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Florent Rivals

Institut Català de Paleoecologia Humana i Evolució Social (IPHES)
Avinguda Catalunya 35
43002 Tarragona, Spain
florent.rivals-at-gmail.com

PUBLICATIONS LIST


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Journal articles

2012
Jordi Rosell, Isabel Cáceres, Ruth Blasco, María Bennàsar, Pilar Bravo, Gerard Campeny, Montserrat Esteban-Nadal, M Cristina Fernández-Laso, M Joana Gabucio, Rosa Huguet, Nuria Ibáñez, Patricia Martín, Florent Rivals, Antonio Rodríguez-Hidalgo, Palmira Saladié (2012)  A zooarchaeological contribution to establish occupational patterns at Level J of Abric Romaní (Barcelona, Spain).   Quaternary International 247: 69-84  
Abstract: In recent decades, many research questions focus on occupational patterns in the Middle Palaeolithic. Many discussions on that topic address the temporal resolution of formation of assemblages and the character of palimpsests that can be frequently observed at archaeological sites. The Abric Romaní site (Barcelona, Spain) is a good example to analyze questions related to temporal resolution of the human occupations. This site is filled by a succession of travertine platforms of rapid formation, which clearly separate different archaeological levels. With the objective of characterizing the settlement patterns developed at this site, a combination of zooarchaeological and spatial data from Level J (sublevels Ja and Jb) is presented here. On one hand, primary and immediate anthropogenic accesses to the carcasses (mainly horses and red deer) and well-established patterns of animal processing are observed. On the other hand, these activities are always developed around hearths of inhabitable areas. From a nutritional perspective, faunal refits suggest synchronic relationships between the different activity areas and, therefore, the anthropic occupation of the total surface of the rock-shelter during the formation of Level J. However, this certain synchrony is combined with the presence of other elements, such as overlapped hearths, which show a succession of several occupations with the same settlement patterns. In this way, Level J of Abric Romaní could be understood as the succession of several occupations formed by groups that maintained the same traditions through time. From this perspective, the aim of this paper is to improve data to understand the settlement patterns and the social organization systems of the Neanderthal groups during MIS 3.
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Francesc Burjachs, Juan Manuel López-García, Ethel Allué, Hugues-Alexandre Blain, Florent Rivals, Maria Bennàsar, Isabel Expósito (2012)  Palaeoecology of Neanderthals during Dansgaard-Oeschger cycles in northeastern Iberia (Abric Romaní): from regional to global scale.   Quaternary International 247: 26-37  
Abstract: The sequence from Abric Romaní has provided several palaeoecological studies. This paper presents a synthesis of the results yielded by palynology, including the data from the entire non-excavated deposits, and the data obtained through the study of large mammals, micromammals, herpetofauna, and anthracology from the excavated layers. The palaeobotanical studies show that the sequence is mainly related to pine forests, and near or within these dominant forests there were smaller plant formations and/or secondary species with a Mediterranean and/or Atlantic character. Nevertheless, continuous forest did not dominate the entire region. The fauna occupying the territory indicate open spaces in which seasonal grasslands proliferated, dry assemblages in sunny areas and humid ones in mountain shadows. The non-excavated archaeological layers correspond to MIS 4, initially characterized by temperate and humid conditions, and thereafter cool and humid until HS 6 and MIS 3. Afterwards, at the beginning of the MIS 3, the DansgaardâOeschger cycles, â¼1500 yr rapid and abrupt climatic changes, occurred during the deposition of the archaeological levels O, N, M, L, K and J. Later, these events were interrupted by a cold and dry phase, between ca 49 and 47 ka BP, which affected levels Ja, I and H, culminating with HS 5. Finally, this cold episode was followed by the Hengelo interstadial or DO 12, characterized by a temperate and humid climate identified in levels E and D.
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Florent Rivals (2012)  Ungulate feeding ecology and middle Pleistocene paleoenvironments at Hundsheim and Deutsch-Altenburg 1 (eastern Austria).   Palaeogeography, Palaeoclimatology, Palaeoecology 217-218: 27-31  
Abstract: The purpose of this study is to analyze the dietary traits in the ungulate communities from two middle Pleistocene localities in eastern Austria, Deutsch-Altenburg 1 and Hundsheim, and to revise the paleoecological reconstructions at each site using tooth wear analyses (mesowear and microwear). The data will also be used to test for intra- and inter-site differences and analyze resource partitioning among ungulates. Intra-site comparisons show significant differences at the two localities and reveal the use of different resources for the various ungulates studied. This suggests the existence of dietary partitioning and niche segregation that would not be detected using traditional morphological proxies. The local environment at the two sites is characterized by the availability of both open and forested habitats. This study indicates the presence of mixed environmental conditions which is a different conclusion from previous paleoenvironmental reconstructions.
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Florent Rivals, Gina M Semprebon (2012)  Paleoindian subsistence strategies and late Pleistocene paleoenvironments in the northeastern and southwestern United States : a tooth wear analysis.   Journal of Archaeological Science 39: 5. 1608-1617  
Abstract: Paleoindians of North America entered a continent undergoing rapid climatic and environmental changes. This paper is a preliminary contribution toward obtaining a better picture regarding how climate and environmental change might have impacted the first settlers of North America. The Paleoindian sites we analyzed are, from the oldest to the youngest, Ingleside (Texas), Blackwater Draw Locality No. 1 (New Mexico), Hiscock Site (New York), and Plainview Quarry (Texas). Paleoenvironmental reconstruction involves identifying the dietary traits of ungulate species that might reflect the environmental conditions where they were living, and also where they might have been hunted by Paleoindians. Such an approach is realized through tooth microwear and mesowear analyses. Results indicate that a variety of food resources were available for the ungulates at these sites as well as the likely presence of mosaic environments around these sites which allowed Paleoindians to exploit resources in a large variety of habitats. The application of a method that allows for the estimation of the duration of occupation at archaeological sites reveals that results for Blackwater Draw and Hiscock Site indicate a long-term occupation of probably several months. However, during short events, Paleoindians were most likely hunting herds of horse and bison when these prey were available near the site. Results indicate that Plainview Quarry was likely used only for short-term occupations, with large game hunting focusing on bison. These patterns identified at the archeological sites studied are related to the fact that Paleoindians follow a high-technology forager model and frequently shifted their territory depending on the composition and distribution of the large mammal fauna.
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Juan Manuel López-García, Hugues-Alexandre Blain, Francesc Burjachs, Anna Ballesteros, Ethel Allué, Gloria Ericka Cuevas-Ruiz, Florent Rivals, Ruth Blasco, Juan Ignacio Morales, Antonio Rodríguez Hidalgo, Eudald Carbonell, David Serrat, Jordi Rosell (2012)  A multidisciplinary approach to reconstructing the chronology and environment of southwestern European Neanderthals: the contribution of Teixoneres cave (Moià, Barcelona, Spain).   Quaternary Science Reviews 43: 33-44  
Abstract: According to pollen analysis and the 18O-isotope curve, the first part of the Late Pleistocene (ca 128 +/- 30 ka) is mainly characterized by a dynamic that alternates cold phases (Heinrich Events) and temperate phases (interstadials). These rapid fluctuations provide the context for the Neanderthal occupations in the northeastern part of the Iberian Peninsula. In this paper we present the chronological, environmental and climatic data obtained by analyzing the pollen, the charcoal, the small vertebrates (amphibians, squamates and small mammals) and the large-mammal dental wear at the Neanderthal site of Teixoneres cave, Northeastern Iberia. Levels II and III from this cavity have provided Mousterian industries and other evidence of Neanderthal occupations, such as cut-marks in large-mammal bones. A multiproxy study such as this constitutes a new approach to the chronological, environmental and climatic context in which Neanderthal populations lived in southwestern Europe (Iberian Peninsula). The results allow us to establish a relative chronology for these two levels of between ca 30e90 ka and show that they are associated with different environmental and climatic conditions: temperate and humid for Level III and cold and dry for Level II. This demonstrates that the Neanderthals were well adapted to the territory that they occupied, irrespective of the climatic conditions.
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Florent Rivals, Gina M Semprebon, Adrian Lister (2012)  An examination of dietary diversity patterns in Pleistocene proboscideans (Mammuthus, Palaeoloxodon, and Mammut) from Europe and North America as revealed by dental microwear.   Quaternary International 255: 188-195  
Abstract: Recently, dental wear studies in mammals have emerged as valuable proxies for documenting the existence of geographical and/or temporal variability in diet and vegetation. However, very few paleodietary studies have focused on proboscidean dietary reconstruction. The most sensitive proxy for discerning short-term dietary shifts is dental microwear. Dental microwear analysis was used to study the dietary traits of fossil proboscideans in eleven samples from nine Pleistocene localities from Europe and North America (Palaeoloxodon antiquus, Mammuthus meridionalis, Mammuthus columbi, Mammuthus primigenius, and Mammut americanum). The North American M. americanum was a committed browser in both localities studied. Interestingly, M. americanum and M. columbi from the same locality (Phosphate Beds, South Carolina) were both engaged in browsing. The genus Mammuthus shows a more browsing habit in a sampled population of its early representative (M. meridionalis), with a probable trend through rather variable mixed feeding in late Mammuthus trogontherii, to grazing specialization in some, but possibly not all, M. primigenius. P. antiquus from Europe also demonstrates dietary plasticity. However, P. antiquus apparently shows a trend toward including a larger component of browse in its diet following its entry into Europe. These different dietary trends in the latter two genera, as far as current data allow, might be the result of a trend towards niche differentiation in these taxa.
