Article

Soft tissue reconstruction of Homotherium latidens (Mammalia, Carnivora, Felidae). Implications for the possibility of representations in Palaeolithic art

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Abstract

We reconstruct the life appearance of the head and body of the sabretoothed felid Homotherium latidens through the study of osteological correlates of soft tissue attributes, incorporating data from the dissection of several large felids and using the Extant Phylogenetic Bracket to infer the condition of unpreserved attributes where morphological evidence is inconclusive. Our reconstruction shows that Homotherium would have differed from modern felids in aspects of the general proportions of the head, having a straighter dorsal outline and a long, square muzzle with an angular ‘‘chin’’, although large pantherines may mimic to some degree the angular shape of the machairodontine mandibular symphysis with the growth of long hair in the chin area. The tips of the sabres of Homotherium would have been visible in life, protruding beyond the lips. Our reconstructed body proportions of Homotherium imply a sloping back. These conclusions contradict a previous proposal that postulated a different soft tissue anatomy for Homotherium, and which led to interpretation of a Palaeolithic figurine as a rendering of the sabretooth. Our results suggest that the figurine in question is a depiction of a cave lion, and that there is no single known representation of a machairodont in prehistoric art. The implications for our current understanding of the Late Pleistocene large-carnivore guild are discussed.

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... In most previous accounts of Middle Pleistocene H. latidens, the distal humeri were figured, but in no case was the most important defining character discussed, although it was also used in soft tissue fore limb reconstructions by Ant on et al. (2009). Based on the complete and well-preserved Plio-/Early Pleistocene H. crenatidens humerus figured by Ballesio (1963) ( Fig. 3.1), the supracondylar ridge is absent in this genus, which does not allow a rotation of the paw (cf. ...
... Based on the complete and well-preserved Plio-/Early Pleistocene H. crenatidens humerus figured by Ballesio (1963) ( Fig. 3.1), the supracondylar ridge is absent in this genus, which does not allow a rotation of the paw (cf. Ant on et al., 2009). Here, in one example figured ( Fig. 4A), for a small lion (most probaly female) from the Late Pleistocene of the Hermann's Cave (Harz Mountain Range), the ridge is strong and about 2e3 mm wide e a level of development similar to that seen in modern and Middle to Late Pleistocene lions (P. ...
... On this ridge, the rotator musculature (Brachiradialis) for the lower forelimb is attached to the radius (e.g. Frewein and Vollmerhaus, 1994;Ant on et al., 2009) ( Fig. 4A). This allows lions to catch its prey with the forelimb paws ( Ant on et al., 2009). ...
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Four new saber-tooth cat (Homotherium) sites in Germany with new dental and postcranial bone material are different in their taphonomic context: 1. The Archaeological Middle Palaeolithic (MIS 9e-Interglacial) Schöningen Lake site with remains of a cub carcass, 2. The Middle Palaeolithic (MIS 5e-9) Archaeological/cave bear den site of Balve Cave yielding a lower canine tooth of an older individual, 3. The Zoolithen Cave (MIS 3–9) cave bear/hyena den with one distal half humerus of an adult, 4. The Ketsch open air Rhine River terrace site which has provided another distal humerus of an adult saber-tooth cat. Whereas only the Schöningen site is precisely dated as Holsteinian Interglacial (approx. 330.000–315.000 BP), all other material seems to come from the same Middle Pleistocene warm period, or few younger Saalian interstadials (MIS 7a, e) deposits, and did not extend over the last MIS 7 glacial into the Late Pleistocene. Homotherium as hyena-like slow moving cat seems to have disappeared within the Saalian due to competition with other scavengers like Ice Age spotted and brown hyenas (Crocuta crocuta praespelaea/ultima and Pachycrocuta brunnea mosbachensis). The juvenile saber-tooth cat cub from Schöningen might be in archaeological context or represent only a carnivore kill. At the Zoolithen Cave, the single bone must have been imported into a hyena prey bone assemblage. The situation is possibly similar at the two other sites Ketsch and Balve Cave. The formerly described Schöningen “saber-tooth cat” humerus is revised, such as other opposite as lion humeri described material from different European sites. The presence of the well-developed supracondylar ridge distinguishes Homotherium well from Middle/Late Pleistocene lions Panthera leo (e.g. spelaea, fossilis). The Schöningen lion humrus has been chew-cut first most probably by a stripped hyena whose cutting scissor teeth produced a diagonal bite cut and P⁴/M¹ impact marks around the trochlea. 1–2 mm small, mostly triangular-oval bite marks on the lion humerus shaft compacta results from a second scavenger and not from “Neanderthal tool use”. Those bite mark sizes are produced mainly of the upper molar teeth of a the red wolf Cuon alpinus subsp. (or small fox Vulpes praecorsac), which were present in the region within the Holsteinian/Saalian.
