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Other tarsals from Moula-Guercy. (A) M-E1-222 from left to right, inferior view (distal is up), proximal view (dorsal is left), medial view (distal is left), lateral view (distal is left). For B–D views are arranged as follows: from left to right, distal view, proximal view, medial view, lateral view; for distal and proximal views, dorsal is left; for medial and lateral views, distal is left. (B) M-G2-464; (C) M-S-18; (D) M-I2-8. Scale is 2 cm. Natural size.
Source publication
The hand and foot remains from Moula-Guercy cave (Ardèche, France) comprise 24 specimens of Eemian age (ca. 120 ka). The specimens include primarily complete elements, which are rare among the Moula-Guercy postcrania. The hand remains have several characteristic Neanderthal traits including a laterally facing (parasagittally oriented) second metaca...
Contexts in source publication
Context 1
... is a nearly complete mature right cuboid with some lateral, medial, and dorsal erosion (Fig. 6A). Though there is some plantaromedial erosion, the pres- ence of a navicular facet is ...
Context 2
... and M-S-18 are mature complete right and left naviculars with some minor erosion of cortical bone (Fig. 6B,C). Mature status is based on overall size. Met- ric data are presented in Table ...
Context 3
... is a complete mature left medial cuneiform with minor erosion, particularly on the medial surface (Fig. 6D). Mature status is based on overall size (see Supporting Information Table S4 for metric data). The medial plantar tubercle is expanded, extending 4.5 mm medial to the navicular facet. The articulation for the intermediate cuneiform is a single facet extending in an arc proximodistally from the dorsolateral corner of the navicular facet ...
Similar publications
Here, we provide a complete, updated, and illustrated inventory, as well as a comprehensive study, of the tarsals (rearfoot) recovered from the Middle Pleistocene site of Sima de los Huesos (SH, Atapuerca, Spain) in comparison to other Homo comparative samples, both extant and fossil. The minimum number of individuals (MNI) estimated from the tarsa...
Citations
... Thus, the phalanx I2-104 fits in well with the Neandertal health pattern already known, although the proximal epiphysis in I2-104 shows no alteration. Other hand phalanges and skeletal remains from Moula-Guercy do not show osteoarthritis or any other pathological changes (Mersey et al., 2013a(Mersey et al., , 2013bRichards et al., 2021). Nonetheless, it should be noted that only a small part of level XV has been excavated, and it is highly possible that other remains are still in situ. ...
Objective: To discuss a Neandertal pathological adult first pollical proximal phalanx (I2-104) from the Baume de Moula-Guercy (Ardèche, France) and evaluate the possible causes of this pathology.
Methods: Macroscopic analyses of external features, as well as CT imaging, were used in the analysis.
Results: The presence of asymmetric eburnation on the distal epiphysis associated with an osteophyte on the palmar surface, as well as the absence of periosteal bone reaction visible on CT images, is consistent with osteoarthritis.
Conclusion: Osteoarthritis (OA) can have different origins and the cause is difficult to identify. The pathology of the Moula-Guercy I2-104 phalanx may be due to a genetic predisposition for OA known in Neandertals and associated with short limb bones. The OA could have been aggravated by the age of this individual and by an inflammatory reaction caused by repeated movements and intense vibrations provoked by high-frequency knapping or by other use of the hands.
Significance: The I2-104 phalanx is the first Neandertal pollical phalanx known to display OA, although joints of this bone are frequently affected by this pathology in modern humans. Thus, greater insight into the presence and consequences of Neandertal behaviors is offered.
Limitation: It is impossible to give a definitive conclusion on the cause(s) of the OA in this case.
Suggestions for further research: More data is needed concerning OA within Neandertals and its relationship with behavior and genetics.
... As detailed by Richards et al. (2021), the Guercy 1 late adolescent female cranium possesses a total morphological pattern (Le Gros Clark, 1964a) that is consistent with the morphological features defined for the Early Neanderthal subgroup (Bräuer et al., 2020;Couture & Hublin, 2005;Dean et al., 1998;Guipert et al., 2011;Sergi, 1944;Vandermeersch, 1978). The dental, axial, and appendicular remains have only been assessed relative to Neanderthals in general (Hlusko et al., 2013;Mersey, Jabbour, et al., 2013). ...
