and Tighennif (Algeria)

20 Pages·2016·1.54 MB·French

Bulletins et mémoires de la Société
d’Anthropologie de Paris 

19 (3-4) | 2007
2007(3-4)
Comparative analysis of the Gran Dolina-TD6
(Spain) and Tighennif (Algeria) hominin mandibles
Analyse comparée des mandibules d’hominidés de Tighennif (Algérie) et de
Gran-Dolina-TD6 (Espagne)
José-Maria Bermúdez de Castro, María Martinón-Torres, Aida Gómez-
Robles, Leyre Prado et Susana Sarmiento
Édition électronique
URL : https://journals.openedition.org/bmsap/4623
DOI : 10.4000/bmsap.4623
ISSN : 1777-5469
Éditeur
Société d'Anthropologie de Paris
Édition imprimée
Date de publication : 1 décembre 2007
Pagination : 149-167
ISSN : 0037-8984

Référence électronique
José-Maria Bermúdez de Castro, María Martinón-Torres, Aida Gómez-Robles, Leyre Prado et Susana
Sarmiento, « Comparative analysis of the Gran Dolina-TD6 (Spain) and Tighennif (Algeria) hominin
mandibles », Bulletins et mémoires de la Société d’Anthropologie de Paris [En ligne], 19 (3-4) | 2007, mis
en ligne le 04 janvier 2011, consulté le 01 juin 2021. URL : http://journals.openedition.org/bmsap/4623
; DOI : https://doi.org/10.4000/bmsap.4623
Les contenus des Bulletins et mémoires de la Société d’Anthropologie de Paris sont mis à disposition
selon les termes de la licence Creative Commons Attribution-NonCommercial-NoDerivatives 4.0
International License.
Bulletins et Mémoires de la Société d’Anthropologie de Paris, n.s., t. 19, 2007, 3-4, p. 149-167
COMPARATIVE ANALYSIS OF THE GRAN DOLINA-TD6 (SPAIN)
AND TIGHENNIF (ALGERIA) HOMININ MANDIBLES
ANALYSE COMPARÉE DES MANDIBULES D’HOMINIDÉS DE TIGHENNIF
(ALGÉRIE) ET DE GRAN-DOLINA-TD6 (ESPAGNE)
José María BERMÚDEZ DE CASTRO 1, María MARTINÓN-TORRES 1, Aida GÓMEZ-ROBLES 1,
Leyre PRADO 1, Susana SARMIENTO 1
ABSTRACT
We present a comparative study of the Tighennif (Algeria) and Gran Dolina-TD6 (Spain) hominin mandibles, which
represent two geographically near and contemporaneous populations separated by the Mediterranean sea, in order to test
the hypothesis that these populations belong to the same evolutionary lineage, as it has been suggested by some authors.
The Tighennif mandibles show a clear primitive structural pattern, derived in some features with regard to the oldest Homo
specimens from H. habilis, as well as from the Javanese H. erectus. In addition, the Tighennif specimens share all these
derived features with H. ergaster and, some of them, with H. antecessor. However, the Gran Dolina-TD6 specimens are
remarkably smaller than those of Tighennif, and lack the robustness which characterizes the African Pleistocene mandibles.
The main difference between both groups in terms of mandibular dimensions can be related to the higher corpus height
characteristic of Tighennif specimens. The dental evidence reveals that North African Middle Pleistocene populations are
morphologically closer to African Early Pleistocene populations than to TD6 fossils. We conclude that the Spanish and
Algerian hominins belong to different hominin lineages. The Tighennif hominins, together with other contemporaneous
(Thomas Quarry and Oulad Hamida 1), and perhaps later North African specimens (Sidi Abderrahaman, Salé, and Rabat
[Kebitat]) should be considered as a subspecies of the H. ergaster species, i.e. H. ergaster mauritanicus, and may be the
result of an evolution in isolation in this African area. In agreement with the mandibular and dental evidences, the Gran
Dolina-TD6 hominins could belong to an exclusive Eurasian lineage.
Keywords: Early Pleistocene, North Africa, Europe, hominins, mandibles.
