001.18 RO17 M f 7.7 7.3 4.6 7.7 6.7 3.6
194 2001.19 RO18 F f 7.8 7.5 4.7 7.6 7.0 3.7
195 2001.20 RO19 F u 7.9 7.2 4.5 7.9 6.6 3.4
196 2001.21 RO20 F u 7.3 6.5 4.3 7.8 6.5 3.4
197 2001.24 RO23 F f 7.9 7.3 4.4 7.8 6.3 3.3
198 2001.25 RO24 M u 8.2 6.9 4.5 7.7 6.4 3.6
199 2001.26 RO25 F fv 7.9 7.7 4.6 8.3 7.0 3.5
200 2001.46 RO48 F f 7.8 7.2 5.1 8.4 6.7 3.2
201 2001.51 RO53 M f 8.4 7.6 4.5 8.1 7.1 3.6
204 2001.2 RO1 - u 7.5 6.8 4.6 7.6 6.6 3.3
205 2001.7 RO6 - f 8.2 7.8 4.4 8.3 7.0 3.6
206 2001.22 RO21 F f 8.1 7.7 4.6 8.4 7.0 3.5
207 2001.35 RO34 F u 7.8 7.2 4.2 7.9 7.0 3.4
208 2001.43 RO45 F f 7.9 7.5 4.2 8.0 7.2 3.5
40
209 2001.27 RO26 F fv 7.5 7.1 4.6 7.4 6.4 3.4
210 2001.28 RO27 F u 8.0 7.1 4.2 7.8 7.0 3.5
211 2001.42 RO44 M fv 8.0 7.4 4.5 7.8 6.7 3.6
212 2001.52 RO54 F f 7.9 7.4 4.9 7.9 6.8 3.1
213 2001.44 RO46 F f 7.9 7.4 4.9 7.9 7.0 3.4
214 2001.36 RO35 F fv 7.6 7.4 4.3 7.8 7.0 3.5
215 2001.37 RO36 F f 7.5 7.0 4.2 7.9 6.6 3.3
216 2001.29 RO28 M f 9.1 7.8 4.6 8.3 6.8 3.7
217 2001.34 RO33 F f 8.2 7.4 4.5 7.9 6.7 3.3
218 2001.30 RO29 F u 8.2 7.3 4.3 8.2 6.9 3.7
219 2001.31 RO30 F f 7.5 7.3 4.1 7.5 6.8 3.2
220 2001.41 RO43 F f 7.4 6.7 4.3 7.6 6.7 3.2
490 2001.48 RO50 F f 7.8 7.2 4.4 8.2 6.8 3.4
Santarém:
No. MNHN ref. Callou ref. Sex Fusion F1 F E J H I
30 1995.45 SM31 M u 7.1 6.0 3.6 6.8 5.7 3.0
32 1995.45 SM31 M u 6.5 5.5 4.0 6.6 5.2 2.6
34 1995.62 SM48 F u 7.1 5.7 4.0 5.9 5.1 2.9
35 1995.53 SM39 F u 6.6 6.1 3.3 6.9 5.8 3.1
37 1995.40 SM26 M u 7.4 5.9 3.4 6.6 5.5 2.8
38 1995.33 SM19 F u 7.2 6.0 3.4 6.6 5.5 2.8
42 1995.49 SM35 M u 6.8 5.6 3.6 6.2 5.5 2.9
43 1995.63 SM49 M u 7.7 6.5 4.1 7.2 5.9 3.3
44 1995.65 SM51 M u 6.7 5.5 2.9 6.1 5.2 2.6
46 1995.59 SM45 F u 6.9 6.0 4.1 6.4 5.3 3.3
47 1995.41 SM27 F u 7.2 5.9 4.0 6.6 5.6 3.0
48 1995.34 SM20 M u 7.4 5.7 3.8 6.5 5.6 2.9
52 1995.56 SM42 M u 7.3 5.7 3.2 7.0 5.1 2.8
53 1995.61 SM47 M u 6.9 5.9 3.3 6.6 5.4 2.7
54 1995.70 SM56 M u 7.1 6.1 3.6 6.9 5.9 2.6
55 1995.42 SM28 M u 7.0 6.1 4.0 6.9 5.7 2.9
56 1995.35 SM21 M u 7.0 5.9 3.3 6.8 5.6 3.0
65 1995.30 SM16 M f 7.4 6.3 3.7 7.1 5.7 3.0
66 1995.83 SM69 F f 7.2 6.3 3.7 6.6 5.3 3.2
67 1995.15 SM1 F f 7.2 6.1 3.7 7.7 6.1 2.8