... Dutour, Defleur et al., 1999;Hlusko et al., 2013;Mersey, Jabbour, et al., 2013;Defleur et al., 2020, Richards et al., 2021. Expanding the geographic range to Southwestern and South-Central France adds significantly to the Early Neanderthal craniodental sample by the inclusion of Biache-Saint-Vaast (N = 2), La Chaise Abri Suard (N = 52), and La Chaise Bourgeois-Delaunay (N = 23: Piveteau et al., 1982;Vandermeersch, 1982;Condemi, 2001;Rougier, 2003;Guipert et al., 2011). ...
Objectives
This article provides an ontogenetically based comparative description of two immature occipital fragments from Baume Moula‐Guercy (MIS 5e) and examines their affinities to European and Middle Eastern Middle‐to‐Late Pleistocene (≈MIS 14–MIS ≈ 1) Homo .
Materials and methods
Description of the M‐S‐41 and M‐S‐61 occipital fragments (≈6–8 years) is based on observations of original fossils, casts, CT scans, literature descriptions, and virtual ectocranial and endocranial reconstructions. Our ontogenetically based sample represents a Preneanderthal‐Neanderthal group and a Homo sapiens group. These groups are subdivided into (1) Preneanderthals (≈MIS 14–9), Early Neanderthals (MIS 7–5e), and Late Neanderthals (MIS 5d‐3), and (2) Middle (MIS 5), Upper (MIS 3–1), and Late (MIS ≈ 1) Paleolithic H. sapiens . Measurements and developmental age determinations follow standard techniques.
Results
Based on the M‐S‐41/M‐S‐61 composite, the strongly convex upper occipital scale flattens into the vertical suprainiac fossa, as in immature Early Neanderthals. A doubled suprainiac fossa expressing a weakly developed bounding torus and pocking of the central region is typical of immature Neanderthals. The transverse torus is thickened medially, lacks significant lateral development, has a laterally placed protuberance, and is concave medially. A linear (triangular) tubercle, as opposed to an external occipital protuberance, is present. An occipital bun is absent.
Discussion
The occipital remains from Moula‐Guercy make a substantial contribution to the record of Neanderthal cranial evolution because immature Early Neanderthal occipitals are rare, fragmentary, and likely represent only two paleodemes (Krapina, La Chaise‐de‐Vouthon). The Moula‐Guercy occipital possesses characteristics well‐established in European Neanderthals by MIS 7, and it is most similar to the occipitals of other Early Neanderthals.
... (2) contributes new data related to the evolution of Preneanderthals and Neanderthals in Europe. Descriptions of the dentition (Hlusko et al., 2013), axial and appendicular skeleton , and hands and feet (Mersey, Jabbour, et al., 2013) have been published. To date, however, descriptions of the cranial remains from Baume Moula-Guercy have been preliminary assessments (Defleur, 1995;Defleur et al., 1999;Defleur, Dutour, Valladas, Combier, & Vandermeersch, 1993). ...
... In order to discern individuals and individual ages and sexes within the Moula-Guercy craniodental sample, we provide assessment of the minimum number of individuals (MNI). For completeness, we also provide a correlation between these data and those from the dental (Hlusko et al., 2013) and postcranial skeleton Mersey, Jabbour, et al., 2013). Because of the importance of these remains, comparative descriptions of other portions of the Moula-Guercy skull sample are being prepared separately. ...
Objectives:
We provide the first comparative description of the Guercy 1 cranium and isolated cranial fragments from Baume Moula-Guercy and examine their affinities to European Preneanderthals, Neanderthals, and Homo sapiens.
Materials and methods:
The Moula-Guercy hominins derive from deposits chronostratigraphically and biostratigraphically dated to the Eemian Interglacial (MIS 5e). For comparisons we compiled a sample of European and Southwest Asian subadult-adult Middle-to-Late Pleistocene hominins (≈MIS 14-MIS 2; N = 184). This sample represents a Preneanderthal-Neanderthal group and a H. sapiens group, both of which were further divided into three time-successive subgroups defined by associated marine isotope stages (MIS). Metric and morphological observations were made on the original fossils and a virtual reconstruction of Guercy 1. Developmental age and sex and the minimum-maximum number of individuals were assessed.