RÉSUMÉ
Nous présentons une analyse comparée des mandibules des hominidés de Tighennif (Algérie) et de la Gran Dolina-
TD6 (Espagne), qui représentent deux populations proches dans l’espace et le temps mais séparées par la mer Méditerranée,
afi n de vérifi er l’hypothèse que les deux échantillons appartiennent au même lignage évolutif, comme certains auteurs l’ont
1. Centro Nacional de Investigación sobre Evolución Humana, Avenida de la Paz 28, 09005 Burgos, Espagne, e-mail: [email protected]
Manuscrit reçu le 13 septembre 2007, accepté le 27 décembre 2007
150 J.M. BERMÚDEZ DE CASTRO, M. MARTINÓN-TORRES, A. GÓMEZ-ROBLES, L. PRADO, S. SARMIENTO
suggéré. Les mandibules de Tighennif ont un pattern morphologique structurel primitif, avec certains caractères dérivés
par rapport à ceux des spécimens de H. habilis et de H. erectus de Java. Les mandibules de Tighennif partagent tous ces
caractères dérivés avec H. ergaster et quelques uns avec H. antecessor. Néanmoins, les spécimens de la Gran Dolina-
TD6 sont remarquablement plus petits que ceux de Tighennif, et sont moins robustes que les mandibules du Pléistocène
d’Afrique. La principale différence entre les deux échantillons, en ce qui concerne les dimensions mandibulaires, est la
grande hauteur du corps de la mandibule des spécimens de Tighennif. Les caractères dentaires révèlent que les populations
du Pléistocène moyen du Nord de l’Afrique sont, du point de vue morphologique, plus proches des hominidés du Pléistocène
inférieur de l’Afrique que de ceux de la Gran Dolina-TD6. Nous concluons donc que les hominidés de Tighennif et de la
Gran Dolina-TD6 appartiennent à deux lignées différentes. Les hominidés de Tighennif pourraient être réunis avec les
spécimens contemporains de Thomas Quarry et de Oulad Hamida 1, et peut-être avec d’autres plus tardifs comme ceux
de Sidi Abderrahaman, Salé, et Rabat [Kebitat] du Nord de l’Afrique dans une même sous-espèce de H. ergaster, i.e.
H. ergaster mauritanicus, qui serait le résultat d’une évolution locale en isolement relatif dans cette région de l’Afrique. Les
hominidés de la Gran Dolina-TD6 appartiennent à une lignée exclusive de l’Eurasie, conformément aux données dentaires
et mandibulaires.
Mots-clés : Pléistocene inférieur, Nord de l’Afrique, Europe, hominidés, mandibules.
Version abrégée de la Gran Dolina-TD6 ont été inclus dans l’espèce
H. antecessor (Bermúdez de Castro et al. 1997). Certains
Dans ce travail, nous présentons une comparaison auteurs ont considéré la possibilité de rassembler les
métrique et morphologique entre deux échantillons deux échantillons dans la même espèce : H. mauritanicus
de mandibules fossiles : Tighennif (Algérie) et Gran (Hublin 2001 ; Stringer 2003). Cependant, ces auteurs
Dolina-TD6 (Espagne), qui sont proches dans l’espace n’ont pas présenté une étude formelle pour défendre leur
et le temps, mais séparées par la mer Méditerranée. Le alternative. Par conséquent, l’objectif de ce travail est de
gisement de Tighennif est bien connu par la découverte tester l’hypothèse que les hominidés de Tighennif et de la
de trois mandibules complètes, ainsi que par un os pariétal Gran Dolina-TD6 représentent une même lignée évolutive
et quelques dents isolées. Tous ces fossiles humains, sur les deux bords de la mer Méditerranée.
