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Djillali Hadjouis is a specialist in human and animal anatomy of populations of the past. He has been, in turn, associate professor at universities, research director, departmental archaeologist and lecturer, training dozens of students from Europe, Africa and Asia.
Introduction xi
Part 1. The Skull of Fossil and Present-day Quadruped Vertebrates: Craniofacial Structure and Postural Balance 1
Chapter 1. Proboscideans: The Mammoth (Mammuthus primigenius) 3
1.1. Chronological, geographical and morphological indications of the species 3
1.2. Mammoth discoveries in Île-de-France 5
1.3. A young mammoth in Maisons-Alfort 5
1.4. A woolly mammoth skull in the reserves 6
1.5. A mammoth skull with removed tusks 7
1.6. A particular tooth eruption 8
Chapter 2. Equidae 11
2.1. The horse (Equus caballus) 11
2.1.1. Chronological, geographical and morphological indications of species 11
2.1.2. A fossil horse in Africa: paleogeographic and biostratigraphic distributions 15
2.1.3. The postural balance of Equidae 17
2.1.4. Joint pathologies in service horses 18
2.1.5. Introduction to animal bone pathologies and zoonoses 20
2.1.6. The horse's status over the centuries 20
2.2. The donkey (Equus asinus) 21
2.2.1. Chronological, geographical and morphological indications of species 22
2.2.2. The status of the donkey over the centuries 23
Chapter 3. Bovidae 25
3.1. Aurochs (Bos primigenius) 25
3.1.1. Chronological, geographical and morphological indications of species 25
3.1.2. Cattle (Bos taurus) 27
3.1.3. The status of cattle over the centuries 28
3.2. The bison (Bison priscus): chronological, geographical and morphological indications of the species 28
3.3. The buffalo (Syncerus antiquus) 29
3.3.1. Chronological, geographical and morphological indications of the current Syncerus and Bubalus buffaloes 29
3.3.2. Chronological, geographical and morphological indications of fossil species 30
3.3.3. Bos/Syncerus dental distinction criteria 35
3.3.4. Postural balance and paleoecology of Bovidae 38
3.3.5. Polymorphism and dimorphism in Bovidae 39
3.3.6. Osteoarticular abnormalities and bone pathologies in Bovidae 41
3.4. The common eland (Taurotragus oryx) 43
3.4.1. Chronological, geographical and morphological indications of species 43
3.4.2. Posture and locomotor adaptation 46
3.5. The hartebeest (Alcelaphus buselaphus) 48
3.5.1. Chronological, geographical and morphological indications of species 48
3.5.2. Postural balance 49
3.6. Gazelles (Gazella) 50
3.6.1. Chronological, geographical and morphological indications of species 50
3.6.2. Postural balance 51
Chapter 4. Cervidae 53
4.1. The red deer (Cervus elaphus) 53
4.1.1. Chronological, geographical and morphological indications of species 53
4.1.2. The status of deer developing over the centuries 58
4.2. The Algerian thick-cheeked deer (Megaceroides algericus) 59
4.2.1. Several species from Europe, the Mediterranean islands and one species from the Maghreb 60
4.2.2. Size of Megaceroides algericus 63
Chapter 5. Suidae 65
5.1. The wild boar (Sus scrofa) 65
5.1.1. Chronological, geographical and morphological indications of species 65
5.1.2. The status of the boar over the centuries 67
5.1.3. Postural balance of the boar 67
5.2. The warthog (Phacochoerus aethiopicus or africanus) 70
5.2.1. Chronological, geographical and morphological indications of species 71
5.2.2. A particular tooth eruption 74
5.2.3. Postural balance of the warthog 76
5.2.4. Pathologies in warthogs 77
5.2.5. A catastrophic mortality curve 78
Chapter 6. Carnivores 81
6.1. The lion (Panthera leo) 81
6.1.1. Chronological, geographical and morphological indications of the species 81
6.1.2. Occlusal posture and the lion's balance on the ground 83
6.2. The panther or leopard (Panthera pardus) 84
6.2.1. Chronological, geographical and morphological indications of species 85
6.2.2. Occlusal posture and postural balance of the panther on the ground 85
6.3. The spotted hyena (Crocuta crocuta): chronological, geographical and morphological indications of the species 87
6.4. The striped hyena (Hyaena hyaena) 89
6.4.1. Chronological, geographical and morphological indications of species 89
6.4.2. Occlusal posture and postural balance of hyenas on the ground 90