... Nowadays, some fan artists have proposed a new facial restoration of Smilodon and other machairodontines. Despite previews works with solid conclusions about sabertooth felids appearance (Turner & Antón, 1997;Antón et al., 1998Antón et al., , 2009 there is a new attempt to recreate these animals. One of the most adventurous illustrators is Duane Nash which has hypothesized about a new kind of restoration for Smilodon and relatives. ...
... In this sense, some artists had published methodologies and instructions for reconstructing the appearance of fossil fauna (Scott, 1913;Paul, 1987;Antón and Sánchez, 2004). Other artists, among which Gregory S. Paul and Mauricio Antón stand out, have not just created images defining the species they represent but have also made multiple scientific publications focusing on their reconstruction (Paul, 1988a;Olshevsky, 1991;Antón et al. 1998Antón et al. , 2009Antón, 2007;Pérez González et al. 2009;Ansón and Hernández Fernández, 2013), even suggesting new hypothesis, like the defensive behavior of Triceratops pictured by Mark Hallett (Wexo, 1987), or anticipating scientific knowledge that can be subsequently verified by fossil evidence, as the predatory behaviour of Tyrannosaurus on Triceratops (Paul, 1988b;Erickson and Olson, 1996). ...
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Paleoart is a widely used term in the paleontological world. This word can be found in different media, used in relation to different kinds of artistic manifestations of paleontological theme. Paleoart has a long history and has helped paleontology to became one of the most popular sciences. The production of this artwork means that paleoartists must have high skills in both disciplines, arts and paleontology. Due to its scientist, artistic and cultural significance, paleoart and paleoartists must be recognized and valued in both paleontology and fine arts. In this situation paleoart needs a clear definition that distinguishes it from other artistic representations. Taking this into account, we have consulted to a series of paleontologists, and this survey has allowed us to outline such a definition. All the original artistic manifestations that pretend to reconstruct o depict prehistoric life according to the current knowledge and scientific evidence at the moment of creating the artwork can be considered paleoart.
... Quaternary Science Reviews j o u r n a l h o me p a g e : w w w . e l s e v i e r. co m/ lo ca t e / q u a s ci re v unique within Cenozoic palaeontology for not having named any new taxa throughout his career e although he came close with the revision of Paramachairodus and the revival of Promegantereon in collaboration with Mauricio Ant on and Manuel Salesa (Salesa et al. 2010). ...
... Each muscle was first created as a Blender new mesh (3D object) with a low number of polygons in order to improve the computer performance. Afterward we started the modeling process following the muscle insertion areas clearly evident on the skulls (Antón et al., 2009). The volume of the muscle mass was reconstructed taking into account the spatial distance between the temporal surface and the zygomatic arch for the temporal muscle and the distance between the external surface of the zygomatic arch and the horizontal branch of the mandible for the masseter muscle. ...
... Current views tend to recognise a single variable species, Homotherium latidens, in the Plio-Pleistocene of Eurasia (Antón et al., 2005(Antón et al., , 2009. Turner (1999) argued that the differences that led Ficcarelli (1979) to retain H. crenatidens and H. latidens as different species, namely the smaller size and slightly different upper canines of the latter, were simply invalid. ...