Results:
Guercy 1 represents the remains of a late stage adolescent (≈15-16.0 years) female. Morphological and metric data combine to associate the total morphological pattern expressed in Guercy 1 with our MIS 7-MIS 5e ("Early Neanderthal") subgroup. Some features, especially those related to the frontal, suggest linkage to a paleodeme comprising the Moula-Guercy, Artenac, La Chaise Abri Suard and, possibly, the Biache-Saint-Vaast samples.
Discussion:
Remains of MIS 7-MIS 5e Neanderthals are rare and fragmentary, especially those dated to the Last Interglacial. The Baume Moula-Guercy sample provides new insights into the total morphological pattern expressed in MIS 5e Neanderthals. Further, our results support earlier suggestions that MIS 7-MIS 5e European hominins represent a morphotype that is distinct from both earlier and later members of the Preneanderthal-Neanderthal group.
... Processus médial plantaire moins volumineux Trinkaus, 1983 Tubercule médial en sailli Tubercule médial plus arrondi Pablos et al ., 2019a Tubérosité large proximodistalement Tubérosité moins large Trinkaus, 1983;Mersey et al ., 2013;Pomeroy et al ., 2017 Portion latérale peu épaisse Portion latérale plus épaisse Larges Graciles Trinkaus, 1975 ;1983;Trinkaus et Hilton, 1996 ;Mersey et al ., 2013 ; Premier métatarsien ayant une large insertion pour le tendon musculaire du peroneus longus Premier métatarsien ayant une insertion plus réduite Trinkaus, 1983 Bases des métatarsiens latéraux (3 à 5) larges ...
... Processus médial plantaire moins volumineux Trinkaus, 1983 Tubercule médial en sailli Tubercule médial plus arrondi Pablos et al ., 2019a Tubérosité large proximodistalement Tubérosité moins large Trinkaus, 1983;Mersey et al ., 2013;Pomeroy et al ., 2017 Portion latérale peu épaisse Portion latérale plus épaisse Larges Graciles Trinkaus, 1975 ;1983;Trinkaus et Hilton, 1996 ;Mersey et al ., 2013 ; Premier métatarsien ayant une large insertion pour le tendon musculaire du peroneus longus Premier métatarsien ayant une insertion plus réduite Trinkaus, 1983 Bases des métatarsiens latéraux (3 à 5) larges ...
... Bases des métatarsiens latéraux (3 à 5) plus étroites Pablos et al ., 2017;2019a Rapport de longueur entre le cinquième et le troisième métatarsiens important Rapport de longueur entre le cinquième et le troisième métatarsiens plus faible Pablos et al. , 2019a Phalanges larges médiolatéralement Phalanges plus graciles Trinkaus, 1975 ;1983;Trinkaus et Hilton, 1996 ;Pablos et al ., 2019a ;2019b Phalanges proximales ayant de larges bases (suggérant une aponévrose plantaire développée) Phalanges proximales ayant des bases moins larges Trinkaus, 1983 Phalanges proximales courtes de manière absolue et relatives par rapport aux phalanges intermédiaires et distales Phalanges proximales plus longues Trinkaus, 1983 ;Trinkaus et Hilton, 1996;Mersey et al ., 2013 ;Pablos et al. , 2019b Phalanges distales hautes dans le plan dosrso-plantaire Phalanges distales moins hautes Pablos et al. , 2019b Phalanges distales de l'hallux ayant une tubérosité large Annexes IV : Morphométrie des empreintes de pieds expérimentales 374 Figure A4.2 : Variation de la stature en fonction des variables morphométriques des empreintes expérimentales. ...