68 1995.72 SM58 M f 8.1 6.9 3.9 7.3 6.2 3.1
69 1995.66 SM52 M f 7.9 7.1 4.3 7.9 6.1 3.4
70 1995.64 SM50 F f 7.5 6.0 3.6 6.8 5.8 3.2
71 1995.60 SM46 M f 7.1 6.6 3.9 7.1 6.2 2.9
72 1995.54 SM40 F f 7.6 6.4 3.5 7.1 6.0 3.3
73 1995.46 SM32 M f 7.2 6.1 3.2 6.9 5.5 3.3
360 1995.76 SM62 F f 6.9 6.0 3.7 7.0 5.7 3.1
362 1995.29 SM15 F f 7.3 6.4 3.4 6.8 5.9 3.0
364 1995.68 SM54 F f 7.7 6.7 3.8 7.5 6.0 3.3
368 1995.24 SM10 M f 7.4 6.0 3.4 6.7 5.6 2.9
41
378 1995.22 SM8 F f 7.0 6.1 3.3 6.6 5.2 2.9
379 1995.23 SM9 F f 7.4 6.6 3.8 7.4 5.9 2.8
380 1995.77 SM63 F f 7.0 6.3 3.5 6.9 5.8 2.9
381 1995.21 SM7 F f 7.3 6.3 3.7 6.7 5.6 2.8
382 1995.71 SM57 M fv 7.5 6.0 3.5 7.0 5.2 3.0
383 1995.67 SM53 M f 7.0 6.2 3.5 6.7 5.7 3.1
384 1995.75 SM61 M f 7.1 6.3 3.7 6.8 5.7 3.2
385 1995.20 SM6 M f 7.3 5.9 3.6 6.8 5.6 3.1
386 1995.74 SM60 F f 7.1 6.4 3.4 6.8 5.9 2.7
387 1995.73 SM59 M f 7.4 6.5 3.4 7.1 5.7 2.8
388 1995.69 SM55 F f 8.0 6.4 4.1 6.9 6.1 2.9
468 1995.79 SM65 M f 6.5 5.8 3.3 6.6 5.7 3.0
469 1995.87 SM73 M f 7.2 6.5 3.6 6.9 5.6 3.1
470 1995.82 SM68 F f 7.2 6.7 4.1 7.2 5.5 3.5
471 1995.37 SM23 F f 7.5 6.2 3.6 7.3 5.8 3.0
472 1995.38 SM24 M f 7.3 6.3 3.5 7.1 5.6 3.0
473 1995.85 SM71 M f 7.1 6.3 3.6 7.2 6.0 3.2
474 1995.78 SM64 F f 6.8 6.3 3.6 6.4 5.5 2.5
475 1995.25 SM11 F f 7.7 6.7 3.9 7.2 6.1 3.1
476 1995.43 SM29 F f 7.1 6.2 3.5 6.5 5.5 2.9
477 1995.44 SM30 F f 7.3 6.3 3.3 6.7 5.3 3.4
478 1995.27 SM13 M f 7.0 6.2 4.0 6.9 6.0 3.4
479 1995.19 SM5 F f 6.4 5.9 3.4 6.8 5.3 2.7
480 1995.84 SM70 F f 7.6 6.5 4.2 6.9 5.8 3.4
481 1995.86 SM72 M f 7.3 6.1 3.5 6.6 5.3 3.2
482 1995.80 SM66 F f 7.4 6.4 3.7 7.2 5.7 2.8
483 1995.81 SM67 M f 7.7 6.3 3.7 6.5 5.9 3.1
484 1995.28 SM14 F f 6.5 6.3 3.9 6.6 5.5 3.2
485 1995.16 SM2 M f 7.4 6.7 3.7 7.5 5.8 3.4
486 1995.17 SM3 F f 7.2 6.4 3.6 7.0 5.7 3.1
487 1995.26 SM12 M f 8.4 7.2 4.0 6.8 5.8 3.2
488 1995.88 SM74 M f 7.5 6.4 3.6 7.1 5.8 3.3
489 1995.18 SM4 F f 7.1 6.5 3.6 6.7 5.8 2.8
Sesma:
No. MNHN ref. Callou ref. Sex Fusion F1 F E J H I
279 2001.132 SE - f 8.1 7.3 4.0 8.2 6.6 3.3
Tudela:
No. MNHN ref. Callou ref. Sex Fusion F1 F E J H I
351 2001.106 TU13 F f 7.8 6.9 4.1 7.6 5.8 3.3