... Although based on a detailed consideration of Smilodon, their conclusions have clear applicability across the range of machairodontine taxa, and imply not only a cat-like appearance but also a cat-like pattern of eating, with the incisors pulling off soft tissues and the cheek teeth employed in slicing. A later facial reconstruction of Homotherium (Antón et al., 2009) showed a combination of broadly cat-like appearance with a unique head shape resulting from unmistakable homotheriin cranial proportions such as a straight dorsal profile, a projecting incisor battery and a deep mandibular symphysis implying an elongated, tall and square muzzle (fig. 12). ...
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Sabre-toothed felids, the machairodontines, have attracted much attention among palaeontologists for many decades, not only because of their spectacular morphology but also because they are a striking example of convergent evolution that is most probably linked to strong selective pressures. In this paper we provide a summary of the changing interpretations of their functional anatomy and evolution, from early hypotheses proposing a stabbing mode of attack and a pleiotropic control of the complex of machairodont morphologies, to the current views favouring the canine shear-bite model and a mosaic evolution of anatomical features. Los félidos de dientes de sable, los macairodontinos, han ejercido una especial atracción entre los paleontólogos durante muchas décadas, no sólo por su espectacular morfología, sino también debido a que son un llamativo ejemplo de evolución convergente, probablemente ligada a una fuerte presión selectiva. En este trabajo suministramos una recopilación de los cambios de interpretación acerca de su anatomía functional y evolución, desde las primeras hipótesis en las que se proponía un ataque por apuñalamiento y un control pleitrópico del complejo morfológico macairodontino, hasta los actuales puntos de vista que favorecen un modelo de mordedura muy especializado y una evolución en mosaico de los carácteres anatómicos.
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Homotherium is one of the sabre-toothed felid genera with a more extensive overlap in space and time with species of our own genus Homo, who must have been familiar with the animal, but now we only have its fossil remains to infer its life appearance. A revised reconstruction of the soft tissue and life appearance of Homotherium latidens is proposed here on the basis of new observations on the anatomy of extant carnivorans and a re-evaluation of the fairly preserved skull and mandible from the classical Late Pliocene site of Perrier (France). This fossil specimen provides some of the best information available about the morphology of the skull and mandible. Like other large early specimens of Homotherium, it has enormous upper canines relative to skull size and high-crowned enough to protrude beyond the lips in the living animal. On the other hand, observations of facial expressions in living big cats and dissected specimens show that, contrary to previous conclusions, the soft tissue around the mouth and the lower lip in particular can cover the upper canines of large felids, even when those are considerably high-crowned. Such observations lead us to propose a revised hypothesis about the life appearance of Homotherium and other sabre-toothed carnivorans, where the upper canines may have been covered in life when the mouth was completely closed.
Thesis
This Thesis deals with the palaeoecology of the fossil carnivorans of Greece and, in particular, with their diet. Carnivora don’t eat exclusively meat, but they frequently are omnivores, insectivores or even herbivores. In this study extant Carnivora were divided into 12 dietary categories. The studied fossil material belongs to 47 species from 8 families, coming from two periods of geological time: the Late Miocene (11.6-5.3 Mya) and the Villafranchian (3.5.-0.8 Mya). These periods include some of the richest fossiliferous localities in Greece, providing enough material to apply the necessary methods. To test the made assumptions, a comparative sample of 75 species belonging to 13 families of extant carnivorans was used. The main focus of this work is to calculate a number of proxies which are connected to the diet of these species. 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The third case of coexistence was the felids of Pikermi (Pristifelis attica, Metailurus parvulus, Metailurus major, Paramachairodus orientalis and Amphimachairodus giganteus). These species were able to coexist, because they did not have the same body size. Therefore, their prey also had a relevant body size. The only species of the same size were Paramachairodus and Metailurus major, which represent two different evolutionary stages of sabertooths. Thus, these two taxa must have been competitive with each other and maybe that’s the reason for their infrequency. Another part was the small-sized mustelids of Turolian (Martes woodwardi, Promeles palaeattica, Promephitis lartetii, Parataxidea maraghana, Sinictis pentelici and a new species of mustelid). These species seemed to cover similar niches and having similar body sizes, with the exception of the smaller Promephitis. 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The first lineage was that of Crocuta-like bone-crushing hyenas: Dinocrocuta - Adcrocuta - Pliohyaena - Pachycrocuta - Crocuta and the second was that of large felids Amphimachairodus / Paramachairodus / Metailurus major - Homotherium / Megantereon / Panthera gombaszoegensis - Panthera leo / pardus. Finally, the dietary category for every species was defined in a table and a suggestion for its possible prey genera (based on its already known associated faunas) was made, accompanied with the reference of a modern analogue species.
