No full-text available
To read the full-text of this research,
you can request a copy directly from the authors.
Solounias, N. and B. Dawson-Saunder. Dietary adaptations and paleoecology of the late Miocene ruminants from Pikermi and Samos in Greece. Paleogeogr. Paleoclimatol. Paleoecol
Abstract
The late Miocene (Turolian) localities of Pikermi and Samos in Greece present numerous ruminant species which have been interpreted to represent savanna-adapted taxa. The masticatory morphology of these extinct ruminants is compared qualitatively to twenty-seven extant ruminant species from Asia and Africa using thirteen masticatory features. Statistical analysis of these features on the modern species shows significant differences between the morphology of browsers, intermediates and grazers and suggests that intermediate and grazing adaptations evolved independently more than once. In addition, certain extant ruminant browsers and intermediates may have assumed such diets from ancestral grazing species recently. Application of this method to the extinct species from Pikermi and Samos shows that, with the exception of one species, the bovids were either browsers or intermediates. Parurmiatherium rugosifrons from Samos was probably the only grazer. Pachytragus laticeps, a presumably advanced species, has masticatory morphology very similar to Miotragocerus species, commonly interpreted as primitive species. Samotherium boissieri and Helladotherium duvernoyi were intermediate feeders, although giraffids. We propose that the absence of grazers at these localities suggests that savanna-like dietary adaptations for ruminants were not as widespread as now. The presence of savannas similar to those of East Africa at Pikermi, Samos and other Miocene localities is unlikely. Such savannas were originally proposed because of modernization of fossil taxa, the presence of numerous ungulate and hyaenid species, the misconception that hipparions represent zebra-like species, and the abundance of fossil specimens. Ruminant faunal comparisons using masticatory morphology point out a greater similarity of Pikermi and Samos to extant forest and woodland ruminants; the extinct ruminant faunas were not similar to extant savanna-adapted species. A good physiognomic model for the Pikermi-Samos ruminants is the forest-woodland ruminants of Sichuan, China, Southeast Asia, and Kanha, India. The hypothesis that the Pikermi ruminants evolved in forested environments is supported by analysis of the local paleofloras. The Pikermi is similar to modern warm mixed riparian evergreen and broadleaf deciduous temperate woodland-forest. The Turolian mountains near Pikermi had mixed forests, although evergreens dominated.
... Each category may be further divided into 2-3 subcategories based on detailed forage selectivity and preference. These categories and subcategories, with slight modifications in terminology and definition, have been widely used to classify the feeding habits of living as well as fossil ruminants (e.g., Gordon and Illius 1988;Janis and Ehrhardt 1988;Langer 1988;Solounias and Dawson-Saunders 1988;Bodmer 1990;Solounias and Moelleken 1993b;Spencer 1995;Dompierre and Churcher 1996;Sponheimer et al. 1999;Pérez-Barbería and Gordon 2001;Clauss et al. 2008;Fraser and Theodor 2011a;Forrest et al. 2018;Supplementary Table S1). A number of studies have alternatively used the percentage of grass in species' diets to quantitatively (as opposed to categorically) place species along the browser-grazer spectrum (e.g., Clauss et al. 2003;Pérez-Barbería et al. 2004;Kaiser et al. 2011;Codron et al. 2019). ...
... In contrast, the craniomandibular adaptations of grazers are primarily associated with processing food. Grasses occur in greater density than leaves and fruits and can be taken in larger amounts per bite (Solounias and Dawson-Saunders 1988). Grazers generally have wider and flatter muzzles, with wider and more protruding incisors, than browsers (Janis and Ehrhardt 1988;Pérez-Barbería and Gordon 2001). ...
... Hypsodonty is particularly associated with the second and third molars, resulting in a deeper mandibular body under those teeth. Taller teeth also bring occlusal surfaces closer to the condyle, which serves as the fulcrum of the chewing muscular apparatus (Solounias and Dawson-Saunders 1988;Pérez-Barbería and Gordon 1999). Hypsodonty, however, does not always signify increased grass consumption; high wear resistance is also an adaptation for consuming more dust and grit in the diet (Janis 1988;MacFadden et al. 1999;Solounias and Semprebon 2002;Strö mberg 2002;Damuth and Janis 2011). ...