associés à plusieurs centaines de fossiles de mammifères Nos résultats indiquent que les mandibules de
et à des outillages lithiques inclus dans l’Oldowayen et Tighennif ont un pattern morphologique structurel
l’Acheuléen (Arambourg 1954a, b), représentent les primitif, dont certains caractères sont dérivés par rapport
hominidés les plus anciens du Nord de l’Afrique, avec une à ceux de H. habilis et des spécimens de H. erectus
chronologie de ca. 700 000 ans (Geraads et al. 1986). de Java. Les mandibules de Tighennif partagent tous
Le niveau TD6 du gisement de la grotte de la Gran ces caractères avec H. ergaster et quelques-uns avec
Dolina, située dans la Sierra d’Atapuerca, a livré plus H. antecessor. Néanmoins, les spécimens de la Gran
d’une centaine de fossiles humains associés avec des Dolina-TD6 sont remarquablement plus petits que ceux
mammifères fossiles et des outils lithiques de tradition de Tighennif, et sont moins robustes que les mandibules
oldowayenne (Carbonell et al. 1995). Cet ensemble, du Pléistocène d’Afrique. La principale différence entre
localisé sous l’inversion Matuyama/Bruhnes, est daté de les deux échantillons, en ce qui concerne les dimensions
ca. 800 000 ans B.P. (Parés, Pérez-González 1995, 1999 ; mandibulaires, est la grande hauteur du corps de la
Falguères et al. 1999). L’échantillon de fossiles humains mandibule des spécimens de Tighennif. Les caractères
inclut trois fragments de mandibule bien conservés. dentaires révèlent que les populations du Pléistocène
Originellement, les hominidés de Tighennif furent moyen du Nord de l’Afrique sont, du point de vue
inclus dans l’espèce Atlanthropus mauritanicus par morphologique, plus proches des hominidés du Pléistocène
Arambourg (1954a) et postérieurement attribués à l’espèce inférieur de l’Afrique que de ceux de la Gran Dolina-TD6
Homo erectus par Le Gros Clark (1964). Les hominidés (Martinón-Torres et al. 2007).
151 GRAN DOLINA-TD6 AND TIGHENNIF HOMININ MANDIBLES
À partir de ce travail, nous concluons donc que later included into Homo erectus (Le Gros Clark 1964),
les hominidés de Tighennif et de la Gran Dolina-TD6 a denomination that is nowadays preferred by some
appartiennent à deux lignées différentes. Les hominidés authors to include other North African human fossils
de Tighennif pourraient être réunis avec les spécimens such as those from Thomas Quarry and Sidi Abderra-
contemporains de Thomas Quarry et de Oulad Hamida 1 haman (Rightmire 1990). Alternatively, other researchers
et peut-être avec d’autres plus tardifs comme ceux de have preferred to use the denomination H. mauritanicus
Sidi Abderrahaman, Salé, et Rabat [Kebitat] du Nord de (Hublin 2001; Stringer 2003). Furthermore, Hublin (2001)
l’Afrique dans une même sous-espèce de H. ergaster, and Stringer (2003) have considered that the human
i.e. H. ergaster mauritanicus, qui serait le résultat d’une fossils recovered from Aurora Stratum of the level TD6 in
évolution locale en isolement relatif dans cette région the Gran Dolina site of the Sierra de Atapuerca (Carbonell
de l’Afrique. Les hominidés de la Gran Dolina-TD6 et al. 1995) could be also included in H. mauritanicus
appartiennent à un groupe exclusif de l’Eurasie, d’après (considering the priority of the name assigned by
la morphologie mandibulaire et dentaire. Arambourg). However, these authors have not presented a
formal study to test their hypothesis.
The cave of Gran Dolina (TD) in the Railway Trench
of the Sierra de Atapuerca (northern Spain) contains an
INTRODUCTION
18 meters high section formed by a succession of eleven
The sand pit of Tighennif (formerly Ternifi ne) levels, probably deposited from the late Early Pleistocene
to the end of the Middle Pleistocene (Parés and Perez-
is located in the village of Palikao, 22 km east from
González 1999). The lowermost stratigraphic levels, TD1
Mascara in Algeria. Excavations during the 1950s in the
and TD2, contain sediments of interior facies typical of
Middle Pleistocene sandy sediments of an ancient small
a closed cave. The paleontological record (pollen and
lake basin yielded thousands of mammal fossils, as well
faunal remains) is continuous from TD3 to TD11 (except
as Oldowan and Acheulean stone tools (Arambourg
TD9), and some levels (TD4, 5, 6, 7, 10, and 11) contain
1954a, b). The Tighennif site is particularly known by the
fi nding of three well preserved mandibles, some isolated abundant stone tools. During the 1994-96 fi eld seasons,
near one hundred human fossils were found together with
teeth and one parietal bone, which represent the oldest
more than one thousand macrovertebrate fossil remains,
fossil hominins recovered so far from northwest Africa.