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2011
Florent Rivals, Armelle Gardeisen, Jean Cantuel (2011)  Domestic and wild ungulate dietary traits at Kouphovouno (Sparta, Greece): implications for livestock management and paleoenvironment in the Neolithic.   Journal of Archaeological Science 38: 528-537  
Abstract: The objective for this study is to explore interspecific variations in domestic and wild ungulate diets and management at the Neolithic site of Kouphovouno (Sparta, southern Greece). We tested four hypotheses related to environmental context and livestock management using, for the first time, a combination of mesowear and microwear analyses on a Neolithic site. We identified interspecific differences between ungulate taxa, especially between wild and domestic ungulates, however there is no significant difference between Ovis and Capra. There is no diachronic change in diet from the Middle Neolithic to the Late Neolithic. Changes in animal utilization are not reflected in their diet. Whatever the purpose of the husbandry, animals had access to the same food resources. The study of dental wear patterns gave some insights regarding the management practices for the domestic livestock as well as paleoenvironment through the study of wild ungulates.
Notes: ⺠Differences in dental wear exist between wild and domestic ungulates. ⺠Differences in dietary traits are not significant between sheep and goats. ⺠Dietary traits of the livestock did not change from the Middle to the Late Neolithic. ⺠Changes in animal management and utilization are not reflected in their diet.
Matthew C Mihlbachler, Florent Rivals, Nikos Solounias, Gina M Semprebon (2011)  Dietary change and evolution of horses in North America.   Science 331: 1178-1181  
Abstract: The evolution of high-crowned molars among horses (Family Equidae) is thought to be an adaptation for abrasive diets associated with the spread of grasslands. The sharpness and relief of the worn cusp apices of teeth (mesowear) are a measure of dietary abrasion. We collected mesowear data for North American Equidae for the past 55.5 million years to test the association of molar height and dietary abrasion. Mesowear trends in horses are reflective of global cooling and associated vegetation changes. There is a strong correlation between mesowear and crown height in horses; however, most horse paleopopulations had highly variable amounts of dietary abrasion, suggesting that selective pressures for crown height may have been weak much of the time. However, instances of higher abrasion were observed in some paleopopulations, suggesting intervals of stronger selection for the evolution of dentitions, including the early Miocene shortly before the first appearance of Equinae, the horse subfamily in which high-crowned dentitions evolved.
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Florent Rivals, Nikos Solounias, George B Schaller (2011)  Diet of Mongolian gazelles and Tibetan antelopes from steppe habitats using premaxillary shape, tooth mesowear, and microwear analyses.   Mammalian Biology 76: 358-364  
Abstract: The diets of Procapra gutturosa (Mongolian gazelle) and Pantholops hodgsoni (Tibetan antelope) are investigated through premaxillary shape, mesowear, and microwear analyses. The objective of the study is to test the hypothesis that the two species have similarities in dietary habits and also similarities with arctic or subarctic ungulates such as muskox (Ovibos moschatus) and caribou (Rangifer tarandus). Results indicate similarities between Procapra gutturosa and Pantholops hodgsoni, and also some variability in dietary habits related to seasonal and/or regional changes in their habitats. Dental wear patterns for the two species are similar to another ungulate from the arctic steppe environments, the caribou (Rangifer tarandus). This supports the hypothesis that such similarities are reflected in dietary habits because the plant items consumed by those species have similar properties in terms of abrasiveness which are reflected in the tooth wear patterns observed through mesowear and microwear analyses. These results reveal a consistent dietary habit for certain ungulates from this type of arid or dry habitat and climate.
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Marie-Hélène Moncel, Florent Rivals (2011)  On the question of short-term Neanderthal site occupations: Payre, France (MIS 8-7), and Taubach/Weimar, Germany (MIS 5).   Journal of Anthropological Research 67: 47-75  
Abstract: We analyze and compare the evidence of human behavior from two Middle Paleolithic localities with short-term (seasonal) occupations: Payre in France (level F, correlated to MIS 8-7) and Taubach in Germany (correlated to MIS 5e). We focus on the lithic assemblages from these occupation levels. Our analysis takes the density of lithic material, technological choices, and the typological composition of the assemblages in the two localities into account. In light of previously published models, the results are partially consistent with various types of land-use as supported by analysis of the lithic assemblages. Our results confirm that Neanderthals were able to develop diverse behaviors in different locations. Although flexible and highly adaptable among the different seasons and landscapes of Western Europe, different types of short occupations may indicate the same kinds of technical and typological strategies.
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Florent Rivals, Gina M Semprebon (2011)  Dietary plasticity in ungulates: Insight from tooth microwear analysis.   Quaternary International 245: 279-284  
Abstract: In recent years, tooth microwear has been used as a powerful tool for investigating mammalian diets in paleontological or archaeological contexts. Tooth microwear techniques were applied to a number of late Pleistocene assemblages of bison (Bison antiquus) from North America to analyze bison dietary traits, but more particularly, to test for dietary plasticity of the fossil species compared to their modern relatives. Modern bison species are known to be grazers from their ecology. However, the results from tooth wear analysis indicate that dietary traits were more diverse in the fossil bison than in their modern relatives. Bison paleodiets range from pure grazing to mixed feeding. The results illustrate not only the dietary plasticity for that species, but also the potential implications involved when using modern species as analogues for reconstructing the ecology of fossil species. Tooth microwear is a good proxy in archaeological contexts because it gives an insight on the diet of the last days of an animalâs life. The intra-population variability in diet is discussed in relation to the duration of formation of the assemblages (natural assemblages versus archaeological Paleo-Indian sites).
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2010
María-Cristina Fernández-Laso, Florent Rivals, Jordi Rosell (2010)  Intra-site changes in seasonality and their consequences on the faunal assemblages from Abric Romaní (Middle Palaeolithic, Spain).   Quaternaire 21: 155-163  
Abstract: The Middle Palaeolithic site of Abric Romaní (North Eastern Spain) was selected to test the hypothesis that seasonal behaviour of Neanderthal groups will induce differences in the faunal assemblages. The site has a long stratigraphical sequence representing a short chronology, but we focused on levels K, L, and M. The study of seasonality in ungulates indicates that the game was hunted at different periods of the year in each level. Combination of zooarchaeological analyses shows that in all levels the pattern of game procurement and transport to the shelter is similar. In the same way, there is no significant difference related to animal processing. The carcasses introduced into the shelter are used in an exhaustive and systematic way. Although seasonality in the occupation are changing, there are no significant differences in the activities of the Neanderthal groups in the three levels. We conclude that the presence of Neanderthals at Abric Romaní is highly seasonal and their displacements are strongly conditioned by the presence of game preys in the territory.
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Jordi Rosell, Ruth Blasco, Florent Rivals, María Gema Chacón, Leticia Menéndez, Juan Ignacio Morales, Antonio Rodríguez-Hidalgo, Arturo Cebrià, Eudald Carbonell, David Serrat (2010)  A stop along the way: the role of Neanderthal groups at level III of Teixoneres Cave (Moià, Barcelona, Spain).   Quaternaire 21: 139-154  
Abstract: Level III of Teixoneres Cave (Moià, Barcelona, Spain) has provided a significant record belonging to the first half of the Late Pleistocene. It corresponds to both human and carnivorous activities. This assemblage is analyzed through an interdisciplinary approach with the objective to discriminate the remains left by these actors and to evaluate the degree of interaction existing between them. The data indicate hunting activities by hominids as well as by carnivores and a minimal relationship or temporal contact between them inside the cave. This allows the characterization of the human occupations at level III in relation to the composition of the groups and their high mobility in the territory.
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Florent Rivals, Matthew C Mihlbachler, Nikos Solounias, Dick Mol, Gina M Semprebon, John de Vos, Daniela C Kalthoff (2010)  Palaeoecology of the mammoth steppe fauna from the late Pleistocene of the North Sea and Alaska: separating species preferences from geographic influence in paleoecological dental wear analysis.   Palaeogeography, Palaeoclimatology, Palaeoecology 286: 42-54  
Abstract: The paleodietary ecology of Late Pleistocene ungulate faunas of the Mammoth Steppe ecosystem was investigated at Fairbanks (Alaska) and Brown Bank (North Sea) through dental mesowear and microwear analysis. The purpose of the study is to address questions concerning the paleoecology of the Mammoth Steppe, an ecosystem that has no extant analog. Dental wear patterns indicate that the niche partitioning at Brown Bank (BB) region was consistent with ecosystem dynamics found in diverse ungulate faunas in recent time. In contrast, despite the lower numbers of extinct taxa, the Fairbanks (FB) fauna is ecologically bizarre. In general, the microwear of this fauna includes excessive numbers of very (narrow) fine scratches that are atypical for extant mixed feeders and grazers. Moreover, the mesowear signal suggests unusual paleodiets (niche dynamics), where low-crowned mixed feeding cervids, Rangifer and Cervus, adopted diets that are similar to Equus in terms of mesowear (abrasion) and much more abrasive than the diets of either Bison or Ovis. We suggest that the anomalies may relate to unique, possibly disharmonious, ecosystem dynamics during the Pleistocene-Holocene transition. In addition to these findings, it was discovered (found) that dental wear patterns, particularly microwear variables such as scratch frequency and scratch width are strongly affected by geographic region, irrespective of species-specific dietary preferences
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Florent Rivals, Almudena Arellano (2010)  Les faunes des sites du Pléistocène supérieur du Caucase méridional : Grotte de Sakažhia, Grotte d’Ortvala et Grotte du Bronze (République de Géorgie).   L'Anthropologie 114: 305-323  
Abstract: The paleontological analysis of the fauna from three Late Pleistocene localities in Southern Caucasus (Sakažhia, Ortvala, and Bronze cave) confirmed the presence of Ursus spelaeus, Canis lupus, Equus ferus, Rhinocerotidae, Cervus elaphus, Capreolus capreolus, Alces alces, Bison priscus, Capra caucasica, Ovis ammon and Sus scrofa. The study also permitted the identification of new taxa for these localities. There may be the presence of reindeer (Rangifer tarandus) at Sakažhia, however it has not yet been confirmed. Moreover, remains of aurochs (Bos primigenius) were identified, in particular at Bronze cave where its abundance is exceptional. Indeed, Bos and Bison are generally very rare in Southern Caucasus. The analysis of the proportion of the different species in each locality revealed the two types of occupations of the caves, one dominated by hominids and the other by carnivores. At Sakažhia, where fauna is dominated by the presence of cave bear, the cave was seasonally occupied by Neanderthal groups. On the other hand, Bronze cave corresponds to a habitat of hunters, which occupied the site for longer periods.