352 2001.110 TU17 F f 8.1 6.9 4.6 8.0 6.6 3.3
353 2001.115 TU22 F f 8.1 6.9 4.4 8.1 6.1 3.0
354 2001.117 TU24 M f 7.8 7.0 4.2 7.4 6.0 3.1
42
355 2001.95 TU2 - f 7.3 6.6 4.1 7.9 6.4 3.0
356 2001.111 TU18 F f 7.2 6.5 4.0 7.1 5.5 3.1
357 2001.113 TU20 F f 7.7 6.7 4.4 7.9 5.7 3.1
358 2001.116 TU23 F f 7.6 6.5 4.1 7.2 6.3 3.2
359 2001.107 TU14 F f 7.5 6.9 4.1 8.0 6.0 3.1
453 2001.100 TU7 - u 7.1 6.5 4.0 7.3 5.8 3.6
454 2001.114 TU21-19 F f 7.3 6.6 3.7 7.6 5.6 2.9
455 2001.97 TU4 - f 7.6 6.5 3.9 7.2 5.4 2.9
456 2001.96 TU3 - f 7.7 6.9 4.0 7.9 6.1 3.6
457 2001.94 TU1 - f 7.5 6.4 3.7 7.5 5.7 3.1
458 2001.104 TU11 - f 8.1 7.3 4.1 7.9 5.9 3.7
459 2001.102 TU9 - f 7.8 7.1 4.2 7.9 6.0 3.3
460 2001.112 TU19-21 F f 7.4 6.6 3.9 7.5 5.7 3.2
461 2001.105 TU12 - u 6.8 6.1 3.7 6.6 5.0 2.9
462 2001.109 TU16-15 M f 8.3 7.1 4.2 7.4 6.1 3.6
463 2001.103 TU10 - f 7.8 7.3 4.3 8.2 6.6 3.6
464 2001.99 TU6 - u 7.5 7.0 4.3 7.3 5.4 3.3
465 2001.101 TU8 - f 8.4 7.2 3.9 8.2 6.0 3.5
466 2001.98 TU5 - f 8.5 7.5 4.5 8.2 6.3 3.8
467 2001.108 TU15-16 M f 7.9 6.6 4.4 7.5 6.2 3.2
Tour du Valat:
No. MNHN ref. Callou ref. Sex Fusion F1 F E J H I
74 1991.73 C4 M f 7.7 7.2 4.3 7.9 6.7 3.3
75 1991.74 C7 M f 7.1 6.7 3.9 7.9 6.0 3.3
76 1991.70 C9 M f 7.4 6.6 4.2 7.4 6.4 3.2
78 1991.75 C11 F f 7.5 6.6 3.9 7.5 6.2 3.0
79 1991.71 C17 F f 7.3 6.6 3.9 7.5 6.6 2.9
80 1991.77 C15 F f 7.3 7.0 4.4 7.7 6.6 3.0
81 1991.72 C18 F f 7.9 6.6 4.0 7.4 6.2 2.5
135 1991.80 C14 M f 7.1 7.0 3.9 8.1 6.7 3.1
136 2001.92 C29 - f 7.2 6.2 3.6 6.9 5.6 3.2
137 2001.82 C12 F f 7.8 7.6 4.4 8.0 6.2 3.1
138 2001.79 C1 F f 7.9 7.6 4.6 8.5 7.0 3.7
139 2001.87 C10 - f 7.3 6.5 4.0 7.6 6.4 3.2
140 2001.81 C16 M f 7.2 6.6 3.7 7.2 6.2 3.1
141 2001.90 C27 - f 7.2 6.5 4.0 7.2 5.7 3.1
142 2001.85 C6 M f 7.8 7.3 4.4 7.9 7.1 3.2
143 2001.86 C13 M f 7.3 6.5 4.0 7.5 6.2 2.8
144 2001.84 C5 F f 7.2 7.1 3.9 7.7 6.6 2.9
145 2001.83 C2 F f 6.9 6.8 4.1 7.7 6.4 3.5
202 2001.89 C26 - u 6.5 5.5 3.3 6.6 5.1 2.6
43
Versailles:
No. MNHN ref. Callou ref. Sex Fusion F1 F E J H I
93 2001.208 V21 F f 7.5 7.3 4.3 7.6 6.4 3.5
94 2001.209 V22 M f 8.0 7.7 4.3 8.1 6.9 3.4