as well as 268 lithic artefacts included in the Mode 1
The lithic assemblage can be included in the Acheulean
Technology (= Oldowan tradition) (Carbonell et al.
technological complex and consists of choppers, chopping
1995). The human remains were later included in a new
tools, polyhedrical spheroids, trihedrons, cores, retouched
pieces, large and small fl akes, hand axes and cleavers. The Homo species, H. antecessor, considered as possible last
common ancestor to Neanderthals and modern humans
artefacts are made of quartzite, sandstone, limestone, and
fl int (Sahnouni, Derradji in press) lineages (Bermúdez de Castro et al. 1997). These fossils
have been fi rmly dated to the late Early Pleistocene
According to the paleontological and paleomagnetic
(Parés, Pérez González 1995, 1999; Falguères et al.
data, the age of Tighennif has been estimated to be ca.
1999). During the 2003 to 2006 fi eld seasons new fi ndings
700 kyrs at the early Middle Pleistocene (Geraads et al.
have signifi cantly enlarged the hominin fossil hypodigm
1986). Paleomagnetic studies made on the lower deposits
of H. antecessor (Carbonell et al. 2005). The TD6 human
indicated normal polarity, and thus they may be correlated
with the Brunnhes Chron. However, Sahnouni and assemblage now includes three mandibular remains and a
Derradji (in press) do not rule out the correlation with the large dental sample.
Jaramillo Subchron. These authors have pointed out the Although the information is obviously limited, we
similarities between the Tighennif and Thomas Quarry 1 have the possibility of comparing dental and mandibular
(level L) sites concerning the macro-mammals and lithic evidence from two near and almost contemporaneous
assemblage. The age of the last site has been recently Pleistocene populations separated by the Gibraltar Strait.
estimated to be ca. 989 kyrs (Rhodes et al. 2006). The aim of this report is therefore to make a comparative
The Tighennif human fossil remains were originally study of the mandibles and teeth recovered from the
classifi ed as Atlanthropus mauritanicus by Arambourg two sites in order to make possible inferences about the
(1954b) together with other remains such as those from phylogenetic story of the hominins represented in the
Rabat (Thomas, Vallois 1977). These hominins were also Tighennif and Gran Dolina-TD human fossil samples.
152 J.M. BERMÚDEZ DE CASTRO, M. MARTINÓN-TORRES, A. GÓMEZ-ROBLES, L. PRADO, S. SARMIENTO
THE TD6 AND TIGHENNIF SPECIMENS
Figures 1 and 2 show the three Tighennif mandibles.
In Tighennif 1 both the left and right ramus are damaged
and some features of this region are not available. Both the
right and left P3-P4 and M1-M3 series are in situ, as well
as the root and a part of the crown of the lateral incisors.
Tighennif 2 consists of the left half of a well preserved
mandible (including the symphysis) that preserves the
P3-P4 and M1-M3 series in situ. The most complete
mandible is Tighennif 3, which preserves the right I2, C,
Fig. 2—Left lateral view of Tighennif 3.
P3, P4, M1, M2, and M3, and the left P4, M1, M2, and M3
Fig. 2 - Vue latérale gauche de Tighennif 3.
in situ. The three specimens belong to adult individuals.
The TD6 human assemblage includes three
mandibular specimens: ATD6-5 is a fragment of the right
side of the mandibular body of a juvenile specimen, with
aa
M1, M2, and M3 in situ, the latter still enclosed in its
alveolus (fi gure in Rosas, Bermúdez de Castro 1999).
bb
Fig. 3a—Lateral view of ATD6-96;
b—medial view of ATD6-96.