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Florent Rivals, Gina M Semprebon (2010)  What can incisor microwear reveal about the diet of ungulates?   Mammalia 74: 401-406  
Abstract: We analyzed dental wear patterns on molars and incisors of modern plains zebra (Equus quagga) from a single locality in Kenya and from a single breeding population. Our objective was to explore the potential of incisor microwear in terms of elucidating ungulate diet as well as the various chewing patterns of different incisors and molar teeth among various age classes. We found significant inter-tooth variation of the microwear signal among different incisors and, not unexpectedly, that incisors do not reveal the same dietary signal as molar teeth. Such differences are related here to differential chewing forces (shearing vs compression) along the tooth row. We find no significant differences among various zebra age classes in terms of their mesowear and microwear patterns, providing additional evidence that the mesowear signal is consistent and stable throughout the adult lifespan of an individual. These findings indicate that there is no age segregation in terms of access to food resources in this particular breeding population.
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Jordi Rosell, Ruth Blasco, Florent Rivals, Arturo Cebrià, Juan Ignacio Morales, Antonio Rodríguez, David Serrat, Eudald Carbonell (2010)  Las ocupaciones en la Cova de les Teixoneres (Moià, Barcelona): Relaciones espaciales y grado de competencia entre hienas, osos y neandertales durante el Pleistoceno superior.   Zona Arqueológica 13: 392-402  
Abstract: At European caves with Pleistocene deposits, is frequent to find a mixture of human and carnivore activities. This phenomenon has led many researchers to interpret these assemblages as the product of competition in the environment between them, not only for preys or others resources, but also for the space. In the present work we analyze this problem using data from Teixoneres Cave. This cave, located near the town of Moià (Barcelona, Spain), presents a wellstratified sedimentary fill originated during the last quarter of Middle Pleistocene and ended in the second half of Upper Pleistocene. At the moment of their discovery during the 50âs of last century, the cave was interpreted as a hyena den with few visits of hominids. The human presence is documented by some lithic artefacts, faunal remains and presence of isolated hearths. The new research project, developed since 2003, has served to identify the activities of multiple carnivores in the cavity (principally hyena, cave bear, lynx, fox and badger) and the sporadic character of the human occupations. However, the archaeo-stratigraphical works and the spatial distribution of remains suggest a clear temporal separation in the areas used in the cave. On the one hand, human activities tend to be clustered in the main entrance of the cave and are characterized by the scarcity of remains and the limitation of used space. On the other hand, the carnivores develop theirs occupations in interior without showing clear preferences for any particular place. So far, we did not recognise evidences of temporal or spatial relationship among both agents at any stratigraphical level, and therefore, we can not arise the existence of competence between hominids and carnivores at Teixoneres Cave.
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Nikos Solounias, Florent Rivals, Gina Semprebon (2010)  Dietary interpretation and paleoecology of herbivores from Pikermi and Samos (late Miocene of Greece).   Paleobiology 36: 113-136  
Abstract: A large sample of the Pikermi and Samos ungulates was examined by microwear analysis using a light stereomicroscope (561 extinct and 809 extant comparative specimens). The results were used to infer the dietary adaptations of individual species and to evaluate the Pikermian Biome ungulate fauna. Many of the bovids have wear consistent with mixed feeding, although a few mesodont taxa apparently enjoyed an exclusive browsing and or grazing diet. The giraffids spanned the entire dietary spectrum of browsing, mixed feeding, and grazing, but most of the three-toed horses (Hippotherium) were hypsodont grazers. The colobine monkey Mesopithecus pentelici displays microwear consistent with a mixed fruit and leaf diet most likely including some hard objects. Similar results were obtained from prior scanning electron microscopy microwear studies at 500 times magnification and from the light microscope method at 35 times magnification for the same species. Results show that diet can differ between species that have very similar gross tooth morphology. Our results also suggest that the Pikermian Biome was most likely a woodland mosaic that provided a diversity of opportunities for species that depended on browsing as well as species that ate grass. The grasses were most likely C3 grasses that would grow in shaded areas of the woodland, glades, and margins of water. The ungulate component of the Pikermi and Samos fauna was more species-rich and more diverse in diet than the ungulates observed in modern African forests, woodlands, or savannas, yet dietarily most similar to the ungulates found in woodland elements of India and to some extent of Africa. It is unlikely that the Pikermi and Samos ungulates inhabited dense forests because we find no evidence for heavy fruit browsing. Conversely, a pure savanna is unlikely because many mixed feeders are present as well as browsers. Extant woodland African species are morphologically and trophically very similar to the African savanna species. Therefore the evolution of grazing and of hypsodont morphology for Africa may have evolved within the Plio-Pleistocene woodlands of Africa. Our results show that major dietary and morphologic ungulate evolution may take place within woodlands rather than as a consequence of species moving into savannas both during the late Miocene of Pikermi and Samos and during the PleistoceneâRecent of Central Africa.
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Gina M Semprebon, Florent Rivals (2010)  Trends in the paleodietary habits of fossil camels from the Tertiary and Quaternary of North America.   Palaeogeography, Palaeoclimatology, Palaeoecology 295: 131-145  
Abstract: Dietary adaptations of both Tertiary and Quaternary representatives of North American Camelidae were examined through deep evolutionary time (via hypsodonty index), though ecological time (via mesowear analysis), and through the last few days of life (via microwear) by examining molar teeth. Fossil samples are from the Great Plains, Great Basin, Arizona, and Florida and span from the early late Eocene (late Chadronianâearly Orellan) to the late Pleistocene (Rancholabrean). Results were compared to those obtained on modern camels and llamas and other ungulates of known dietary behavior. Camels apparently exploited open habitats early on in their evolution as evidenced by the extreme pitting of their enamel surfaces. Grasses were likely consumed early on in their history (e.g., Poebrotherium) but the vast majority of taxa were committed browsers. Results show that the hypsodonty pattern (deep time adaptation) and mesowear pattern (cumulative abrasion index) are very similar. Hypsodonty indices and mesowear scores decrease in the middle Miocene, a time when a few taxa also incorporate fruit and/or seeds in their browse. Crown height and dietary abrasion increase in the late Miocene and Pliocene, a time when some grazers and mixed feeders also appear, but then decrease in Pleistocene and Recent forms.
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2009
Florent Rivals, Ellen Schulz, Thomas M Kaiser (2009)  Late and Middle Pleistocene ungulates dietary diversity in Western Europe indicate variations of Neanderthal paleoenvironments through time and space.   Quaternary Science Reviews 28: 3388-3400  
Abstract: Mesowear and microwear on enamel from 763 teeth of middle and late Pleistocene ungulates were analysed to infer the potential of dental wear analysis of faunal remains as a paleoenvironmental and paleoclimatic proxy in relation to climatic changes and diversity of vegetation available in the environment. Fossil localities including levels belonging to two glacial and two interglacial stages were selected in Germany, France, and Spain. At a temporal scale, results indicate that the dietary diversity in ungulates is higher during interglacial phases (MIS 5 and 3) than during pleniglacial phases (MIS 8 and 4). Dietary diversity is concluded to be related to climate-driven vegetation changes which during interglacials lead to increased variety of potential food items available to ungulates. At the geographical scale, during interglacials, changes in diet composition are evident along geographical gradients. The corresponding dietary gradients are proposed to be related to climate and vegetation gradients reflecting more arid climates in the Mediterranean area compared to North-Western Europe. Species consistently represented at all localities investigated are Cervus elaphus (Cervidae, Artiodactyla) and Equus ferus (Equidae, Perissodactyla). C. elaphus populations are found to consistently have less abrasive diets than E. ferus populations but dietary traits of both species varied largely, revealing a significant plasticity in the feeding adaptation of both species. Those traits are concluded to be related to differences in vegetation structure at each locality and complement the evidence that ungulates have broader dietary habits than what is usually assumed.
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Florent Rivals, Marie-Hélène Moncel, Marylène Patou-Mathis (2009)  Seasonality and intra-site variation of Neanderthal occupations in the Middle Palaeolithic locality of Payre (Ardeche, France) using dental wear analyses.   Journal of Archaeological Science 36: 1070-1078  
Abstract: The reconstruction of Palaeolithic subsistence and settlement systems at the Middle Palaeolithic site of Payre (France) is undertaken through the application of dental wear analyses combined to zooarchaeological, technological, and ecological indicators. Three archaeological levels were investigated. Level D, dated to MIS 5, could correspond to an occupation during the cold season. In levels G and F, dated to MIS 8-7, animals were probably hunted during a warmer season. According to dental wear analyses, it is likely that level F has actually recorded a succession of short occupations, contrary to what is observed in the levels G and D. Those differences in the duration of site occupation are discussed in the light of previous sedimentological, zooarchaeological, and technological studies.
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Florent Rivals, Ellen Schulz, Thomas M Kaiser (2009)  A new application of dental wear analyses: estimation of duration of hominid occupations in archaeological localities.   Journal of Human Evolution 56: 329-339  
Abstract: Characterization of settlement patterns is one of the core concepts in archeological research. The duration of an occupation is usually estimated through zooarchaeology (e.g., density of remains, cementochronology) and is limited by taphonomic processes and sample size. We propose a new application of dental wear methods for estimating the relative duration of hominid settlements in Paleolithic sites. Dental microwear is known to be sensitive to seasonal changes in diet. In this new application we use microwear scratch counts to estimate the variation in the dietary signal of various ungulate species. We propose that this variation is correlated to the duration of site occupation. Each season presents a limited and different set of food resources available in the environment. If animals are sampled only during a specific season (i.e., during a short term occupation) then they would be expected to have a dental wear signal with little variation. On the other hand, a greater diversity of food is available across different seasons. Therefore, if game animals are hunted through various seasons during long occupation periods, then they would be expected to have more variable dental wear. The application of this technique to the Middle Paleolithic site of Arago Cave (France), where various types of occupations occurred, supports this hypothesis. When combined with multidisciplinary studies of archaeological localities (seasonality in particular), this new application of dental wear analysis presents valuable information about hominid settlements and behavior. We contextualize our data with results from lithic and zooarchaeological analyses from Arago. These results reveal the presence of both high and low mobility groups of Homo heidelbergensis throughout the sequence of the Arago Cave.