95 2001.204 V17 F f 7.7 7.1 4.4 7.9 6.6 3.4
96 2001.194 V7 M f 8.1 7.4 4.3 7.7 6.9 3.7
97 2001.199 V12 F f 7.8 7.1 4.4 7.5 6.6 3.5
98 2001.198 V11 M f 7.1 6.7 4.3 7.4 6.2 3.2
99 2001.203 V16 F f 7.8 7.0 4.9 7.5 6.8 3.4
100 2001.210 V23 M f 7.6 7.1 4.4 7.7 6.3 3.2
101 2001.200 V13 F f 7.6 6.9 4.2 7.4 6.2 3.5
102 2001.205 V18 F f 7.7 7.1 4.8 7.2 6.1 3.3
103 2001.195 V8 F f 8.1 7.2 4.6 7.6 6.6 3.4
104 2001.188 V1 - f 8.5 7.4 4.3 8.0 6.9 3.5
105 2001.191 V4 F f 8.2 7.9 4.5 7.6 7.0 3.5
107 2001.193 V6 F f 8.2 7.5 4.9 7.9 6.6 3.3
108 2001.207 V20 M f 8.1 7.5 5.1 8.0 7.0 3.9
109 2001.197 V10 F f 7.7 7.3 4.5 7.3 6.6 3.3
110 2001.202 V15 F f 8.3 7.0 4.7 7.6 6.4 3.5
111 2001.212 V25 F f 8.3 7.7 4.9 8.2 7.0 3.5
112 2001.189 V2 - u 7.1 6.8 4.2 6.9 6.3 3.5
113 2001.192 V5 F f 7.6 6.8 4.4 7.5 6.6 3.5
114 2001.196 V9 M f 8.0 7.5 4.8 7.6 7.1 3.5
116 2001.211 V24 F f 8.3 7.1 4.5 7.9 7.0 3.4
117 2001.201 V14 M f 7.8 7.3 4.6 7.8 7.1 3.2
Vila Viçosa:
No. MNHN ref. Callou ref. Sex Fusion F1 F E J H I
1 1995.134 VV46 F u 7.6 6.5 3.6 7.0 5.4 3.0
2 1995.99 VV11 F f 7.7 7.2 3.8 7.5 6.2 3.1
3 1995.129 VV41 M fv 6.8 5.9 4.0 6.9 5.6 3.1
4 1995.138 VV50 M f 7.1 6.0 3.8 6.8 5.6 3.4
5 1995.97 VV9 F f 7.0 6.7 3.5 7.2 5.5 3.3
6 1995.125 VV37 F f 6.7 6.4 3.5 6.8 5.8 2.9
7 1995.139 VV51 F f 7.6 6.4 3.8 7.3 5.8 3.3
8 1995.115 VV27 M f 7.1 6.4 3.9 7.1 5.8 3.3
9 1995.136 VV48 F f 7.4 6.7 3.7 7.4 6.0 3.1
10 1995.90 VV2 F f 7.1 6.5 3.5 7.2 6.3 3.1
11 1995.100 VV12 M f 7.0 6.1 3.3 6.9 5.6 2.7
12 1995.105 VV17 M f 6.8 6.1 3.3 6.4 5.7 2.6
13 1995.126 VV38 F f 6.9 6.4 3.4 6.9 5.7 3.3
14 1995.127 VV39 F f 7.2 6.4 3.4 7.1 5.7 3.2
15 1995.140 VV52 M f 7.6 6.9 3.7 7.3 5.8 3.5
16 1995.135 VV47 F f 8.2 6.6 3.3 7.6 5.9 3.2
17 1995.114 VV26 F f 7.5 7.1 4.0 6.7 5.9 3.4
44
18 1995.131 VV43 M f 7.1 6.8 3.8 7.1 5.9 3.3
19 1995.91 VV3 F f 7.0 6.6 3.5 7.0 5.8 2.9
20 1995.128 VV40 F f 7.5 6.5 3.7 7.0 5.7 3.1
21 1995.124 VV36 F u 7.2 6.8 3.8 7.0 5.9 3.3
22 1995.102 VV14 F f 7.0 6.9 3.5 7.0 5.9 2.9
23 1995.132 VV44 F f 7.8 7.2 3.6 7.1 5.9 3.2
24 1995.109 VV21 F f 7.2 6.5 4.2 6.8 6.1 3.1
25 1995.137 VV49 M f 7.3 6.2 3.9 7.1 5.6 3.5
26 1995.116 VV28 M f 7.7 6.6 4.2 7.4 6.1 3.6