Fig. 3a - Vue latérale de ATD6-96 ;
b - vue médiale de ATD6-96.
Fig. 1a—Tighennif 1; b—Tighennif 2. The photographs
This specimen, as well as some isolated maxillary
on the originals of Tighennif mandibles were obtained
by Marcel Bovis in 1954 and donated to and mandibular teeth, belongs to Hominid 1 (the holotype
Emiliano Aguirre by Camille Arambourg. of H. antecessor). ATD6-96 (Hominid 7) is the left half of
Fig. 1a - Tighennif 1 ; b - Tighennif 2. Les photographies a small and gracile mandible of an adult individual with
des originaux des mandibules de Tighennif ont été the premolars and molars in place (fi g. 3) (Carbonell et al.
obtenues par Marcel Bovis en 1954 et offertes
à Emiliano Aguirre par Camille Arambourg. 2005). The third specimen, ATD6-113 (Hominid 10) is a
153 GRAN DOLINA-TD6 AND TIGHENNIF HOMININ MANDIBLES
fragment of the left side of the mandibular body with a internal side. Height of the mandibular body was taken
part of the ramus, which preserves the M2 and M3 in situ between the basal and the alveolar borders, keeping the
and it was found in the 2006 excavation season. The third callipers parallel to the external side of the mandibular
molar is fully erupted and exhibits slight occlusal wear body. A scatter biplot is provided in order to illustrate
at the level of the protoconid and metaconid that only the variability of the mandibular corpus robustness (the
affects the enamel. Thus, this mandible may correspond to corpus width at M1 level divided by the corpus height
a young adult. The increased TD6 human hypodigm allow at the same level) against the corpus height. In the D211
us now to do a proper comparison with the Tighennif Dmanisi mandible we had to do an estimation of the corpus
specimens. height since the corpus base is broken at the M1 level.
The slope of the regression line, the correlation coeffi cient
and the p-value of the regression was calculated for those
groups with a sample size large enough to obtain the
METHODS
parameters: H. ergaster, H. erectus, H. heidelbergensis
and H. neanderthalensis.
For comparative purposes, a large number of fossil
The mesiodistal (MD) and buccolingual (BL)
Homo mandibles and teeth were included in our study
dimensions were obtained in the TD6 and Tighennif teeth,
(tables I, II, III). Observations and data were recorded on
according to the technique quoted in Bermúdez de Castro
the following original fossil samples: Atapuerca-Sima de
(1993).
los Huesos, Arago, Atapuerca-TD6, Tighennif, Sangiran,
and Dmanisi, as well as on some high-quality casts. We
have grouped the specimens into different species in
COMPARATIVE ANALYSIS
accordance with the criteria of different authors (e.g.
Wood 1992; Schwartz, Tattersall 1999 ; Tattersall 2000). The mandibles
Thus, we have differentiated taxonomically the Early
Pleistocene East African hominins (H. ergaster) from the As it has been previously pointed out, the Tighennif
Asian Pleistocene hominins (H. erectus). Furthermore, mandibles are extremely massive and robust (Howell
we have distinguished between the Chinese and Javanese 1960; Arambourg, Hoffstetter 1963), whereas the TD6
H. erectus because, according to different authors, they specimens are particularly gracile. Nevertheless, before
show some regional and temporal variations (Rightmire comparing the dimensions of the TD6 mandibles with
1990; Grimaud-Hervé et al. 2000; Antón 2002; Babá those of Tighennif it is necessary to consider the possible
et al. 2003). The species H. heidelbergensis includes the effect of sexual dimorphism in the specimens size. Judging
European Middle Pleistocene hominins, according to the by the remarkable size difference between the teeth of
criteria expressed by Arsuaga et al. (1999) (but see also Hominid 1 (ATD6-5) and Hominid 10 (ATD6-113) from
an alternative opinion in Rightmire 1996). The Dmanisi TD6 on one hand, and Hominid 7 (ATD6-96) on the other
mandibles were not included in a particular species hand (table I), we can assume that the Hominid 1 and 10
because their taxonomic status are a current disputed are probably males, whereas Hominid 7 would be female.
matter among the specialists (e.g. Gabunia,Vekua 1995; The dimensions of the upper canine of Hominid 1 are at
Rosas, Bermúdez de Castro 1998; Gabunia et al. 2002; the upper limit of the variation range of the genus Homo,
Rightmire et al. 2005). which supports the sex status of this immature individual.