6.5. The cave bear (Ursus spelaeus) and the brown bear (Ursus arctos): chronological, geographical and morphological indications of the species 93
6.6. The wolf (Canis lupus): chronological, geographical and morphological indications of the species 95
Chapter 7. Lagomorphs: The Hare (Lepus capensis) 99
7.1. Chronological, geographical and morphological indications of the species 99
7.2. The status of the hare over the centuries 101
Part 2. The Skull of Fossil Bipedal Vertebrates: Craniofacial Structure and Postural Balance 103
Chapter 8. Primates 105
8.1. Occlusal posture, quadrupedic and verticalization of the Hominoid body 106
8.2. Work in dentofacial orthopedics and embryogenesis 108
Chapter 9. Hominoids 111
9.1. Kenyapithecus 112
9.2. Nacholapithecus 113
9.3. Otavipithecus namibiensis 113
Chapter 10. From Hominoids to Hominids 115
10.1. Ardipithecus ramidus kadabba 115
10.2. Praeanthropus tugenensis (= Orrorin tugenensis) 116
10.3. Sahelanthropus tchadensis 116
10.4. Ardipithecus ramidus 117
10.5. Praeanthropus africanus (= Australopithecus anamensis) 118
Chapter 11. Australopithecus 119
11.1. Australopithecus afarensis 120
11.2. Australopithecus africanus 120
11.3. Australopithecus bahrelghazali 120
11.4. Australopithecus garhi 121
11.5. Paranthropus robustus 121
11.6. Australopithecus aethiopicus 121
11.7. Australopithecus boisei 122
Chapter 12. The Genus Homo 123
12.1. Homo habilis 126
12.2. Homo rudolfensis 126
12.3. Homo ergaster and Homo erectus 127
12.4. Homo georgicus 128
12.5. Homo neanderthalensis 129
12.5.1. Plesiomorphic and autapomorphic morphological features 129
12.5.2. Non-Sapiens craniofacial dynamics and posture 130
12.5.3. A permanent labidodental joint 130
12.5.4. The asymmetry of fossil pieces 133
12.6. Homo sapiens 135
Chapter 13. Migration and Paleogeographic Distribution of the Homininae 137
13.1. Australopithecus and Homo habilis: regional African migrations 137
13.2. Homo ergaster and Homo erectus: the first great African-Eurasian journey 139
13.3. Homo neanderthalensis: a Eurasian traveler 141
13.4. Homo sapiens: the second great conquest voyage on all continents 141
Part 3. The Skull of Homo sapiens in All its Diversity 145
Chapter 14. The Craniofacial Puzzle in Motion 147
14.1. Normality and its boundaries with the abnormal and the pathological 147
14.2. The importance of interpreting or reinterpreting (Le Double 1903, 1906) 148
14.3. Craniofacial structural mechanics and dynamics 149
14.3.1. Biodynamics of vault bones 150
14.3.2. Biodynamics of the temporal bone 151
14.3.3. Biodynamics of the occipital bone 151
14.3.4. Biodynamics of the sphenoidal bone 152
14.3.5. Biodynamics of the maxillary bone 152
14.3.6. Biodynamics of the mandibular bone 154
Chapter 15. The Basics of Structural Analysis 157
15.1. Analysis tools using imaging 157
15.2. Maxillo-mandibular dysmorphoses 159
x The Skull of Quadruped and Bipedal Vertebrates
15.2.1. Angle's classification 160
15.3. History of structural mechanics: from geometry to imagery 161
15.3.1. The initiators 161
15.3.2. FDO orthopedists and orthodontists 163
15.3.3. Osteopaths 165
15.3.4. Recent work in human paleontology and paleoanthropology 166
Chapter 16. Identification of Malformation 169
16.1. Craniostenosis, a history of sutures 169
16.2. Craniofacial asymmetries 172
16.2.1. Examples of craniofacial asymmetries 174
16.2.2. The importance of the spine and its effects in basic cranial equilibrium or disequilibrium 180
16.3. Psalidodontia or labidodontia? 181
16.3.1. The behavior of the dental articulation of juvenile Pleistocene and Holocene populations in the Maghreb and the Sahara 184
16.3.2. Dental articulation and extraction of the incisors 187
16.4. Para-masticatory functions of Homo sapiens in Algeria 190
16.5. Occlusal equilibrium and adaptation of regional morphotypes 193
16.5.1. In the Paris Basin 193
16.5.2. In the Maghreb countries 198
16.5.3. Occlusal balance and the regional morphotype in the Maghreb and Sub-Saharan Africa 199
Chapter 17. Ignored Pathologies 205
17.1. Extremely rare craniofacial pathologies 205
17.1.1. Crouzon syndrome 205
17.1.2. Marfan syndrome 205
17.1.3. Cranial thickening and Albers-Schönberg's disease 206
17.1.4. Torticollis 206
17.1.5. Parietal thinning 207
17.1.6. Scurvy 208
17.2. The oldest therapeutic practice: trepanning 209
Conclusion 211
References 213
Index 235
Super-Order Proboscidea.