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2008
Florent Rivals, Ellen Schulz, Thomas M Kaiser (2008)  Climate-related dietary diversity of the ungulate faunas from the middle Pleistocene succession (OIS 14-12) at the Caune de l'Arago (France).   Paleobiology 34: 117-127  
Abstract: The Caune de lâArago Cave (southern France) has yielded one of the best preserved and best documented sedimentary successions of the European Middle Pleistocene (Oxygen Isotopic Stages 14 to 12). Herbivorous ungulates (horse, reindeer, red deer, fallow deer, bison, musk ox, argali, and tahr) are well represented in the three major stratigraphic units CM1, CM2, and CM3. CM1 and CM3 correspond to cold and dry climate and CM2 represents temperate and humid environmental conditions. Dental microwear and mesowear analyses were performed for the ungulates from CM1â3 to test whether these methods of dental wear evaluation were suitable for detecting climate-driven changes in the dietary resources of the Arago ungulate community. We found that both dental mesowear and microwear indicate dietary traits and their relationship to climatic conditions as reflected by vegetation cover and community structure. In all units, even if some species seemto share habitats or resources, it appears that the overlap in their feeding ecology is very low. The CM1 and CM3 units, where pollen analysis indicates that the climate was cold and dry, show the lowest diversity in dietary traits. The CM2, where climate is known to be more temperate and humid, the spectrum of dietary traits is largeâgrazers, browsers, and mixed feeders are present.
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Florent Rivals, Athanassios Athanassiou (2008)  Dietary adaptations in an ungulate community from the late Pliocene of Greece.   Palaeogeography, Palaeoclimatology, Palaeoecology 265: 134-139  
Abstract: The dietary morphological methods of mesowear and microwear were applied to ungulates of the Late Pliocene fauna of Sésklo (Thessaly, Greece). The results provide evidence for the predominance of open grassland in the area, as the most common species, <i>Equus stenonis</i>, was a strict grazer. The rare cervid cf. <i>Croizetoceros ramosus</i> was the only browser. The antelopes (genera <i>Gazella</i> and <i>Gazellospira</i>) yielded discrepant microwear and mesowear results. This is interpreted as an indication of regional or seasonal dietary resource differentiation, inferring that the antelopes were probably mixed feeders that grazed occasionally or periodically.
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Florent Rivals, Ruth Blasco (2008)  Presence of Hemitragus aff. cedrensis (Mammalia, Bovidae) in the Iberian Peninsula: Biochronological and biogeographical implications of its discovery at Bolomor Cave (Valencia, Spain).   C. R. Palevol 7: 391–399  
Abstract: The discovery of new material in late Pleistocene levels at Bolomor Cave (Valencia, Spain) raises some questions about the presence the most ancient record of <I>Hemitragus cedrensis</I> in the peninsula, and its dispersal out of Provence. The morphology and dimensions of some lower teeth confirm the identification of <I>H.</I> aff. <I>cedrensis</I>. Moreover, it presents strong similarities, both morphological and metrical, with the specimens from the Caune de l'Arago and Bau de l'Aubesier (end of OIS 7 to OIS 5e) rather than with the population from the eponymous locality. The data suggest a dispersal event out of Provence towards the Iberian Peninsula during the Eemian. This dispersal was not stopped by natural barriers such large rivers, or mountains. The results presented confirm the biochronological interest of the genus <I>Hemitragus</I> for the Late Pleistocene in Mediterranean Europe.
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2007
Florent Rivals, Nikos Solounias (2007)  Differences in tooth microwear of populations of caribou (Rangifer tarandus, Ruminantia, Mammalia) and implications to ecology, migration, glaciations and dental evolution.   Journal of Mammalian Evolution 14: 182-192  
Abstract: Tooth microwear was analyzed for a large sample of wild-shot barren-ground caribou (Rangifer tarandus groenlandicus) from the Kaminuriak population of eastern Canada. This sample was compared to the microwear of specimens from three Pleistocene localities in North America (Alaska) and western Europe (Caune de lâArago in France and Salzgitter in Germany). The results show that the extant samples from eastern Canada have seasonal variation in microwear and presumably in diet. The differences in microwear between the various seasons may reflect a cyclic migration of the population within a year. The extinct population from Alaska has extremely blunt teeth (mesowear), as blunt as those of modern zebras and bison. This observation is corroborated by the lowest number of microwear pits. The findings are untypical, as most typical caribou teeth have sharper apices, and we interpret this as an indication of a local habitat that was different with animals feeding on non-typical vegetation. The combination of Rangifer from Caune de lâArago and Salzgitter reveals a pattern in microwear variability. The Salzgitter is interglacial and shows a greater diversity of browsing (broad spectrum on average number of pits) than the glacial Caune de lâArago. The interglacial population from Salzgitter is interesting because it shows several different types of browsing. Collectively all the Rangifer teeth show that diet of a brachydont taxon can vary across most of the dietary morphospace of ungulates as represented by tooth microwear. The three Pleistocene samples exhibit microwear that is different from the extant population in question. This observation implies that the recent diet of Rangifer has changed from the typical caribou diet in the past. This indicates dietary change within a species. This is important because it represents dietary evolution without changes in tooth morphology.
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Florent Rivals, Nikos Solounias, Matthew C Mihlbachler (2007)  Evidence for geographic variation in the diets of late Pleistocene and early Holocene Bison in North America, and differences from the diets of recent Bison.   Quaternary Research 68: 338-346  
Abstract: During the late Pleistocene and early Holocene, Bison was widely dispersed across North America and occupied most regions not covered by ice sheets. A dietary study on Bison paleopopulations from Alaska, New Mexico, Florida, and Texas was performed using two methods that relate dental wear patterns to diet, mesowear analysis and microwear analysis. These data were compared to a mixed sample of extant Bison from the North American central plains, extant wood Bison from Alberta (Canada) and a variety of other modern ungulates. Mesowear relates macroscopic molar facet shape to levels of dietary abrasion. The mesowear signature observed on fossil Bison differs significantly from the hyper-abrasive grazing diet of extant Bison. Tooth microwear examines wear on the surface of enamel at a microscopic scale. The microwear signal of fossil samples resembles to modern Bison, but the fossil samples show a greater diversity of features, suggesting that fossil Bison populations regularly consumed food items that are texturally inconsistent with the short-grass diet typical of modern plains Bison. Mesowear and microwear signals of fossil Bison samples most closely resemble a variety of typical mixed feeding ungulates, all with diets that are substantially less abrasive than what is typical for modern plains Bison. Furthermore, statistical tests suggest significant differences between the microwear signatures of the fossil samples, thus revealing geographic variability in Pleistocene Bison diets. This study reveals that fossils are of value in developing an understanding of the dietary breadth and ecological versatility of species that, in recent times, are rare, endangered, and occupy only a small remnant of their former ranges.
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Gina M Semprebon, Florent Rivals (2007)  Was grass more prevalent in the pronghorn past? An assessment of the dietary adaptations of Miocene to recent Antilocapridae (Mammalia : Artiodactyla).   Palaeogeography, Palaeoclimatology, Palaeoecology 253: 332-347  
Abstract: Molar teeth of both Tertiary and Quaternary representatives of the family Antilocapridae were examined using mesowear and light stereomicroscopy microwear. Taxa from the Central and Southern Great Plains and the Northern and Southern Great Basin were included in the analyses and results compared to those obtained on the modern pronghorn (Antilocapra americana). Species included spanned from the early Miocene (late Hemingfordian) to the late Pleistocene (Rancholabrean). Results are concordant with well-known trends toward increasing aridity and shifts in vegetational structure in the late Mioceneâearly Pliocene of North America. Both mesowear and microwear results indicate a shift toward more abrasive diets beginning in the Hemphillian (late MioceneâPliocene) and then a return to a less abrasive dietary regime for the duration of the Pleistocene and into the Recent. The more derived antilocaprines (more hypsodont, relatively longer limbs) apparently depended more on grass than the less advanced merycodontines, but even the earliest of the latter seem to have relied more on grass as a dietary staple than the modern pronghorn. Seasonal grit encroachment on food items encountered by fossil antilocaprids coupled with a heavier reliance on grasses may provide a possible explanation for the extreme hypsodonty present in the modern pronghorn despite its mainly browsing dietary behavior.
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2006
Florent Rivals, Agnès Testu (2006)  Un nouveau gisement paléontologique à Capra caucasica praepyrenaica : la grotte de l’Arche à Bugarach (Aude, France).   C. R. Palevol 5: 711-719  
Abstract: A new palaeontological locality with Capra caucasica praepyrenaica: The Arche cave at Bugarach (Aude, France). The Arche cave is a new palaeontological locality situated in the Corbières (southern France). This cave has yielded an abundant and well-preserved large mammal fauna. A first palaeontological description of the remains from surface excavation is provided. According to biochronological data, the site is dated from MIS 5 to MIS 4 and contains very uncommon species of wild goat, Capra caucasica praepyrenaica, in the eastern part of the Pyrenees.
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Florent Rivals (2006)  Découverte de Capra caucasica et d’Hemitragus cedrensis (Mammalia, Bovidae) dans les niveaux du Pléistocène supérieur de la Caune de l'Arago (Tautavel, France) : implication biochronologique dans le contexte du Bassin Méditerranéen.   Geobios 39: 85-102  
Abstract: Two species of the Caprini tribe, Hemitragus cedrensis and Capra caucasica, were identified in the top deposits of the Caune de l'Arago (Pyrénées-Orientales, France) Morphological and metrical analyses have shown how these two species evolved in this area. Owing to these results the Caune de l'Arago deposits can be correlated with the end of OIS 5 or the beginning of OIS 4 (corresponding to MNQ 25 and MNQ 26 biozones). The size reduction of these Caprinae, observed at the Caune de l'Arago and in two other sites, suggests a possible local evolution.