Features and data of several mandibular specimens Thus, we think that males and females are present at
were confi rmed or obtained from tables 5, 6, and 7 of least in the TD6 sample. Interestingly, the thickness and
Rosas (2001) and table 2 of Rosas and Bermúdez de height of the ATD6-113 mandible (the largest of the Gran
Castro (1999). Our methodological approach is cladistic Dolina-TD6 sample) are similar to those of Tighennif 2
and the description, defi nition and evaluation of the (the smallest of the North African sample) (table II).
status (primitive or derived) of the mandibular features The thickness and height of the mandibular body of
(table IV) are taken from Rosas (2001). Thickness of the the Tighennif mandibles are similar to those of H. erectus
mandibular body was measured at the M1 and at lateral from Java, whereas the same variables of the TD6
prominence location (tables II, III). In both cases, the specimens are similar to those of the H. erectus from China
measurement was taken between the external side of the (table III). The ramus breadth of the ATD6-96 mandible
mandibular body and the most prominent point of the (40.6 mm) is lower than those of Tighennif 2 (45.8 mm)
154 J.M. BERMÚDEZ DE CASTRO, M. MARTINÓN-TORRES, A. GÓMEZ-ROBLES, L. PRADO, S. SARMIENTO
and Tighennif 3 (47.3 mm). The height of the ramus in Howell (1960), the total length and bicondylar dimensions
ATD6-96 is 60 mm, whereas in Tighennif 2 and 3 this of Tighennif 3 exceed those of the Mauer mandible.
measurement is 78.5 and 85.6, respectively. According to
Gran Dolina-TD6
Tighennif 1 Tighennif 2 Tighennif 3 H 1 H 7 H 10
Side Tooth MD BL MD BL MD BL MD BL MD BL MD BL
R 12 (5.6) 7.3

L 12 7.0 7.8

R C 7.8 10.3
L C 8.1 10.0

R P3 8.5 10.2 8.8 10.2 8.8 10.6
L P3 8.4 10.1 8.7 11.2 8.0 9.9
R P4 8.4 10.2 8.2 10.2 8.2 10.2
L P4 8.4 10.0 9.2 11.4 7.7 10.2 7.6 9.4
R M1 13.2 12.6 12.4 12.0 12.2 11.8
L M1 12.9 12.3 14.0 13.0 12.4 12.2 10.5 11.0
R M2 13.00 13.2 12.6 12.0 13.5 12.0
L M2 13.2 13.2 14.0 13.6 12.6 12.2 12.3 11.0 13.1 11.4
R M3 12.4 12.2 12.3 11.5
L M3 12.4 12.2 13.2 12.7 12.0 11.0 9.2 8.8 12.5 10.6
Isolated teeth from Tighennif
R Lower I1 - 6.7
R Lower I2 7.1 7.0
L Lower C 9.0 10.0
L Upper dm1 8.3 9.8
L Upper dm2 10.3 11.7
R Upper M1-M2 12.0 14.5
Table I—Dental dimensions (mm) of the Tighennif and Gran Dolina-TD6 mandibles
and isolated teeth. H: Hominids 1 (ATD6-5 mandible and isolated teeth),
7 (ATD6-96 mandible) and 10 (ATD6-113 mandible)
from TD6. MD: mesiodistal; BL: buccolingual.
Tabl. I - Dimensions (mm) des dents isolées et des dents en place des mandibules de Tighennif et
Gran Dolina-TD6. H: Hominidés 1 (mandibule ATD6-5 et dents isolées),
7 (mandibule ATD6-96), et 10 (mandibule ATD6-113) de la Gran Dolina-TD6.