27 1995.141 VV53 F f 7.3 6.4 3.6 7.1 5.7 3.2
49 1995.121 VV33 F u 7.4 6.5 4.1 6.9 5.8 3.6
51 1995.108 VV20 F u 6.9 6.2 3.6 6.7 5.7 3.0
57 1995.101 VV13 M u 7.2 6.1 3.4 6.3 5.5 3.0
58 1995.123 VV35 F u 7.3 6.2 3.6 7.1 5.8 3.5
60 1995.110 VV22 F u 6.6 6.0 3.5 6.5 5.8 3.2
62 1995.111 VV23 M u 6.9 6.3 3.6 6.4 5.3 3.2
361 1995.103 VV15 M f 6.8 6.3 3.5 6.8 5.5 2.5
363 1995.120 VV32 F f 6.6 5.8 3.9 7.2 5.9 2.9
365 1995.133 VV45 F f 7.6 6.4 4.4 7.9 5.9 3.1
366 1995.119 VV31 M f 7.4 6.8 4.0 7.9 6.1 3.2
367 1995.122 VV34 F f 7.7 6.6 3.6 7.7 5.8 3.3
369 1995.94 VV6 M f 7.2 6.9 3.6 7.1 6.2 3.2
370 1995.89 VV1 M f 7.4 6.5 3.8 7.1 5.6 3.3
371 1995.130 VV42 F fv 7.1 6.6 3.9 7.1 6.1 3.6
372 1995.117 VV29 F f 7.2 6.6 3.4 7.5 5.8 2.8
373 1995.93 VV5 F f 7.5 6.8 3.7 7.1 5.4 3.3
374 1995.113 VV25 F f 7.7 6.9 3.8 7.3 6.0 3.3
375 1995.98 VV10 M f 6.8 6.3 3.8 6.6 5.8 3.4
376 1995.104 VV16 M f 7.6 6.7 3.9 7.4 5.7 2.8
377 1995.96 VV8 F f 6.8 6.0 3.4 6.9 5.5 2.9
Zembra:
No. MNHN ref. Callou ref. Sex Fusion F1 F E J H I
148 1991.9 ZA9 - f 7.4 6.4 3.6 7.2 5.4 3.1
149 1991.5 ZA5 - f 7.8 7.0 3.9 7.3 6.3 3.0
151 1991.3 ZA3 - f 8.0 7.1 3.8 7.0 6.1 3.3
152 1991.1 ZA1 - f 7.5 6.8 4.2 7.5 6.0 3.2
156 1991.17 ZA17 - f 7.6 6.7 3.7 7.4 5.9 2.5
157 1991.15 ZA15 - f 7.7 7.2 4.1 7.9 6.3 3.5
158 1991.8 ZA8 - f 7.5 7.0 4.4 7.3 5.9 3.0
160 1991.14 ZA14 - f 7.6 6.6 3.7 7.0 6.1 2.7
165 1991.11 ZA11 - f 8.1 6.7 3.9 6.8 6.1 3.2
166 1991.12 ZA12 - f 7.0 6.2 3.8 6.7 5.5 3.3
169 1991.19 ZA19 - f 6.4 6.2 3.6 6.7 5.5 3.0
174 1991.26 ZA26 - f 7.2 6.5 3.9 6.5 5.6 2.8
175 1991.27 ZA27 - f 8.0 7.0 4.3 7.7 6.6 3.0
45
176 1991.28 ZA28 - f 7.5 6.9 3.9 7.2 6.1 3.3
178 1991.30 ZA30 - f 7.7 6.6 3.7 7.1 6.0 3.3
Domestic (Papillon):
No. MNHN ref. Sex Fusion F1 F E J H I
940 2007-1560 F - 10.0 9.2 6.6 9.9 8.8 4.2
941 2007-1559 M - 9.6 8.7 6.5 9.7 7.9 4.3
Domestic (Gris de Vienne):
No. MNHN ref. Sex Fusion F1 F E J H I
942 2007-1561 M - 11.2 10.1 7.5 11.6 10.1 4.6
943 2007-1562 M - 11.0 10.2 6.5 11.1 9.8 5.0
944 2007-1565 M - 10.7 9.6 6.0 10.4 9.5 4.4
945 2007-1563 M - 9.7 8.3 5.8 8.8 8.1 4.1
947 2007-1564 F f 10.9 9.8 6.4 9.7 8.9 4.6
Cova Matutano:
No. Level Phase F1 F E J H I