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Anne-Marie Moigne, Maria Rita Palombo, Véronique Belda, Djamila Heriech-Briki, Sarah Kacimi, Frédéric Lacombat, Marie-Antoinette de Lumley, José Moutoussamy, Florent Rivals, Jérôme Quilès, Agnès Testu (2006)  Les faunes de grands mammifères de la Caune de l'Arago (Tautavel) dans le cadre biochronologique des faunes du Pléistocène moyen italien.   L’Anthropologie 110: 788-831  
Abstract: The fossiliferous levels of Caune de l'Arago supply a useful tool to investigate the relationships between faunal renewals and climate changes in a restricted area during the Middle Pleistocene. On the other hand the Italian fossil assemblages of the middle Galerian testified changes of palaeoenvironmental conditions. A biochronologic and palaeoecologic comparison among large mammalian faunas of these quite different Mediterranean regions have been performed in the aim to value possible vicariance of taxa and differences of faunal renewal dynamic during the Middle Pleistocene. After the results of our analysis we can correlate the faunal assemblages from the âComplexe moyen, ensemble stratigraphique Iâ, âComplexe moyen, ensembles stratigraphiques II and IIIâ, âComplexe supérieur, sols A and Bâ of Caune de L'Arago sedimentary succession with the Italian Isernia (middle Galerian), Fontana Ranuccio (late Galerian) and Torre in Pietra (early Aurelian) faunal units (FUs) respectively. The French faunal assemblages were characterised by the absence of browsers and, conversely, by the presence of boreal taxa, mainly grazers, that were rare or never occurred in Italy. Accordingly, at no one Italian faunal complex, as expected, we can find any indication of continental cold climatic condition as showed by the faunal complexes of the Caune de l'Arago. Nevertheless, the major climate changes characterising the Middle Pleistocene can be detected in both regions. During the span time of Isernia FU, the temperature was at its lowest and arid conditions prevailed, whereas during the following Fontana Ranuccio FU humidity and temperature increased, as is more evident in central Italy local faunas. Moreover, after the similarity analysis, the Visogliano local fauna (North-Eastern Italy) was more related to the French than to the Italian ones. During the late Middle Pleistocene (Torre in Pietra FU) a moderate faunal reconstruction can be detected in both regions. The renewal was linked to climatic modifications taking place in the Mediterranean area approximately around OIS 11/OIS 10 transition, when interstadials became progressively milder and average humidity increased.
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Florent Rivals, Agnès Testu, Anne-Maire Moigne, Henry de Lumley (2006)  The Middle Pleistocene argali (Ovis ammon antiqua) assemblages at the Caune de l'Arago (Tautavel, Pyrénées-Orientales, France) : are prehistoric hunters or carnivores responsible for their accumulation?   International Journal of Osteoarchaeology 16: 249-268  
Abstract: The argali (Ovis ammon antiqua) assemblages from the Middle Pleistocene site of the Caune de l'Arago (Tautavel, southern France) were studied in terms of zooarchaeology and taphonomy. It is possible to discern palaeobiological information lost during fossilisation, as well as the palaeoethology of the bone collector, by the observation of taphonomic details preserved on the bone assemblages. The observations leave no doubt that both humans and carnivores were involved in the accumulation of argali carcasses in the cave. In some assemblages, the type of bones found in articulation and the gnawing marks observed are characteristic of carnivores. In other levels, the intense fracturing of the major limb bones in relation to their marrow content and mineral density, and butchering marks found on specimens in the earlier levels, are in favour of human accumulation, the modalities of which are discussed. The results suggest that the degree of carnivore activity seems to have been higher in levels M, N and O than in level F.
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2005
Florent Rivals, Brigitte Deniaux (2005)  Investigation of human hunting seasonality through dental microwear analysis of two Caprinae in late Pleistocene localities in Southern France.   Journal of Archaeological Science 32: 1603-1612  
Abstract: Dental microwear analysis was used to interpret the diet of two species of Caprinae, Capra caucasica and Rupicapra pyrenaica in three Mousterian sites (120â35 Kyr) of Southern France. Both species, which are representative of the hunted faunal assemblages, were found to be seasonal mixed-feeders. During a phase of cold climate, the differences observed in the proportion of various microwear features can be explained only by differences in diet. These microwear changes were found to be correlated with hunting periods which are different between the sites. Our findings confirm former observations made using other methods and show how dental microwear analysis can be applied to explore seasonality in zooarchaeological and paleoenvironmental reconstructions.
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2004
Henry de Lumley, Sophie Grégoire, Déborah Barsky, Gérard Batalla, Salvador Bailon, Véronique Belda, Djamila Briki, Louise Byrne, Emmanuel Desclaux, Khalid El Guenouni, Alain Fournier, Sarah Kacimi, Frédéric Lacombat, Marie-Antoinette de Lumley, Anne-Marie Moigne, José Moutoussamy, Cristina Paunescu, Christian Perrenoud, Véronique Pois, Jérôme Quiles, Florent Rivals, Thierry Roger, Agnès Testu (2004)  Habitat et mode de vie des chasseurs paléolithiques de la Caune de l’Arago (600 000 - 400 000 ans).   L'Anthropologie 108: 159-184  
Abstract: Several human occupations have been identified within the deposits of the Caune de lâArago Cave site, dated from 700 000 years to 100 000 years old. The exceptional wealth of these archeological levels provides an opportunity for the study of different types of habitat and behavioral patterns of Paleolithic hunters between 600 000 and 400 000 years ago. Results from preliminary studies, carried out over the last 30 years on these archeological levels, allow for the proposition of a typology for cave habitats during the Middle Pleistocene in the south of France. In fact, four types of habitat were identified and defined. They have been described according to different characteristics such as, the thickness of the archeological accumulation, the animal species represented, the season during which the animals were hunted, the state of conservation of the fossils discovered, the morpho-technological and typological characteristics of the lithics, the categories of raw materials used and the territory exploited by the hominids inhabiting the cave, as defined by the raw materials represented. The four habitation types defined are: the long duration habitat, where the cave was used as a home base; the temporary seasonal habitat, in which case the cave served as a secondary campsite; the hunting stopover and the bivouac, during which the cave was used as a refuge for short term stays. Behavioral patterns appear to be directly related to the type of habitat and oriented towards a principal activity: hunting. The activities performed by the inhabitants of the cave seem to be more diversified during prolonged or seasonal occupations, whereas they appear to be reduced or very specialised during brief stays. Generally, The characteristics studied show that the Caune de lâArago Cave site had multiple uses over time for Anteneandertalian family groups and hunters who left traces of their activities, allowing us to better understand their lifestyle.
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Florent Rivals, Sarah Kacimi, José Moutoussamy (2004)  Artiodactyls, favourite game of prehistoric hunters at the Caune de l'Arago Cave (Tautavel, France). Opportunist or selective hunting strategies?   European Journal of Wildlife Research 50: 25-32  
Abstract: An archaeological study of the artiodactyls was carried out in the richest layers of the Caune de l'Arago Cave in France, namely levels L, J, G (upper) and F. The fauna of these levels is dominated by the reindeer, the Cervini (red deer and fallow deer), the musk ox and the argali, respectively. Selective hunting, among other factors, can be responsible for the relative abundance of some population categories, such as males or females, young or adults, but other factors can also be responsible. We used the eco-ethological information obtained by studying present-day wild herds to determine the structure (age and sex) of a theoretical population for a specified season. The season of hunting is estimated through the study of tooth eruption, and this season is shown to have a severe impact on the composition of the studied animal populations. By comparing the theoretical population obtained for the selected season with our results on fossil materials, we demonstrate that there is no selection by hunters on the site.
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2003
Florent Rivals, Brigitte Deniaux (2003)  Dental microwear analysis for investigating the diet of an argali population (Ovis ammon antiqua) of mid-Pleistocene age, Caune de l'Arago cave, eastern Pyrenees, France.   Palaeogeography, Palaeoclimatology, Palaeoecology 193: 443-455  
Abstract: In recent years, dental microwear analysis has been widely used as a method for reconstructing ancient diet. The current study reconstructs the diet of Ovis ammon antiqua, a 440 000 year old argali-like sheep fossil from the Pleistocene site of the Caune de lâArago in Southern France. The microscopic wear was observed on second upper molars with an environmental scanning electron microscope, using a magnification of Ã500. Data analysis was performed on the following microwear variables: feature density, pit density, striation density, pit/striation ratio, pit size, and striation length and width. The microwear pattern of O. ammon antiqua was compared with those of extant ruminant taxa. Fossil wild sheep have significantly lower pit densities but similar scratch densities. The fossil animal shows shorter and narrower striations than extant herbivores. Our results suggest that fossil argali was a typical grazer, more so than extant ruminants. As far as diet variations are concerned, the diet of extant Caprinae varies according to season and region. In our fossil record, diet variations reflect some of the climatic changes that occurred during the Quaternary. This study shows that microwear analysis is not only a good tool for determining diet adaptations, but that it also brings indirect evidence of climatic variations.
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2002
Florent Rivals, Anne-Marie Moigne, Henry de Lumley (2002)  La chasse aux petits bovidés à la Caune de l'Arago (Tautavel, France) : opportunisme ou sélection des proies?   Anthropozoologica 36: 3-12  
Abstract: The Caune de lâArago, a Lower Palaeolithic cave, contains several archaeological levels with a large assemblage of argali sheep and tahr. The study of bone remain spatial distribution was made possible by using a computer method. Three levels are individualised in the middle complex, the layers E, F and G, and the latter is subdivided into three archaeostratigraphical levels. Fossil remains from all identified levels were studied in order to determine the number of hunted animals, their age and sex, and the hunting season. These data characterize argali and tahr populations for each level. A comparative study of these populations helped to verify that hunting was selective and to better understand the behaviour and living mode of the Lower Palaeolithic hunters.