Les empreintes de pieds d’hominines représentent des vestiges uniques ouvrant une fenêtre sur de brefs moments de vie de groupes disparus. Leur étude donne des informations directes sur la taille et la composition de ces groupes, paramètres essentiels à leur succès adaptatif mais rarement accessibles à partir du registre fossile. Toutefois, l’étude des empreintes est rendue complexe par le nombre de facteurs (caractéristiques corporelles et biomécaniques, nature du substrat, taphonomie) impactant leur morphologie et leur relative rareté au sein du registre fossile. Depuis 2012, plusieurs centaines d’empreintes de pieds potentielles attribuables à des néandertaliens ont été découvertes associées à un riche matériel archéologique dans 5 niveaux datés à 80 000 ans de la paléodune du Rozel (Manche, France). La découverte de ces empreintes offre l’opportunité de s’intéresser à la taille et à la composition des groupes sociaux néandertaliens, problématique centrale de ce doctorat. Dans un premier temps, l’analyse des empreintes potentielles découvertes entre 2012 et 2017 a permis d’identifier 257 empreintes de pieds et 8 empreintes de mains ce qui représente à ce jour le plus gros corpus ichnologique associé aux Néandertaliens. Puis, grâce au développement d’une approche combinant morphométrie et expérimentation, la taille et la composition des groupes a été estimée à partir des empreintes numérisées en 3D. Les empreintes de pieds étudiées reflètent ainsi différentes classes d’âge allant du très jeune enfant (à partir de 1 an) à l’adulte. Les empreintes issues du niveau le plus dense ichnologiquement représentent un groupe de petite taille, probablement composé de 10 à 13 individus, dont 90% étaient des enfants et des adolescents. Les empreintes de pieds du Rozel fournissent ainsi des informations uniques sur la taille et la composition des groupes néandertaliens permettant de mieux comprendre les occupations paléolithiques au Rozel il y a 80 000 ans.
... Anatomical studies on the crania and mandibles, teeth (Hlusko et al., 2013) and postcranial remains (Mersey et al., 2013a(Mersey et al., , 2013b, identified characteristic Neanderthal traits, while no evidence for the presence of any of other hominid taxa was found. Detailed forensic examinations on the human remains showed that all individuals, including a 4-yearold child, had been cannibalised. ...
... In addition, the Manot Cave specimen displays a uniquely small talus and calcaneus when compared with the long metatarsals. The Neanderthal foot bones, in contrast, are characterized by their enhanced robustness, especially evident in the talus and the first and fifth metatarsals (Trinkaus, 1975;Mersey et al., 2013). The AMH foot bones, best represented by fossils from Skhul and Qafzeh (~120-90kyr ago) display an intermediate robusticity between modern humans and Neanderthals (McCown and Keith, 1939;Vandermeersch, 1981). ...
The transition from the Middle Paleolithic to the Upper Paleolithic in the Levant represents a major event in human prehistory with regards to the dispersal of modern human populations. Unfortunately, the scarcity of human remains from this period has hampered our ability to study the anatomy of Upper Paleolithic populations. This study describes and examines pedal bones recovered from the Early Upper Paleolithic period at Manot Cave, Israel, from 2014 to 2017. The Manot Cave foot bones include a partial, left foot skeleton comprising a talus, a calcaneus, a cuboid, a first metatarsal, a second metatarsal, a fifth metatarsal, and a hallucal sesamoid. All these remains were found in the same archaeological unit of the cave and belong to a young adult. Shape and size comparisons with Neanderthals, Anatomically Modern Human and modern human foot bones indicate a modern human morphology. In some characteristics, however, the Manot Cave foot bones display a Neanderthal-like pattern. Notably, the Manot Cave foot is remarkable in its overall gracility. A healed traumatic injury in the second metatarsal (Lisfranc's fracture) is most likely due to a remote impact to the dorsum of the foot. This injury, its subsequent debility, and the individual's apparent recovery suggest that the members of the Manot Cave community had a supportive environment, one with mutual responsibilities among the members.
... -Unlike Krapina and other sites, such as the Sima de los Huesos (Bocquet-Appel and Arsuaga, 1999), at BMG layer XV the individuals were not selected from any age group. Younger people aged under 15-16 years represent 2/3 of cannibalized individuals, along with an elderly robust individual and another gracile youngadult possibly female (Defleur et al., 1999;Hlusko et al., 2013;Mersey et al., 2013aMersey et al., , 2013b. All age classes are close to natural mortality cases observed in recent hunter-gatherers (Gurven and Kaplan, 2007). ...