Order Elephantoidea.
Family Elephantidae.
Genus Mammuthus Brooks, 1828.
Species Mammuthus primigenius Blumenbach, 1799 (woolly mammoth).
In the species of the Elephantidae Family, the orbits are very advanced on the antero-posterior axis of the skull and open in front of the jugal teeth; these are anatomical characteristics of animals without snouts. In general, in Mammals, the orbits open above the last molars (Lecointre and Le Guyader 2001).
The nose and upper lip are replaced by a flexible tube used for breathing, drinking, picking things up, etc. The lower canines are lost and the upper incisors are transformed into tusks that are continuously developing (Figure 1.1).
The species belongs to one of the three related genera regrouping mammoths and elephants (Mammuthus, Elephas and Loxodonta) included in the Family Elephantidae and classified in the Super-Order Proboscidea. These trunk bearers have provided no less than 170 fossil species, the oldest dating back 55 million years (Eocene). The first Elephantidae appeared in Africa 7 million years ago, the first mammoths 3-4 million years ago. Great confusion reigned for a long time among specialists for the classification of the mammoth and its origin. Today everyone is unanimous in classifying the first mammoths in Africa.
The arrival of mammoths in Europe and Asia took place around 2.6 million years ago. Three main Euro-Asian species followed one another. The southern mammoth (Mammuthus meridionalis), known in Saint-Vallier, Senèze and Chilhac, had a size exceeding 3 meters at the withers, large slightly curved tusks and jugal teeth whose hypsodontic character was still weak. The steppe mammoth (Mammuthus trogontherii), descendant of the previous one, was known in Sussenborn and Mosbach (Germany), in Abbeville and Saint-Acheul (France) and in Great Britain. With a height of more than 4 meters at the withers, this mammoth was considered the largest species in Europe. The woolly or Siberian mammoth (Mammuthus primigenius) is the best known. It appeared about 190,000 years ago and remains the most documented of all fossils due to the many discoveries of whole animals preserved in the permafrost (Guérin 1996a).
Figure 1.1. Skull of a young present-day Asian elephant in left lateral view. Note the position of the orbits above the maxilla and not behind it (© Éditions Belin/Dominique Visset)
The Fifth International Conference on Mammoths and their Families, held in 2010 in Le Puy-en-Velay, brought together the discoveries of this trunk fossil in more than 20 countries of Eurasia, Africa and America. Even though the Île-de-France region was not represented at this meeting, there have been a large number of remains of woolly mammoths in the departments of Hauts-de-Seine, Seine-Saint-Denis, Val-de-Marne, Yvelines, Essonne, Seine-et-Marne and Val-d'Oise, most of which were cleared during the 19th and 20th centuries as a result of quarrying on the outskirts of Paris. The latter also had its share of discoveries, sometimes complete skeletons, such as that of the Montholon Square near the Montmartre cemetery. In the department of Val-de-Marne, several bony and dental remains of woolly mammoths (M. primigenius) have been found since the end of the 19th century on the banks of the Seine and its confluence with the Marne (Ardouin et al. 2009; Hadjouis 2020a). Thanks to the land development of the last 40 years carried out along the river banks, preventive archeological operations have brought to light new discoveries in well-dated biostratigraphic contexts.
Although the remains of ancient mammoths and elephants unearthed in this small department of south-eastern Paris have been numerous (remains preserved at the Muséum national d'Histoire naturelle, Musée de Saint-Maur-des-Fossés and Musée d'antiquités nationales), most of them were transported and deposited on the banks of the Seine or in the loop of the Marne (Le Perreux, Nogent-sur-Marne, Saint-Maur-des-Fossés, Créteil, Valenton, etc.) without great chronostratigraphic precision.