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Book chapters

2012
2011
Maria Cristina Fernández-Laso, Maria Gema Chacón, Maria Dolores García-Antón, Florent Rivals (2011)  Territorial mobility Neanderthals groups: A case study from level M of Abric Romaní (Capellades, Barcelona, Spain).   In: Neanderthal lifeways, subsistence and technology. Edited by:N.J. Conard and J. Richter. pp. 187-202. Springer, Dordrecht. isbn:9789400704145  
Abstract: The Abric Romaní site is located in the northeast Iberian Peninsula, 50 km from Barcelona 317 m above sea level, in a travertine cliff on the right bank of the Anoia river, which is a tributary of the Llobregat river. This river passes by the Capellades locality, forming a narrow gorge known as âCinglera del Capellóâ. This corridor crosses the mountain range âCordillera Prelitoralâ, and is considered as a natural passage between the inland and coastal Catalonia regions. This situation provides the possibility to exploit a great variety of biotopes. The site yielded an important and complete stratigraphic sequence of European Middle Paleolithic. Fifteen archeological levels have been excavated until today, which appear as thin layers between sterile travertine platforms. This sequence has been dated by U-series between 70 and 40 ka BP, except for the uppermost level, which is attributed to the Early Upper Paleolithic. Pollen analyses have revealed five climatic phases, ranging from the milder conditions at the bottom to an interstadial climate at the top, correlated with the Hengelo interstadial. It is, consequently, a key site for human behavioral studies among Neanderthal groups. This paper presents the results of studies on resource procurements, lithics and animals and on the territory used by Neanderthal groups, through the example of level M dated around 55 ka BP. The results of the lithic analysis and faunal assemblages suggest that the resource procurement and management is focused on a local and semi-local exploitation in a geographic area of about 20 km around the rock-shelter.
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Conference papers

2007
Florent Rivals, Anne-Marie Moigne (2007)  Saison d'occupation et stratégies de chasse au Paléolithique inférieur, l'exemple de la Caune de l'Arago (Tautavel, Pyrénées-Orientales)   In: Aires d'approvisionnement en matières premières et aires d'approvisionnement en ressources alimentaires. Approche intégrée des comportements, Actes du XV Congrès Mondial (Lisbonne, 4-9 Septembre 2006) Vol. 5, Session WS23 Edited by:M-H Moncel, A-M Moigne, M Arzarello, C Peretto. 17-25 British Archaeological Reports, International Series S1725, Archaeopress, Oxford  
Abstract: Zooarchaeological data on game procurement strategies (scavenging, opportunistic hunting, selective hunting) for Lower Palaeolithic sites are scarce. Few localities have been analysed in a zooarchaeological way for two main reasons. First of all, because the study of game procurement strategies by hominids requires large-sized samples. Secondly, the stratigraphic context and origin of the remains must be clearly defined. The Caune de lâArago cave (Tautavel, France) meets those two conditions. The levels E, F, and G, correlated to the OIS 12, correspond to human occupation levels. For those levels, the structure of the hunted populations (age and sex ratio) and seasonality were done of the argali (Ovis ammon antiqua) and the tahr (Hemitragus bonali). Seasonality of hunting was studied using teeth eruption and wear (Rivals, 2004), but also through dental microwear analysis (Rivals and Deniaux, 2005). The relative proportion of males vs. females, and young vs. adults is related to the season when they were killed. In the level F, for example, groups of females and young argali were killed in spring and summer, when the males live alone and away from the group. In this case, mortality season explain the low proportion adult males in the sample. Combined study of the population structure and the mortality season suggest an opportunistic hunting strategy on argali and tahr in several levels at the Caune de lâArago. Environment plays a fundamental role in the analysis of game hunting strategies, and especially the hunting season, which is a key parameter in zooarchaeological studies.
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2005
Florent Rivals, Anne-Marie Moigne, Brigitte Deniaux (2005)  Caractérisation des variations paléo-environnementales d'après l'alimentation des caprinés chassés par les hommes du Paléolithique inférieur et moyen en Languedoc-Roussillon   In: Données récentes sur les modalités de peuplement et sur le cadre chronostratigraphique, géologique et paléogéographique des industries du Paléolithique ancien et moyen en Europe, Archaeopress, Oxford Edited by:N Molines, M-H Moncel, J-L Monnier. 115-123 British Archaeological Reports, International Series S1364  
Abstract: The diet of three species of Caprinae is studied through dental microwear analysis in three Upper Pleistocene sites of the North Mediterranean area. Microwear analysis complement the classical palaeoenvironmental results obtained with the cenogram method. Microwear analysis lead us to detect climatic changes which were not perceived through neither the study of faunas nor sedimentological analysis.
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2004
Jacques Pernaud, Jérôme Quiles, Florent Rivals (2004)  La faune de la fin du Pléistocène dans la haute vallée de l'Aude : L'exemple de la grotte du Castel 2 à Bessède-de-Sault (Aude, France)   In: Acts of the XIVth UISPP Congress, University of Liège, Belgium. Section 3: Paléoécologie / Paleoecology, General sessions and posters. Archaeopress, Oxford Edited by:Le Secrétariat du Congrès. 31-37 British Archaeological Reports BAR S1271  
Abstract: The Castel 2 cave is found at a middle mountain altitude, in the high valley of the river Aude (France). The excavation yielded numerous carnivore remains (hyena and cave bear) and small bovid remains, especially Capra pyrenaica. The different bones show that the site may date of the end of Pleistocene.
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2003
Florent Rivals, Agnès Testu, Véronique Belda (2003)  Le rôle des grands Carnivores dans l'accumulation de l'argali des niveaux inférieurs de la Caune de l'Arago (Pyrénées-Orientales, France)   In: Acts of the XIVth UISPP Congress, University of Liège, Belgium. Archaeopress, Oxford Edited by:M Patou-Mathis, H Bocherens. 163-169 British Archaeological Reports International Series S1105  
Abstract: Were big Carnivores responsible of the argali accumulation in the lower levels of the Caune de l'Arago (Pyrénées-Orientales, France)? The MNO levels from the Caune de l'Arago, recently brought to light and dated about 550,000 years, delivered numerous argali remains (Ovis ammon antiqua) associated with eight species of carnivores (Ursidae, Felidae and Canidae). These bones are very well preserved and were mostly found in anatomical connection. The palaeodemographic study (age and sex profiles, estimation of the living animal weight) reveals a population made of strong size males exclusively. Bone surface examination doesn't show any mark resulting from human activities (cut marks or percussion marks). On the other hand, many carnivore tooth and gnaw marks were revealed. The skeletal part representation indicates that this accumulation was not produced by prehistoric hunters. All these elements tend to assess that the agent responsible for the accumulation of argali remains in the MNO levels was probably the big Carnivores.
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2001
Florent Rivals (2001)  The argali of the "Caune de l'Arago" (Southern France). Palaeoecology of a 440,000 years old population   In: Proceedings of the third international symposium on mouflon. Sopron, Hungary, October 27-29, 2000 Edited by:A Náhlik, W Uloth. 103-113  
Abstract: The âCaune de lâAragoâ is a Middle Pleistocene prehistoric site located at Tautavel, France. As a result of the excavation performed in this cave, a large number of fossils and stone tools were brought to light. Abundant wild sheep remains, discovered together with four complete crania, are attributed to Ovis ammon antiqua (Argali). The vertical distribution of the remains was studied and indicates that Ovis was present from 690,000 to 400,000 years ago. The fossil population was modified by the hunter-gatherer behaviour of Homo erectus. A study of distribution, age and sex parameters and pathologies was made possible by a very large number of bones and teeth in the F level (about 5,300 remains). This level, dating from 440,000 years, contains 75 % of Argali among the large Mammals. Individual age was determined from teeth. The age profile is of hunting mortality type and according to the Stiner method this profile is of catastrophic type i.e. a distribution close to a living assembly. Besides natural disasters, this structure can be the result of ambush hunting when prey selection is determined by chance. Crown height measurements are used to define cohorts among a population. We found cohorts that prove a seasonal periodicity in animal acquisition due either to the Argali behaviour or to Homo erectus movements. Determination of sex was possible on some bones. Pelvis bones give a 0.11 ratio (female/male). These proportions do not correspond to a natural ratio but to a herd of females with juvenile individuals during spring or summer. Our Argali population is composed of 27% juvenile, 64% adult females, and 9% adult males. Weight was estimated by measuring the first lower molar area or the talus area. Our measurements gave an average weight of 130 kg. Some pathologies were observed on teeth. Type and frequency allow to conclude on the good sanitary state of the fossil population. All these methods provide a first overview of this 440,000 year old population.
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Published abstracts

2010
Florent Rivals, Gina M Semprebon, Nikos Solounias (2010)  Advances in ungulate dental wear techniques reveal new patterns of niche breadth and expansion throughout the Cenozoic   Journal of Vertebrate Paleontology, SVP Program and Abstracts Book, 2010: 151-152A [Published abstracts]  
Abstract: In the last few decades, dietary ecological reconstructions have been used as powerful tools in gaining insight into local and global environmental trends. Ungulate mesowear and microwear studies in particular serve as useful proxies for demonstrating the existence of geographical and/or temporal variability in diet and vegetation structure. Improvements in these techniques reveal patterns involving Cenozoic vegetation and climatic structure as well as aspects of mammalian niche utilization. Results here indicate a congruity of responses to climate and aridity shifts among different ungulates and a much greater plasticity in dietary behavior among forms with similar tooth crown heights and morphology than previously supposed. Different diets are observed by the same taxon from different localities and brachydont forms are shown to engage in grazing while hypsodont forms are shown to engage in leaf or even fruit browsing. Specific examples from five ungulate families (Antilocapridae, Equidae, Camelidae, Dromomerycidae, and Merycoidodontidae) are used to illustrate these trends. These results are intriguing as increased hypsodonty acquisition through time is generally explained as a mechanism for niche displacement (i.e., browsing forms become grazing forms when displaced from their relatively closed habitats to relatively open ones) such as might have been the case in the evolution of more hypsodont ungulates from brachydont forms. Results here suggest that this classical explanation for increased crown height is too simplistic and not necessarily correct. Increased crown height is considered here as most likely serving the purpose of expanding the dietary breadth of a taxon. That is, crown height augmentation may serve as a mechanism to allow a species to exploit new habitats and to expand its niche but not necessarily to shift exclusively to a new dietary regime.