3408 N4 97-113 C 8.9 7.7 4.1 7.7 6.2 3.6
3517 N7 165-171 B 8.6 7.1 3.8 8.0 6.2 3.2
3518 N7 165-171 B 7.4 6.4 3.5 6.7 6.3 3.0
3520 N7 165-171 B 8.2 7.0 4.0 8.0 6.0 3.6
3805 N8 201-208 A 9.0 7.3 4.5 7.6 6.7 3.7
3806 N8 201-208 A 8.8 7.8 4.3 8.4 6.8 3.9
3807 N8 201-208 A 8.4 7.5 4.1 8.2 6.4 3.8
3812 N8 201-208 A 7.9 7.3 4.1 7.4 6.2 3.6
3813 N8 201-208 A 8.6 7.1 4.1 7.5 6.4 3.6
3820 N8 201-208 A 8.2 7.1 3.9 7.7 6.3 3.9
3822 N8 201-208 A 8.4 7.0 4.2 7.6 6.2 3.6
3824 N8 201-208 A 9.4 7.7 4.5 8.4 6.9 3.1
3829 N8 201-208 A 8.8 7.5 4.5 8.0 6.6 3.6
3855 N8 201-208 A 8.3 7.4 4.2 7.7 6.3 3.6
3856 N8 201-208 A 8.2 7.2 3.9 8.0 6.4 3.6
3858 N8 201-208 A 8.7 7.3 4.2 8.0 6.5 3.9
3859 N8 201-208 A 8.9 7.6 4.3 8.4 7.0 3.6
3862 N8 201-208 A 7.7 7.0 4.1 7.9 6.1 3.1
3865 N8 201-208 A 8.3 7.2 3.9 7.4 6.5 3.8
4477 N3 77-97 C 7.9 7.7 4.0 8.2 6.9 3.7
4490 N3 77-97 C 8.2 6.9 3.8 7.4 6.2 3.1
4512 N3 77-97 C 9.4 7.8 4.5 8.8 7.4 3.5
4729 N1 48-68 D 8.7 7.7 4.2 7.9 6.7 3.5
46
4985 N7 177-187 B 8.3 7.1 4.0 7.5 6.6 3.1
4987 N7 177-187 B 8.0 7.3 4.1 8.2 6.1 3.3
4988 N7 177-187 B 8.3 7.3 4.1 7.9 6.0 3.7
4990 N7 177-187 B 8.6 7.3 4.2 8.0 6.8 3.7
5006 N7 177-187 B 8.8 8.0 4.3 8.6 6.9 3.5
5012 N7 177-187 B 8.6 7.1 3.8 8.5 6.4 3.1
5014 N7 177-187 B 8.2 7.3 4.2 7.8 6.5 3.5
5018 N7 177-187 B 7.8 6.6 3.4 7.0 5.9 3.3
5299 N5 133-155 C 7.8 6.9 3.9 7.3 5.9 3.2
5303 N5 133-155 C 9.0 7.9 4.5 8.9 6.6 3.6
5308 N5 133-155 C 9.0 7.8 4.6 7.9 7.1 3.3
5315 N5 133-155 C 8.0 6.9 3.8 7.5 6.2 3.7
5509 N4 113-133 C 8.5 7.0 5.0 8.0 6.9 3.7
5516 N4 113-133 C 8.7 7.1 4.3 8.0 6.5 3.6
5517 N4 113-133 C 8.2 7.3 4.2 8.0 6.3 3.2
5519 N4 113-133 C 7.6 7.0 4.0 7.7 6.2 3.3
5526 N4 113-133 C 7.5 7.3 4.0 8.5 6.5 3.4
5527 N4 113-133 C 7.3 5.8 3.5 6.8 5.3 2.4
5736 N6 153-162 B 7.5 7.0 3.5 7.0 6.0 3.0
5737 N6 153-162 B 8.4 7.5 4.3 9.0 6.6 3.7
5740 N6 153-162 B 7.9 6.8 3.9 7.5 6.1 3.2
5762 N6 153-162 B 8.4 7.6 4.1 8.3 6.8 3.4
5765 N6 153-162 B 8.2 7.5 4.1 7.7 6.7 3.4
5767 N6 153-162 B 8.0 7.2 4.1 7.8 6.1 3.6
5771 N6 153-162 B 7.7 6.8 3.3 7.2 6.0 3.5
6051 N7 171-177 B 9.1 7.6 4.5 8.6 6.8 3.8
6188 N2 68-77 D 7.8 6.8 4.0 7.5 6.3 3.5
6189 N2 68-77 D 8.1 7.1 4.0 7.9 6.7 3.3