Appearing in Eastern Siberia around 800,000 years ago, the woolly mammoth's existence was known in Western Europe 200,000 years ago in chronocultural contexts of the Middle Paleolithic. With the exception of some spectacular specimens, it is mainly mandibles, cranial portions or isolated teeth that are found during archeological excavations or chance discoveries. Among the fossils recently dated by radiometric methods (uranium/thorium and carbon 14), two stand out: the young mammoth from Maisons-Alfort and the skull from Bonneuil-sur-Marne.
The development operations of the Seine-Marne confluence, which began in the 1980s as part of the Seine-Amont project, have highlighted major sites such as Alfort 1 in Maisons-Alfort, located on the left bank of the Marne, a few hundred meters from its confluence with the Seine. One of the plots excavated in 1995 revealed a Neanderthal occupation of the Middle Paleolithic within the alluvial formations (Durbet et al. 1997). The association of identified animal species (aurochs, bison, red deer, Mosbach horse, wolf, mammoth) and the presence of a lithic industry shaped by Neanderthals allowed the reconstruction of paleoenvironments suggesting a cold and dry glacial climate. Several datings on fossil remains ranging from 160,000 to 200,000 years BCE give an image of a glacial environment in the Paris region inhabited by both humans and animals.
Usually, the attribution of Proboscidian species (Elephas, Loxodonta and Mammuthus) is based on complete third molars. It is indeed this last molar which delivers the greatest number of distinctive dental criteria (tooth shape, number of blade teeth and their index, enamel character and their plicature, sinus morphology, etc.). The milk tooth of the Maisons-Alfort baby mammoth was an incomplete lower third milky premolar (D3) with four blades (length: 36.2 mm; height: 44.6 mm).
The inventory of Vertebrate faunas carried out at the request of the director of the Val-de-Marne Departmental Archaeology Laboratory, Philippe Andrieux, in the 1990s included a fossil, which exceeded in volume and significance all other bone and dental remains. The skull without mandible of a woolly mammoth, perfectly preserved, had been waiting to be studied since 1923.
Without the short note Les gros blocs quaternaires du port de Bonneuil by Paul Lemoine and Teilhard de Chardin published in the journal La Nature in 1923, no one would know the exact origin of this Propboscidian. The authors describe the work carried out on Barbière Island, a commune of Bonneuil, located between the Marne and Morbras rivers, in order to dig the future coal port of Paris. The dredging of alluvium and blocks of local origin (coarse limestone with ceriths, Champigny limestone, Brie millstone and Fontainebleau sandstone) had yielded the fossil remains of M. primigenius and Bos. The woolly mammoth skull discovered during these dredging operations was not described in this note, but was obviously part of the port works.
It was in the 2000s that we studied it, starting with radiocarbon dating. C14 analyses of the radiocarbon laboratory of Villeurbanne gave dates around 45,000 years BCE, the Middle Paleolithic period. Although more recent than the dates of the Middle Paleolithic site of Alfort 1 (190,000 BC), which also yielded mammoth remains, the fact remains that the skull of Bonneuil figures as a spectacular piece within the Quaternary bestiary of Île-de-France. Although it shows traces of anthropic activity that suggest human intervention, there is nothing to distinguish between hunting and natural death. Similarly, the intervention on its carcass must have taken place in two stages after death.
Another individual was found in the same conditions in Bonneuil-sur-Marne, notably a tusk with traces of anthropogenic cutting.
As mentioned above, the Bonneuil-sur-Marne fossil was not found in welldocumented stratigraphic levels or in a chronocultural context of the Paris Basin. However, its fortuitous discovery during the dredging of the coal port works in 1923 does not in any way detract from its exceptional character. The 45,000 year dating corresponds to a period of cold and dry climate in the Middle Paleolithic. Anthropic indications found on this skull without a mandible suggest hunting practices or the recovery of tusks from a dead animal.
The morphological criteria found on the maxillary teeth show the typical characters of the woolly mammoth (weak enamel with sinuous sinuses, high hypsodontia index and strongly folded enamel ribbons (Hadjouis 2016b)). The age given to this animal at the time of death is estimated to be between 22 and 34 years. It is based on criteria such as the partial wear of the molars, the clearing of their roots and the absence of wear of the last molar, some of whose distal blade teeth were still embedded in the alveolus. Three...
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