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2009
Florent Rivals, Dick Mol, John de Vos (2009)  Thirty meters under sea level: Dietary traits and paleoecology of woolly mammoths, rhinos, and other late Pleistocene ungulates fished in the North Sea   Journal of Vertebrate Paleontology, SVP Program and Abstracts Book, 2009: 172A [Published abstracts]  
Abstract: The glacial/interglacial climatic cycles of the Pleistocene resulted in significant variations in sea level. During glacial periods sea levels fell dramatically. The southern North Sea became a dry-land area with both rivers and lakes. Hundreds of thousands of fossil bones of Pleistocene mammals, both terrestrial and marine, have been fished from the bottom of the North Sea between Great Britain and The Netherlands. AMS radiocarbon dates were obtained from the Brown Bank locality and range from ca. 45,000 to 34,000 BP. We examined dietary traits of the late Pleistocene ungulates with dental wear patterns (mesowear and microwear) that relate to diet abrasion. Eight species were analyzed: Mammuthus primigenius, Coelodonta antiquitatis, Equus ferus, Megaloceros giganteus, Alces alces, Cervus elaphus, Rangifer tarandus, Bos primigenius, and Bison priscus. Considering mesowear, all cervids fall among the ungulates with low abrasive diets. For the bovids, mesowear signals indicate dietary habits with relatively high levels of abrasion. However, Bos displays significantly lower abrasion in its diet than Bison. Equus is observed among the modern grazers with a very high level of abrasion. Considering microwear, dietary traits form a continuum from browsing to grazing. In the low abrasion range (grazers), we observe Megaloceros, Cervus, Rangifer, Bos, and Coelodonta. Mammuthus is found among the mixed feeders, and finally Equus and Bison among the grazers (high abrasion). All these taxa have relatively low numbers of pits. The continuum of dietary traits indicates the presence of the wide diversity in the vegetal resources available and used by the ungulates. Diet thus reflects the presence of a dominant steppe-tundra with some deciduous trees or shrubs. The fauna could reflect the transition stadial/interstadial during the Pleniglacial. Alces, however, has a diet significantly different from the other species and falls among the dirty browsers which might be explained by (1) the use of a different resource (resource partitioning because of high competition) or (2) a different chronology (beginning of an interstadial).
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2008
Florent Rivals, Ellen Schulz, Thomas M Kaiser (2008)  Ungulate dietary adaptations and diversity in the Middle and Late Pleistocene of Western Europe: Variation through time and space   Journal of Vertebrate Paleontology 28(3): 132A [Published abstracts]  
Abstract: Dental wear patterns of Pleistocene ungulates (equids, cervids, and bovids) were analyzed as a paleoenvironmental and paleoclimatic proxy in relation to climatic changes and habitat diversity. The mesowear and microwear analysis includes 763 specimens from seven localities. The material belongs to nine archaeological levels corresponding to Neanderthal activities in Western Europe, and four different marine isotopic stages (MIS) 8 to MIS 3. The sites are located in two main geographic areas in Western Europe. The three German localities, Salzgitter Lebenstedt, Taubach, and Wallertheim are located in the Northern part of Western Europe and correspond to open-air sites. The localities Payre, Portel-Ouest, Ermitons, and Abric Romaní are caves located in the southern, Mediterranean area of France and Spain. Those sites were selected to analyze changes through time and space. (1) We compared changes through time during two glacial and two interglacial phases. Interglacial phases (MIS 5 and 3) are characterized by a high diversity in dietary adaptations of the ungulate fauna, whereas during glacial periods (MIS 8 and 4) the dietary diversity decreased significantly. The variations observed through time are certainly related to a reduction in diversity of the available vegetation during the cold and dry periods of the Pleistocene. (2) We compared geographical changes in dietary adaptations during interglacial periods, when North-Western Europe was not covered by ice sheets and Neanderthals colonized the area. During MIS 5 and 3, there is a significant increase in abrasiveness of the diet from North-Western to Southern Europe, corresponding to an increase towards more grazing. This corresponds to a latitudinal gradient in vegetation from an evergreen forest in North-Western Europe to temperate grassland in the Mediterranean area evidenced by pollen data. Dental wear analyzes reveals significant changes on both temporal and geographical scales in relation to climate and vegetation changes.
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2007
Gina M Semprebon, Florent Rivals (2007)  Hypsodont browsing: Camel and pronghorn dietary reconstruction through deep time   Journal of Vertebrate Paleontology 27: 144A [Published abstracts]  
Abstract: Dietary adaptations of both Tertiary and Quaternary representatives of North American Antilocapridae and Camelidae were examined through deep evolutionary time (via hypsodonty index), though ecological time (via mesowear analysis), and through the last few days of life (microwear). Fossil samples were from the Great Plains, Great Basin, and Florida and span from the early Miocene to the Recent. Mesowear patterns are strikingly similar in antilocaprids and camels and demonstrate a clear shift toward more abrasive diets beginning in the late Hemphillian (Late Miocene) and then a return to a less abrasive dietary regime beginning in the Pliocene and continuing into the recent. However, microwear patterns are strikingly dissimilar in antiliocaprids versus camelids. In addition, hypsodonty indices fall short in terms of accurately predicting most probable modern and paleodiet. This study underscores the need to use an eclectic dietary approach to assess ancient diets and offers new insight into the origin of hypsodonty in these two groups.
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2006
Agnès Testu, Florent Rivals (2006)  Taphonomy and paleobiology of carnivores and argali from a Middle Pleistocene assemblage (Caune de l'Arago, Tautavel, France)   Journal of Vertebrate Paleontology 26: 131A [Published abstracts]  
Abstract: Observation of taphonomic details preserved on bone assemblages gives access to the paleobiological information lost during fossilization and of the paleoethology of the bone collector. An argali (Ovis ammon antiqua) assemblage from the Middle Pleistocene cave of the Caune de l'Arago (Tautavel, southern France) is studied in terms of taphonomy. The level analyzed is 550,000 years old. Previous observations leave no doubt that carnivores are responsible for this accumulation and non-selectively transported argali carcasses into the cave. The type of bones in articulation and the gnawing marks are characteristic of carnivores. Analytical study of this accumulation reveals hunting rather than scavenging as procurement mode of carcasses. This is consistent with the association of carnivores from these levels, where the hyena is absent, represented by ursids, and strict carnivores such as felids (lion cave, leopard with uncia-like features) and canids (wolf), whose body size and food habits are compatible with argali as game. The hypothesis tested consists in defining the species involved in this process. A similar association can be observed today in the Himalayan Mountains. In addition, the topography of the site (plateau, cliffs), as well as the climatic conditions (cold and dry) contemporaneous to the deposit of these levels favored such an association. This unique assemblage provides relevant elements for interspecific competition between carnivores, in southern Europe, during the lower middle Pleistocene.
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Florent Rivals, Agnès Testu (2006)  Local evolution of Late Pleistocene Caprinae and Hyaenidae in the northeastern part of the Pyrenean Mountains (France)   Journal of Vertebrate Paleontology 26: 115A-116A [Published abstracts]  
Abstract: The Pyrenean Mountains are characterized by high levels of endemism in modern faunas and floras (e.g. Pyrenean desman or water-mole, Galemys pyrenaicus; Pyrenean salamander, Euproctus asper; or Pyrenean rock lizard, Lacerta bonnali). This endemism was to be the result of glaciations which geographically isolated some areas of southern Europe during Pleistocene. We tested this hypothesis by studying the evolution of Caprinae and Hyaenidae found in three late Pleistocene localities of the northeastern Pyrenean Mountains (France), dated from 90,000 to 60,000 years B.P. â Caune de l'Arago, Arche cave, and Portel-Ouest cave. The data obtained on populations of tahr, (Hemitragus cedrensis), ibex (Capra caucasica), and hyena (Crocuta crocuta spelaea) were compared with those of the French Massif Central populations (Bouxès, Hortus, Baume Moula-Guercy, Saint-Marcel-d'Ardèche) and Provence ones (Adaouste, Bau de l'Aubesier). This study is mainly based upon dental and postcranial characters and measurements. Typical dental features were found on the northeastern Pyrenees species only, for Caprinae and hyena as well. Ibex, for instance, shows a reduced metastylar wing on the M3, narrow parastyle and metastyle with low relief. The two species of Caprinae present also a reduction in size of all anatomical parts, especially on metatarsal and phalanx. Populations of the Massif Central and Provence do not display such a reduction. Pyrenean populations of hyena show a narrow P4 in its anterior part and a p4 with a short protoconid. All these characters suggest a local evolution of Caprinae and Hyaenidae during the late Pleistocene, supporting the hypothesis that the northeastern part of the Pyrenees was geographically isolated from the Iberian Peninsula and the rest of continental Europe during the coldest periods of the late Pleistocene.