Earth's climate experienced a major warming during the last interglacial period (Eemian, MIS 5e, LIG 128 to 114 ky). The rapid climate change altered ecosystems causing a geographical redistribution of flora and fauna. Due to the scarcity of archaeological sites representing this period, the effect of these events on the behavior of Neanderthal hunter-gatherers in Western Europe has been poorly understood. New evidence from a well preserved archaeological layer (XV) at Baume (cave) Moula-Guercy in Southeastern France, attributed to the optimum Eemian Interglacial, unparalleled on the European continent, allows us to consider the challenges Neanderthals faced as these new ecosystems and ecological communities formed. We argue that, on the European continent, the human population collapsed, maintaining itself only in a few regions. We further suggest that these environmental upheavals, including depletion of prey biomass at the beginning of the Upper Pleistocene, contributed to the rise of cannibalistic behavior in Neanderthals, as exhibited among remains found at the Baume Moula-Guercy.
... Stabilization of the lateral tarsometatarsal joint is a key bipedal innovation in early hominins and thus the flattening of this joint in Ardipithecus ) supports its identification as a bipedal hominin. Trinkaus, 1975;Latimer et al., 1982;Pearson, Royer, Grine, & Fleagle, 2008;Lu et al., 2011;Mersey, Jabbour, Brudvik, & Defleur, 2013;Harcourt-Smith et al., 2015;Pablos, Pantoja-Pérez, Martínez, Lorenzo, & Arsuaga, 2017 , 2015). Naviculars from Ardipithecus (ARA-VP-6/503: White et al., 2009;Lovejoy et al., 2009), StW 623 (Clarke, 2013) and KNM-ER 64062 (Jungers et al., 2015) have been discussed in print but are not yet formally described. ...
... Begun (2004) suggested that dorsal canting also occurs in the phalanges of nonbipedal hominoids, including Sivapithecus, and cited Rose (1986 Trinkaus, 1975;Latimer et al., 1982;Lorenzo, Arsuaga, & Carretero, 1999;Pearson et al., 2008 Note. Data in this table from original fossils and measurements reported in Day & Leakey, 1974;Latimer et al., 1982;Ward et al., 1999;Semaw, Simpson, Quade, & Renne, 2005;Mednikova, 2011;Haile-Selassie et al., 2012;Mersey et al., 2013;Vernon, 2013;Harcourt-Smith et al., 2015;Trinkaus and Patel, 2017. PPs from Ardipithecus (ARA-VP-1/2677, 6/1006, 6/1005: White et al., 2009), and from Gona (Simpson et al., 2018), and ASI-VP-2/215 (White et al., 2006) have been discussed in print but are not yet described in detail or are too fragmentary to yield the measurements listed above. ...
... However, as in later hominins, the midfoot (cuboid) is proximodistally long. Image courtesy of Tim White and Gen Suwa Day & Leakey, 1974;Trinkaus, 1975;Latimer et al., 1982;Day, 1986;Lorenzo et al., 1999;Pearson et al., 2008;Lovejoy et al., 2009;Mersey et al., 2013;Harcourt-Smith et al., 2015. DiPs from Ardipithecus (ARA-VP-6/500-033, -052, -099, 6/1008, 6/1009: White et al., 2009), Dmanisi (D2670 and D3877; Lordkipanidze et al., 2006;Patel et al., 2015), Jinniushan (Lu et al., 2011) have been discussed in print but are not yet described in detail or are too fragmentary to yield the measurements listed above. ...