6194 N2 68-77 D 7.5 6.8 3.7 7.3 6.3 3.4
6469 N8 A 8.3 7.2 4.6 8.0 6.7 3.7
6472 N8 A 7.5 7.1 3.6 7.4 6.3 3.1
6486 N8 A 7.8 7.4 4.3 7.9 6.4 3.0
6487 N8 A 9.0 8.0 4.4 8.6 7.0 3.2
6505 N8 A 8.8 7.8 4.0 8.2 6.9 3.6
6507 N8 A 8.0 6.7 3.9 7.2 6.1 3.2
6508 N8 A 8.0 6.9 4.2 7.6 6.5 3.6
6514 N8 A 9.1 7.3 4.1 7.0 6.0 3.5
6517 N8 A 8.2 7.4 3.9 8.3 6.4 3.1
6530 N8 A 7.5 6.7 4.1 7.8 6.4 3.5
6531 N8 A 8.3 7.3 4.5 7.6 6.9 3.4
6532 N8 A 9.5 8.0 4.6 8.6 6.9 4.0
6535 N8 A 7.5 6.8 3.9 7.7 6.2 3.3
6536 N8 A 8.0 6.7 4.4 7.8 5.8 3.8
6742 NS 8-48 E 8.8 7.1 4.2 8.1 6.3 3.3
6747 NS 8-48 E 8.3 7.3 4.1 7.9 6.7 3.5
6754 NS 8-48 E 8.5 7.3 4.4 7.9 6.6 3.4
6755 NS 8-48 E 8.6 7.6 4.5 8.3 7.0 3.8
47
Beja:
No. Site ref. no. F1 F E J H I
495 Silo 65 65/84 XIV 8.4 6.7 4.3 7.7 6.4 3.1
497 Silo 65 65/84 XIV 8.3 7.4 4.2 7.7 6.3 3.6
498 Silo 65 65/84 XIV 7.9 6.8 3.7 6.9 6.0 3.6
499 Silo 65 65/84 XIV 8.3 6.8 3.7 7.2 6.2 3.1
500 Silo 65 65/84 XIV 6.7 6.3 3.5 6.4 5.1 2.8
501 Silo 65 65/84 XIV 7.3 6.2 3.9 6.9 5.5 3.6
502 Silo 65 65/84 XIV 7.1 5.9 3.6 6.3 5.2 3.3
503 Silo 65 65/84 XIV 7.8 6.9 4.0 7.1 5.9 3.3
504 Silo 65 65/84 XIV 7.5 6.5 3.8 7.1 6.1 3.3
505 Silo 65 65/84 XIV 7.6 6.5 3.9 7.0 5.7 3.4
507 Silo 65 65/84 XIV 6.8 6.1 3.5 6.5 5.3 3.3
508 Silo 65 65/84 XXI 7.7 6.1 3.2 6.8 5.6 3.0
509 Silo 65 65/84 XXI 7.1 6.0 3.9 6.5 5.5 3.0
514 Silo 80 80/114 XXVI 7.3 6.4 3.6 7.1 5.6 3.0
515 Silo 80 80/114 XXVI 7.3 6.1 3.4 6.8 5.2 2.8
516 Silo 80 80/116 XXXV 7.1 6.1 3.6 6.8 5.6 3.2
517 Silo 80 80/116 XXXV 7.9 6.6 3.6 7.2 5.8 3.3
518 Silo 80 80/116 XXXV 7.5 6.6 3.6 6.9 5.5 3.1
519 Silo 80 80/118 LI 7.9 7.1 4.0 7.3 6.1 3.9
520 Silo 80 80/118 LI 8.7 7.6 4.5 8.1 6.5 3.7
521 Silo 86 86/119 II 7.6 6.6 4.0 7.3 6.2 3.1
522 Silo 89 89/129 XVI 7.2 5.9 3.9 7.0 5.4 3.0
524 Silo 86 86/123 XL 8.1 7.2 4.0 7.3 6.4 3.3
527 Silo 86 86/120 XIV 7.9 7.0 3.9 7.6 5.8 3.4
528 Silo 86 86/126 L 7.1 6.4 3.5 7.1 5.9 3.3
529 Silo 86 86/126 L 8.1 7.4 4.5 7.6 6.8 3.4
530 Silo 128 128/175 XLVI 7.4 6.7 3.9 6.7 5.9 3.4
531 Silo 128 128/173 XXIX 7.6 6.5 3.8 6.8 5.7 3.0
532 Silo 128 128/173 XXIX 8.1 7.1 4.5 6.9 6.4 3.6
534 Silo 128 128/176 LII 7.5 6.5 3.7 6.9 5.9 3.1