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In Canis and Ursus the largest proportion of attachments of muscles of the shoulder and brachium on the scapula and humerus is direct; fewer attachments are aponeurotic or tendinous. In both genera most attachments can be associated with superficial osteological features (scars or delimitable surfaces); attachments that lack such features are direct. Most aponeurotic attachments are associated with rugose scarring whereas tendinous attachments are often associated with smooth surfaces. Although most attachments can be associated with osteological features the areal extent of attachment is often not inferrable from the bone. The inference of muscle size or functional significance from osteological features is problematic. The amount of myological information that can be deciphered from the osteology in Canis and Ursus is greater than that reported for particular members of other vertebrate groups which suggests that there may be differences in the degree to which muscles can be reconstructed from superficial osteology alone. Nonetheless, even in mammals such as the Carnivora, detailed muscular reconstructions in extinct taxa cannot be achieved without reference to the musculature of extant relatives. Such reconstructions rely on assumptions, that often have not been adequately tested, regarding the similarity of musculature in closely related taxa. This testing and well corroborated hypotheses of phylogenetic relationship are essential for the evaluation of the accuracy of reconstructions of the musculature in fossil vertebrates.
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The Late Miocene (Late Vallesian, MN 10, about 9 Mya) carnivore trap of Batallones-1 (Madrid, Spain) has yielded a large sample of two species of sabre-toothed cats: the puma-sized Paramachairodus ogygia and the tiger-sized Machairodus aphanistus. This has allowed, for the first time, complete studies of the biomechanics and comparative anatomy of these animals. Focusing our study on the small species, Par. ogygia, the most richly represented and best known carnivore from Batallones-1, we attempt to infer some aspects of the behaviour and ecology of this early sabre-toothed cat, such as breeding behaviour, the degree of social interaction between individuals, sexual dimorphism, preferred habitat and prey size. Our results suggest that Par. ogygia was a solitary felid with a low sexual dimorphism index, which in turn indicates low competition between males for access to females, and some degree of tolerance between adults, so that young adults were allowed to share the territory of their mothers for some time after maturity. The machairodont adaptations of Par. ogygia indicate that this species was able to subdue and kill prey in less time than pantherines do, thus minimizing the risk of injury and the energetic costs of this action. In a wider context, the carnivore guild of Batallones-1 and the overall mammal community indicate that the landscape around the trap was a wooded habitat. Batallones-1 is thus establishing itself as one of the most important European Late Miocene fossil localities, not only for the study of the anatomy and biomechanics of the early sabre-toothed cats but also for our understanding of the intra- and inter-specific ecological relationships of the first members of this specialized sub-family of felids.
Article
The skull and cervical anatomy of the sabre-toothed felid Paramachairodus ogygia (Kaup, 1832) is described in this paper, with special attention paid to its functional morphology. Because of the scarcity of fossil remains, the anatomy of this felid has been very poorly known. However, the recently discovered Miocene carnivore trap of Batallones-1, near Madrid, Spain, has yielded almost complete skeletons of this animal, which is now one of the best known machairodontines. Consequently, the machairodont adaptations of this primitive sabre-toothed felid can be assessed for the first time. Some characters, such as the morphology of the mastoid area, reveal an intermediate state between that of felines and machairodontines, while others, such as the flattened upper canines and verticalized mandibular symphysis, show clear machairodont affinities. © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society, 2005, 144, 363−377.