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Florent Rivals, Nikos Solounias, G B Schaller (2006)  Similarities in the diets of Mongolian gazelles, Tibetan antelopes and ungulates from arctic/subartic habitats using mesowear and microwear analyses   Mammalian Biology 71 suppl: 24 [Published abstracts]  
Abstract: The diet of Mongolian gazelles and Tibetan antelopes is subject to discussion. Mongolian gazelles (Procapra gutturosa) are classified either as grazers or mixed feeders, whereas Tibetan antelopes are known to be browsers or mixed feeders. The diet of these two ungulates is evaluated in this study by dental microwear (i.e., immediate wear) and mesowear (i.e., cumulative wear), and compared the results with ungulates steppe habitats and arctic/subartic habitats. The high pit densities observed for both Pantholops and Procapra can be related to the presence of grit: grit encroachment on the vegetation and grit or soil ingested because these animals are feeding on short vegetation and can ingest some soil with it. Multivariate analysis, using both microwear and mesowear data, shows a good discrimination between grazers and browsers. The samples we analyzed plotted outside of the area occupied by grazers and browsers and fall into intermediate areas occupied by mixed feeders. The analysis reveals that both species plot close to the musk-ox (Ovibos moschatus) and the caribou (Rangifer tarandus). The diets of these four species have similar properties in terms of abrasiveness because they give very similar mesowear and microwear signatures. These similarities are also identified by biologists who suggested that the arctic or subarctic regions inhabited by musk-ox and caribou are comparable to the landscapes used by Pantholops and Procapra. The diet of caribou and musk-ox appears to be highly variable on regional or seasonal basis. Similar pattern were observed for Mongolian gazelles with significant changes related to seasonal modifications in food availability. Both ecological observations and tooth wear analysis reflect this seasonal variability. Because of the similarities between Pantholops, Procapra and two artic/sub-artic mammals we can suggest that the two species we analyzed are mixed feeders, their diet varying regionally and/or seasonally.
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2005
Florent Rivals, Gina Semprebon (2005)  Dental wear and dietary evolution of Antilocapridae during the Miocene-Pleistocene climatic transformation   Journal of Vertebrate Paleontology 25: 105A [Published abstracts]  
Abstract: </i>Dietary adaptations of both Tertiary and Quaternary representatives of the family Antilocapridae were examined using both enamel microwear and mesowear analyses of molar teeth. Taxa assessed span from the early Miocene (late Hemingfordian) to the late Pleistocene and include samples from the Central and Southern Great Plains and the Northern and Southern Great Basin. Results are compared to those of the modern pronghorn (<i>Antilocapra americana</i>) and depict a surprising array of dietary behavior given the fairly stereotypic gross morphology of molar teeth found throughout the family (hypsodonty). Both microwear and mesowear results show a shift toward more abrasive diets beginning in the Hemphillian (late Miocene-Pliocene). Results are concordant with well-known trends toward increasing aridity and shifts in vegetational structure in the late Miocene-Early Pliocene of North America and demonstrate that fossil pronghorn were sensitive to changes in the abundance and quality of their food resources. A return to a less abrasive dietary regime is noted for the duration of the Pleistocene. The less derived merycodontines apparently depended less on grass than the more advanced antilocaprines (more hypsodont, longer limbs), yet even the earliest merycodontines studied apparently relied more on grass as a dietary staple than the modern pronghorn, underscoring the importance such dietary analyses hold for overturning uniformitarian assumptions based upon well-known Holocene forms that might not prove to be good analogs for fossil forms and providing glimpses into a possible reason for the extreme hypsodonty present in the modern pronghorn despite its mainly browsing dietary behavior.
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2004
Nikos Solounias, Matthew C Mihlbachler, Florent Rivals, Cécile Blondel, Dale R Guthrie (2004)  Evaluation of diet in mammoths and other Late Pleistocene and Early Holocene ungulates from Alaska   Journal of Vertebrate Paleontology 24: 116A [Published abstracts]  
Abstract: The dietary ecology of species across the Pleistocene-Holocene boundary has the potential to reveal ecological trends associated with the end of the last ice age and/or reveal interspecies niche dynamics that are dissimilar to today due to associations with extinct species or unique paleoenvironments. We examined diet of late Pleistocene and early Holocene ungulates from Alaska with dental wear patterns that relate to diet abrasion. Browsers have low abrasion diets while grazers have high abrasion diets. Characterizing labial molar wear facets as sharp, rounded, or blunt (mesowear) characterizes dietary abrasion over a long period of ecological time (years, months), whereas the relative number of pits and scratches on occlusal surfaces (microwear) characterizes diet over a shorter amount of time (days, weeks). Dental wear patterns of late Pleistocene species, Mammuthus, Equus, Bison, and Rangifer, consistently suggest a surprisingly homogenous diet with levels of abrasion reflecting a significant grassy component. Dental wear patterns of early Holocene ungulates, Bison ,Rangifer , and Cervus , indicate similar levels of abrasion. However, early Holocene Alces diet was significantly less abrasive, suggesting that it was the sole browser. Bison and Equus, typical grazers, tend to have finer scratches and a higher proportion of rounded cusps than modern reference samples, suggesting a somewhat less abrasive diet for these species than is typical of today. In contrast, the three deer species, Rangifer, Cervus, and Alces, typical mixed feeders and browsers, show dental wear patterns that reflect somewhat more abrasive diets than that found in modern reference samples. However, among the deer, the relative diet pattern was the same as the modern samples, with Rangifer having the most abrasive diet, Alces having the least abrasive diet, and Cervus with an intermediate diet.
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Florent Rivals, Nikos Solounias (2004)  Bison dietary changes in North America during the Pleistocene-Holocene transition   Journal of Vertebrate Paleontology 24: 104A [Published abstracts]  
Abstract: Bison is a common fossil taxon in North America. Bison inhabited large regions of North America south of the margin of the Pleistocene ice sheets. Because of the widespread distribution of Bison, some differences should exist during the Pleistocene from the northern to the southern habitats. Four samples of Bison from Pleistocene localities were studied: Fairbanks Area (Alaska), Folsom Quarry (New Mexico), Seminole Field (Florida) and Dalhart Sideroad Pit (Texas). They in turn were compared to a mixed sample of extant Bison from the central plains. A dietary study was performed using tooth mesowear and microwear of the upper second molars. Mesowear: All extinct samples had mainly rounded molar apices (between 93 and 100%). This mesowear signature is strongly different from that of modem Bison where it is 26.6% rounded and 73.3% blunt and most similar to Hippotragus equinus and Kobus ellipsiprymnus (African bush and water margin grazers). The microwear analysis shows that Bison from New Mexico, and Texas were grazers similar to Equus burchelli. Bison from Florida was similar to Gazella granti (a mixed feeder) or fruit dominated browsers. The extinct Alaska Bison had a diet similar to recent Cewus canadensis (a mixed feeder). Elk graze part of the time as mixed feeders. Two additional observations are possible. (1) Because of the high longitude of Alaska and the Pleistocene temperatures, the fine scratches observed on this sample suggest this extinct Bison population was grazing on C3 grasses. (2) Such C3 grasses produce very fine microwear scratches. The results show geographically variable diets for the extinct Bison. This study reveals the importance of such analyses which are necessary to understand the dietary breadth of species and escape uniformitarian assumptions based upon the well-known but in certain cases unrealistic Holocene scenarios.
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Masters theses

1999
Florent Rivals (1999)  Les petits bovidés pléistocènes de la Caune de l'Arago (Tautavel, Pyrénées-Orientales). Etude paléontologique, taphonomique, archéozoologique et paléoécologique   Université de Perpignan (France)  
Abstract: The Lower Palaeolithic site of Caune de l'Arago (Tautavel, France), presents a remarkable diversity and abundance of small bovids (Ovis ammon antiqua, Hemitragus bonali, Rupicapra sp., and Capra ibex). Spatial distribution study of remains had allowed to subdivide the G level in three sub-levels. Seasonality study show that bovids were hunted and brought to the cave from late summer to late fall. Wild sheep (O. antiqua) are abundant in the F level, where they represent 74 % of hunted animals (56 young and 150 adults). Tahr (H. bonali) is well represented at the base of the G level. Mortality patterns confirm the hunting activity of the Tautavel hominids. Wild sheep were hunted whatever their age, whereas the tahr show a selective hunting pattern with young animals being very rare or absent.
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PhD theses

2002
Florent Rivals (2002)  Les petits bovidés pléistocènes dans le bassin méditerranéen et le Caucase. Etude paléontologique, biostratigraphique, archéozoologique et paléoécologique   Université de Perpignan (France)  
Abstract: In the Middle and Upper Pleistocene sites studied in the Mediterranean area and the Caucasus, small bovids are represented by 4 genera (Ovis, Hemitragus, Capra and Rupicapra). Ovis ammon antiqua and Capra aff. caucasica are similar to those known in other European and Caucasian populations. Two species of thar are present at the Caune de l'Arago, namely Hemitragus bonali in the Middle Pleistocene levels and Hemitragus cedrensis in the Upper Pleistocene level. Rupicapra from the Caune de l'Arago is the oldest sample of the genera in Europe. This taxa would be phylogenetically linked to Rupicapra pyrenaica. A zooarchaeological approach was applied to the levels of units I and III of the Caune de l'Arago. The levels M, N and O from unit I correspond to carnivore occupations while in most of the levels of unit III, argali accumulation is due to human occupation. In level F dated from about 440,000 years, the cavity was occupied from the end of spring to the beginning of summer by groups which practised non selective hunting and argali was the main prey. The study of locomotive and dietary adaptations shows that fossil populations had very comparable adaptations to those of current populations. Dental microwear shows the seasonal changes in food selection which varied with the hunting season of small bovids and with palaeoenvironmental conditions.
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Book

2004
Florent Rivals (2004)  Les petits bovidés (Caprini et Rupicaprini) pléistocènes dans le bassin méditerranéen et le Caucase. Etude paléontologique, biostratigraphique, archéozoologique et paléoécologique   [Book]  
Abstract: A detailed study of the exploitation of small bovine species based on faunal assemblages from eight sites in the east Pyrenees, the Languedoc region of France, southern Turkey and the Caucasus. Four main species are identified and analysed, with comparisons drawn between the sites in the study, and with a view to investigating the subsistence behaviour of people living in these areas during the Pleistocene. The effects of changing environmental conditions on species levels and human behaviour is also covered.
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Online papers

2005
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