MD: mésio-distal ; BL : bucco-lingual.
155 GRAN DOLINA-TD6 AND TIGHENNIF HOMININ MANDIBLES
Tighennif 1 Tighennif 2 Tighennif 3 ATD6-5 ATD6-96 ATD6-113
T H T H T H T H T H T H
Corpus at M1 19.8 36.2 16.6 33.4 19.5 37.0 16.3 26.7 16.6 28.5 19.0* 31.0*
Corpus at LP 23.2 37.5 22.2 31.2 23.7 36.0 – – 18.6 27.5 20.0 32.0
Ramus breadth – 45.8 47.3 – 40.6 –
Ramus height – 78.5 85.6 – 60.0 –
Symphysis height 36.0 33.3 37.5 – – –
Table II—Measurements (mm) of the Tighennif and Gran Dolina-TD6 mandibles. T: thickness; H: height.
Tabl. II - Mesures (mm) des mandibules de Tighennif et de la Gran Dolina-TD6. T : épaisseur ; H : hauteur.
Species Thickness, mm (n) Height, mm (n) Robustness (n)
Homo habilis 19.7 ± 2.3 (5) 29.3 ± 2.2 (4) 64.5 ± 5.3 (4)
H. ergaster# 19.8 ± 1.4 (8) 31.2 ± 2.8 (8) 63.8 ± 5.0 (8)
H. erectus (Java)* 21.3 ± 3.9 (5) 38.8 ± 5.3 (5) 54.9 ± 5.8 (5)
H. erectus (China)** 16.5 ± 1.8 (7) 28.6 ± 3.3 (7) 58.0 ± 5.6 (7)
Tighennif 18.6 (3) 35.5 (3) 52.4 (3)
Sidi Abderraman 17.0 35.8 47.5
H. antecessor 17.3 (3) 28.7 (3) 60.2 (3)
H. heidelbergensis+ 16.4 ± 1.7 (16) 30.8 ± 3.4 (16) 53.4 ± 5.1 (16)
H. neanderthalensis++ 15.3 ± 1.7 (21) 32.1 ± 3.3 (21) 47.9 ± 5.1 (21)
Data are given as means ± SD
# This sample includes East African Early and Middle Pleistocene adult specimens KNM-ER 730, KNM-ER 731, KNM-ER 992, KNM-BK 67,
KNM-BK 8518, OH 22, OH 23, and OH 51.
* This sample includes the Sangiran mandibles 1B, 5, 8, and 9.
** This sample includes the mandibles from Zhoukoudian, Lantian, Hexian, and PA86.
+ This sample includes the Arago, Atapuerca-Sima de los Huesos, Mauer and Montmaurin specimens (see Rosas, Bermúdez de Castro 1999).
++ See Rosas and Bermúdez de Castro (1999) for the composition of this sample.
Table III—Measurements of the mandibular body at the level of M1 in some Homo species and specimens.
Tabl. III - Mesures du corps mandibulaire (épaisseur, hauteur et robustesse) au niveau de la première molaire (M1)
chez quelques espèces et spécimens du genre Homo.
156 J.M. BERMÚDEZ DE CASTRO, M. MARTINÓN-TORRES, A. GÓMEZ-ROBLES, L. PRADO, S. SARMIENTO
Mandibular features State A State B State C
1 - Position of mental foramen P3-P4 P4-M1 M1
2 - Position of the anterior marginal tubercle P3-P4 P4-M1 M1
3 - Position of the lateral prominence M2 M2-M3 M3
4 - M3 in relation to ramus (retromolar space) Coverted Partially coverted Uncovered
5 - Inclination of the retromolar area Vertical Oblique Horizontal
6 - Position of mylohyoid line in relation to alveolar margin at
Low Medium High
M3 level
7 - Trajectory of mylohyoid line in relation to alveolar margin Parallel Intermediate Diagonal
8 - Relief of masseteric fossa Deep Shallow Flat
9 - Relief of pterigoyd fossa Shallow Flat –
10 - Gonion profile Expanded Regular Truncated
11 - Position of condyle in relation to mandibular notch Lateral Medial –
12 - Posterior subalveolar fossa Shallow Moderately hollowed Deep
13 - Alveolar prominence Present Absent –
Table IV—State of the mandibular features analyzed in table V [partially modifi ed (features and scores)
from Rosas (2001)].