Article
Recent research at the Oxford Radiocarbon Accelerator Unit (ORAU) has shown that ultrafiltration of gelatin from archaeological bone can, in many instances, remove low molecular weight contaminants. These can sometimes be of a different radiocarbon age and, unless removed, may severely influence results, particularly when dating bones greater than two to three half-lives of 14C. In this study this methodology is applied to samples of Late Middle and Early Upper Palaeolithic age from the British Isles. In many instances the results of redating invite serious reconsideration of the chronology for these periods. Copyright © 2006 John Wiley & Sons, Ltd.
Article
Life reconstructions are a useful means of providing a package of information about morphology, functionality, behaviour, biology and ecological characteristics of an extinct organism. These reconstructions are of interest not only to researchers but also to a wider public. Reconstructions of sabretoothed cats in particular are widely published and exhibited, despite the absence of a general consensus on either how their canines were used or the prey sought. Cat-like restorations of the American Pleistocene sabretooth genus Smilodon prepared by Charles Knight under the direction of J.C. Merriam were accepted as valid for over three decades until GJ. Miller criticized them, claiming that Smilodon should have looked very different from modern felids. In particular, he argued for a longer mouth opening and lip line to provide a wider gape, a retracted nose and ears set relatively lower on a head with a straighter dorsal profile. These arguments were accepted by many authors employing reconstructions, and have lead to depictions of bizarre appearance and interpretations of rather specialized feeding behaviour. We believe that phylogenetic, anatomical and functional considerations point to substantial flaws in the basis for such depictions, and argue for a return to more felid-like morphology and to interpretations of broadly cat-like eating patterns.
Article
Muscle attachments in the mastoid region of the skull of extant felids are studied through dissection of two adult tigers Panthera tigris (Linnaeus, 1758) Pocock, 1930, a lion Panthera leo (Linnaeus, 1758) Pocock, 1930 and a puma Puma concolor (Linnaeus, 1771) Jardine, 1834, providing for the first time an adequate reference for the study of the evolution of that region in sabretoothed felids. Our study supports the inference by W. Akersten that the main muscles inserting in the mastoid process in sabretooths were those originating in the atlas, rather than those from the posterior neck, sternum and forelimb. Those inferences were based on the anatomy of the giant panda, Ailuropoda melanoleuca (David, 1869) Milne-Edwards, 1870, raising uncertainties about homology, which were founded, as revealed by our results. The mastoid muscle insertions in extant felids differ in important details from those described for Ailuropoda, but agree with those described for domestic cats, hyenas and dogs. The large, antero-ventrally projected mastoid process of pantherines allows a moderate implication of the m. obliquus capitis anterior in head-flexion. This contradicts the widespread notion that the function of this muscle in carnivores is to extend the atlanto-cranial joint and to flex it laterally, but supports previous inferences about the head-flexing function of atlanto-mastoid muscles in machairodontines. Sabretooth mastoid morphology implies larger and longer-fibred atlanto-mastoid muscles than in pantherines, and that most of their fibres ran inferior to the axis of rotation of the atlanto-occipital joint, emphasizing head-flexing action. © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 140, 207–221.
Article
Human occupants of Europe shared food resources with a number of larger Carnivora, and their coexistence with two lion-sized felids, the lion and the scimitar-toothed machairodont Homotherium latidens, poses intriguing paleoecological problems. We investigate the ecology of Homotherium latidens using an exceptional sample of postcrania from the Spanish Early Pleistocene site of Incarcal, making comparisons with modern cats and with other machairodont species. Evidence of cursorial adaptations in Homotherium suggests a hunting technique different from modern cats or smilodontine sabre-tooths. Some, like reduction of the claws, would have limited the ability of individual homotheres to bring down large prey, implying group action. Homotherium would also have been disadvantaged in direct confrontation with Pleistocene lions by smaller body mass, reduced forepaw muscle strength, smaller claws and more fragile dentition. Its hunting technique would have worked best in more open habitats, but competition from lions would have forced it to seek moderate cover. Among factors that could de-stabilise coexistence of the two big cat species in Pleistocene Europe we invoke a decrease in environmental mosaicism associated with stepped climatic change over the last million years, and the increased importance of humans within the larger predator guild.