535 Silo 128 128/176 LII 7.9 6.7 3.6 7.3 6.3 3.3
536 Silo 128 128/176 LII 7.5 6.5 4.3 7.0 5.5 2.9
537 Silo 128 128/176 LII 7.9 7.1 4.1 7.5 6.2 3.6
538 Silo 128 128/176 LII 7.6 6.3 3.5 7.0 5.9 3.1
539 Silo 128 128/176 LII 7.3 6.4 3.6 6.8 6.2 3.4
540 Silo 128 128/170 VI 8.7 7.0 4.2 8.1 6.6 3.6
541 Silo 128 128/170 VI 8.1 7.2 4.0 7.6 6.0 3.2
542 Silo 128 128/170 VI 7.3 6.2 3.7 7.0 5.9 3.5
543 Silo 128 128/170 VI 7.5 6.5 3.7 7.2 6.0 3.1
544 Silo 128 128/170 VI 7.2 6.2 3.6 7.0 5.6 2.9
545 Silo 128 128/172 XXII 7.5 7.1 3.9 7.4 6.3 3.0
546 Silo 128 128/172 XXII 7.8 7.1 3.8 7.0 6.5 3.4
547 Silo 128 128/172 XXII 7.7 6.5 4.1 6.7 5.7 3.2
549 Silo 128 128/172 XXII 8.2 7.0 3.9 7.1 5.6 3.2
550 Silo 128 128/172 XXII 7.8 7.1 4.1 7.6 6.4 3.0
48
Silves:
No. Site ref. no. F1 F E J H I
592 Quad-07 Est-8 Cam1032 7.1 6.5 3.7 7.0 5.7 2.9
593 Quad-L7 Est-5 Cam1034 7.5 6.5 3.8 6.8 5.3 3.2
594 Quad-J7 Est-5 Cam1039 7.4 6.9 4.0 7.4 6.0 3.0
595 Quad-O7 Est-7 Cam1050 7.4 6.6 3.2 6.6 5.5 2.9
598 Quad-M7 Est-8 Cam1034 7.7 6.2 3.7 7.2 6.1 3.3
599 Quad-N7 Est-11 Cam1031 7.0 6.1 3.5 6.6 5.4 2.8
600 Quad-N7 Est-10 Cam1030 6.9 6.5 3.7 6.7 5.5 3.0
601 Quad-K7 Est-6 Cam1043 7.3 6.5 3.8 7.0 5.6 2.9
602 Quad-L7 Est-5 Cam1034 7.9 6.4 3.5 7.1 5.9 3.0
603 Quad-L7 Est-5 Cam1034 7.5 6.7 3.8 7.0 5.8 3.3
604 Quad-L7 Est-5 Cam1034 7.7 6.6 4.0 7.2 6.2 3.0
605 Quad-M7 Est-4A Cam1034 7.9 6.9 3.6 7.3 5.9 3.3
606 Quad-M7 Est-4A Cam1034 7.7 6.6 3.8 7.4 5.9 3.3
607 Quad-M7 Est-8A Cam1034 7.1 6.3 3.5 7.6 5.7 3.0
608 Quad-L7 Est-5 Cam1034 7.4 6.6 4.0 7.3 5.8 3.0
609 Quad-K7 Est-6 Cam1043 7.4 6.5 3.6 6.6 5.6 3.1
610 Quad-N7 Est-7 Cam1050 7.0 6.2 3.3 6.2 5.5 3.1
611 Quad-M7 Est-4A Cam1034 7.3 6.2 3.5 6.7 6.3 3.1
612 Quad-M7 Est-8 Cam1034 7.4 6.0 3.7 7.1 5.7 3.0
613 Quad-M7 Est-8 Cam1034 7.9 6.5 3.6 6.9 5.6 2.6
614 Quad-M7 Est-8 Cam1034 7.1 6.3 3.8 6.5 5.5 2.9
615 Quad-M7 Est-8 Cam1034 6.9 6.2 3.3 6.5 5.4 2.8
616 Quad-M7 Est-8 Cam1034 7.6 6.5 3.7 7.1 6.1 3.3
617 Quad-M7 Est-6 Cam1036 7.2 6.7 3.7 7.2 5.8 3.2
618 Quad-O6 Est-5 Cam1036 7.3 6.5 4.0 7.1 5.7 3.6
619 Quad-L7 Est-4 Cam1035 7.9 6.4 3.4 6.9 5.6 2.8
620 Quad-N6 Est-5a1 Cam1034 7.1 6.6 3.7 6.9 5.9 2.9
49