Tabl. IV - État des caractéristiques mandibulaires analysées dans le tableau V [modifi é partiellement
(caractéristiques et état) d’après Rosas (2001)].
Figure 4 shows a clear relationship between outliers such as Sangiran 6. However, H. ergaster line
increasing robusticity and decreasing corpus height in presents a signifi cant regression model (p-value = 0.003),
Pleistocene hominin mandibles. The plot shows two with a correlation coeffi cient of 0.735.
pairs of regression lines clearly differentiated. The The slopes of the regression lines are almost the same
lowest two lines correspond to H. neanderthalensis for all the groups analyzed, but the lines corresponding
and H. heidelbergensis, although they do not show to the more primitive groups are above the ones
especially high correlation coeffi cients (0.264 and 0.318, corresponding to later species, such as H. heidelbergensis
respectively). These lines are almost superimposed, and H. neanderthalensis. This could imply that in more
with a slope of -0.744 (p-value = 0.020) and a constant recent groups, a given corpus width tends to be related
of 71.87 for H. neanderthalensis and a slope of to a lower corpus height, whereas the same corpus width
-0.835 (p-value = 0.023) and a constant of 79.15 would correspond to a higher height in more primitive
for H. heidelbergensis. The upper lines correspond groups.
to H. erectus and H. ergaster but, although they are Regarding to the species analyzed in this study, we
superimposed, the line corresponding to H. erectus is can observe that Tighennif and Sidi Abderraman specimens
not signifi cant (p-value = 0.301) due to the presence of are located nearby the H. ergaster regression line (they
157 GRAN DOLINA-TD6 AND TIGHENNIF HOMININ MANDIBLES
Fig. 4—Robusticity index versus mandibular height. A. Regression line for H. ergaster, H. erectus, H. heidelbergensis and
H. neanderthalensis. B. Confi dence intervals at 95% for H. ergaster and H. neanderthalensis. It is important to note the position
of Tighennif and Gran Dolina-TD6 in the plot (see text for explanation), as well as the big difference in the corpus index/corpus
height ratio between D 2600 and both D 211 and D 2735 Dmanisi mandibles, as it was previously noticed (Skinner et al. 2006;
Bermúdez de Castro et al. submitted). D 2735 is located over H. ergaster line, whereas D 211 is located on the H. neanderthalensis
line. Both individuals, however, display low corpus height compared with their corpus width, whereas D 2600 is out of the main
point cloud due to its elevated corpus height. NAFMP: North Africa Middle Pleistocene fossils (Tighennif and Thomas Quarry).
Fig. 4 - Graphique qui met en rapport l’indice de robustesse et la hauteur du corps mandibulaire. A. Ligne de régression pour H. ergaster, H. erectus,
H. heidelbergensis et H. neanderthalensis. B. Intervalles de confi ance à 95 % pour H. ergaster et H. neanderthalensis. Notez la position des spécimens
de Tighennif et de la Gran Dolina-TD6 dans le graphique (voir le texte pour une explication détaillée) ainsi que la grande différence entre D 2600,
d’une part, et D 211 et D 2735 d’autre part, comme l’ont signalé d’autres auteurs (Skinner et al. 2006 ; Bermúdez de Castro et al. soumis). D 2735
est placé au-dessus de la ligne de H. ergaster, tandis que D 211 est placé sur la ligne de H. neanderthalensis. Ces deux spécimens de Dmanisi
présentent cependant une grande épaisseur du corps mandibulaire par rapport à la hauteur, tandis que D 2600 se situe à l’écart, du fait de la
grande hauteur de son corps mandibulaire. NAFMP : fossiles du Pléistocène moyen du Nord de l’Afrique (Tighennif et Thomas Quarry).