Article
Kent's Cavern has long been known as potentially among the oldest Palaeolithic sites in the country, with the basal Breccia deposit containing a sparse Lower Palaeolithic industry. The sediment consists of a chaotic clayey conglomerate emplaced as a series of debris flows, which entered the cave via blocked entrances at its southwest end. The Breccia contains a fauna dominated by the bear Ursus deningeri, with lion Felis leo and the voles Arvicola cantiana and Microtus oeconomus, establishing a late Cromerian age for the deposit. The artefacts comprise an industry of crudely manufactured handaxes and flakes, and show damage suggesting that they were brought into the cave by the debris flows, and may thus predate the sediment and fauna. We demonstrate an age of >340 ka for the Breccia using two independant dating methods, consistent with existing models of the age of the British Middle Pleistocene sequence.
Article
This study was carried out to assess the breast cancer knowledge, attitudes, and awareness of women age 40 to 74 in Alberta, a Canadian province of 2.4 million people. This analysis compares the attributes of 538 rural women, defined as those living between 1 and 3 hr drive from the major cities in Alberta, and 735 urban women who lived in one of these two cities. Rural women were found to have the same basic knowledge of breast cancer or perceptions of barriers to mammography, but had more negative attitudes about breast cancer itself. Despite their similar access to physician care, they were less likely to have had a recent clinical breast examination or mammogram (P less than 0.001). These differences remained when adjustment was made for demographic background variables; the adjusted prevalence rate ratio for a screening mammogram in the past 2 years was 0.52 (95% C.I., 0.43, 0.64), and for intention to have a mammogram in the next 2 years, 0.75 (0.63, 0.90). The results suggest particular program delivery strategies when planning for provision of breast screening information and service to the large subgroup of rural women.
Pleistocene Mammals of Europe On a supposed prehistoric representation of the Pleistocene scimitar cat, Homotherium Fabrini, 1890 (Mammalia; Machairodontidae)
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Comments on the reconstructions of fossil vertebrates from Lothagam Lothagam: Dawn of Huma-nity in East Africa
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Antón, M., 2003. Comments on the reconstructions of fossil vertebrates from Lothagam. In: Harris, J.M., Leakey, M. (Eds.), Lothagam: Dawn of Huma-nity in East Africa. Columbia University Press, New York, pp. 661–665.
La Caverne de Font-de-Gaume aux Eyzies (Dordogne) La Grotte Chauvet : l'art des Origines Observations on the artefacts from the Breccia at Kent's Cavern
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Les grands félins dans l'art de notre préhistoire
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Witmer, L.M., 1995. The extant phylogenetic bracket and the importance of reconstructing soft tissues in fossils. In: Thomason, J.J. (Ed.), Functional Morphology in Vertebrate Paleontology. University of Cambridge Press, Cambridge, pp. 19–33.
The Saber-toothed Cat of the North Sea Age of Middle Pleistocene fauna and Lower Palaeolithic industries from Kent's Cavern, Devon
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Mol, D., van Logchem, W., van Hooijdonk, K, Bakker, R., 2008. The Saber-toothed Cat of the North Sea. DrukWare, Norg. Proctor, C.J., Berridge, P.J., Bishop, M.J., Richards, D.A., Smart, P.L., 2005. Age of Middle Pleistocene fauna and Lower Palaeolithic industries from Kent's Cavern, Devon. Quaternary Science Reviews 24, 1243–1252.
The Saber-toothed Cat of the North Sea
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Mol, D., van Logchem, W., van Hooijdonk, K, Bakker, R., 2008. The Sabertoothed Cat of the North Sea. DrukWare, Norg.
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La Grotte Chauvet : l'art des Origines
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Clottes, J., 2001. La Grotte Chauvet : l'art des Origines. Éditions du Seuil, Paris.
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Los carnívoros de los yacimientos Pleistocenos de la Sierra de Atapuerca. Ph.D. Disertación, Departamento de Biología Animal I, Facultad de Ciencias Biológicas
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Comments on the reconstructions of fossil vertebrates from Lothagam
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