Bipedalism is a hallmark of being human and the human foot is modified to reflect this unique form of locomotion. Leonardo da Vinci is credited with calling the human foot “a masterpiece of engineering and a work of art.” However, a scientific approach to human origins has revealed that our feet are products of a long, evolutionary history in which a mobile, grasping organ has been converted into a propulsive structure adapted for the rigors of bipedal locomotion. Reconstructing the evolutionary history of foot anatomy benefits from a fossil record; yet, prior to 1960, the only hominin foot bones recovered were from Neandertals. Even into the 1990s, the human foot fossil record consisted mostly of fragmentary remains. However, in the last two decades, the human foot fossil record has quadrupled, and these new discoveries have fostered fresh new perspectives on how our feet evolved. In this review, we document anatomical differences between extant ape and human foot bones, and comprehensively examine the hominin foot fossil record. Additionally, we take a novel approach and conduct a cladistics analysis on foot fossils (n = 19 taxa; n = 80 characters), and find strong evidence for mosaic evolution of the foot, and a variety of anatomically and functionally distinct foot forms as bipedal locomotion evolved.
... Nevertheless, among those deserving special mention is Baume Moula-Guercy, 80 m above the River Rhône, where twenty sedimentary layers span a period that began in MIS-6 and ended in MIS-4. The deep layer XV assigned to MIS-5 contained a "Ferrassie Mousterian" assemblage with "Levallois" core-reduction, and bones showing cut-marks and signs of defleshing (maybe for extracting marrow and brain) of both red deer and six Neanderthals, perhaps implying cannibalism (Defleur et al., , 1998Hlusko et al., 2013;Mersey et al., 2013). ...
This paper provides a background to Neanderthal presence in Western Mediterranean Europe. Habitual tool-use underpinned human survival in late Early Pleistocene western Mediterranean Europe. By the onset of the early Middle Pleistocene, early humans (descendants of Homo antecessor in all likelihood) were exploiting diverse biotopes, sometimes (perhaps often) attaining primary access to large game, and deploying a variety of stone artifacts and rock types, which implies not only manual dexterity but also technical competence and cognitive versatility. Late Early Pleistocene human behaviour foreshadowed that of Middle Pleistocene pre-Neanderthal humans whose background conceivably had deep regional roots. By the mid-Middle Pleistocene Homo heidelbergensis, some of whose anatomical features herald Neanderthal morphology, was exploiting a wide range of natural resources in western Mediterranean environments, including small game and plants. Neanderthal morphology began to emerge during the second half of the Middle Pleistocene, accompanied by increasing technological diversity and an expanding variety of small tools, conjecturally favoured by hafting, perhaps following development of wooden spears (or other tools) and adhesive and binding technologies, and generation and heat-control of fire (which undoubtedly was necessary for activities inside Bruniquel cave). By the onset of the last interglacial period, Neanderthal morphology and Mousterian artifacts are widespread, and there are indications of coordinated retrieval and treatment of body-parts of large ungulates.
... Understanding the relationships between morphology, dexterity, and thermoregulation are also important for interpreting the morphology of skeletal remains from modern humans and extinct hominins. For example, the comparative length of metacarpals between Homo neanderthalensis (Neanderthals) and modern humans indicates that Neanderthals had longer palms relative to their digits (Mersey, Jabbour, Brudvik, & Defleur, 2013;Musgrave, 1971). This would indicate that their hands may have remained warmer in cold conditions, based on the experimental results of Payne et al. (in press), although it is not possible to make inferences about how vasoregulation may have affected Neanderthal extremity temperature. ...
Objectives
The current study investigated whether size and proportions of the hands and digits affect dexterity during severe cold exposure. As wide hands are known to lose less heat than narrow hands, and narrow digits are associated with greater dexterity, this study aimed to test whether there was a direct trade‐off between dexterity and thermoregulation that shapes hand morphology.
Methods
Participants (25 women, 15 men) carried out the Purdue Pegboard test before and after a 3‐min ice‐water immersion of the hand. Their hand length, hand width, digit lengths, and digit widths were measured using standard anthropometric methods.
Results
Wide first and third digits associated with significantly reduced dexterity after immersion relative to individuals with narrower first and third digits. Second digit width positively correlated with average digit temperature after immersion. Hand length and hand width did not influence dexterity.
Conclusion
The current study suggests that digit width influences dexterity in cold conditions, reflecting patterns found at room temperature. Hand and digit morphology may be the product of two significant constraints on the hand: dexterity and thermoregulation. In cold conditions, hand morphology appears to be predominantly constrained by thermal stress, at the expense of dexterity. This may have important implications for interpreting the morphology of extinct and extant hominins.