The mammalian family Bovidae has been widely studied in ecomorphological research, with important applications to paleoecological and paleohabitat reconstructions. Most studies of bovid craniomandibular features in relation to diet have used linear measurements. In this study, we conduct landmark-based geometric-morphometric analyses to evaluate whether different dietary groups can be distinguished by mandibular morphology. Our analysis includes data for 100 species of extant bovids, covering all bovid tribes and two dietary classifications. For the first classification with three feeding categories, we found that browsers (including frugivores), mixed feeders, and grazers are moderately well separated using mandibular shape. A finer dietary classification (frugivore, browser, browser-grazer intermediate, generalist, variable grazer, obligate grazer) proved to be more useful for differentiating dietary extremes (frugivores and obligate grazers) but performed equally or less well for other groups. Notably, frugivorous bovids, which belong in tribe Cephalophini, have a distinct mandibular shape that is readily distinguished from all other dietary groups, yielding a 100% correct classification rate from jackknife cross-validation. The main differences in mandibular shape found among dietary groups are related to the functional needs of species during forage prehension and mastication. Compared to browsers, both frugivores and grazers have mandibles that are adapted for higher biomechanical demand of chewing. Additionally, frugivore mandibles are adapted for selective cropping. Our results call for more work on the feeding ecology and functional morphology of frugivores and offer an approach for reconstructing the diet of extinct bovids.
... Previous publications have demonstrated morphological differences in cranio-dental morphology between bovids with different feeding preferences (e.g., Janis, 1988;Solounias and Dawson-Saunders, 1988;Spencer, 1995aSpencer, ,b, 1997Reed, 1996). In general, browsers eat mainly soft leafy tree and shrub foliage and fruits, while grazers eat predominantly fresh grass and roughage. ...
... Grazing species tend to chew their food more posteriorly and have reduced molar and premolar rows compared to browsers. This brings the food closer to the fulcrum of the mandible and increases the amount of occlusal pressure applied to the plant material during mastication (Solounias and Dawson-Saunders, 1988). ...
... In grazing taxa, we expect to see more convexity on the posterior and ventral borders of the mandibular ramus. In contrast, the posterior and ventral borders of the mandibular ramus are expected to appear concave in browsers (Solounias and Dawson-Saunders, 1988). ...
... The genus Protragelaphus is known from the Late Miocene of Pikermi (Greece), Grebeniki (Ukraine) and Taraclia (Moldova) (Sokolov 1953), thus also suggesting the Ponto-Mediterranean origin of Gazellospira. The paleoenvironments of Protragelaphus from Samos and Pikermi, according to Solounias and Dawson-Saunders (1988), were dominated by mixed non-canopy forming woodlands and forest with a minor sclerophyllous component. The palaeobotanic data are supported by the functional morphology of bovids from Samos and Pikermi, which indicates an adaptation to browsing and mixed feeding (Solounias and Dawson-Saunders 1988;Solounias and Moelleken 1999). ...
... The paleoenvironments of Protragelaphus from Samos and Pikermi, according to Solounias and Dawson-Saunders (1988), were dominated by mixed non-canopy forming woodlands and forest with a minor sclerophyllous component. The palaeobotanic data are supported by the functional morphology of bovids from Samos and Pikermi, which indicates an adaptation to browsing and mixed feeding (Solounias and Dawson-Saunders 1988;Solounias and Moelleken 1999). The further evolution of Gazellospira could be linked to the forested humid biotopes from the Carpathian area and Balkans reported by Eronen and Rook (2004) and Fortelius et al. (2006). ...
The palaeontological material from the Dacian Basin provides a complete and well-represented record of faunal succession during the important faunal turnover called the Pachycrocuta event. The present study describes fossil remains of ruminants (Cervidae, Bovidae, Giraffidae) from the Early Pleistocene of Valea Grăunceanului, Fântâna lui Mitilan and other smaller sites from the Olteț River Valley. The article discuss the taxonomic context, eco-morphology and functional morphology of the described taxa and dynamics of biogeographic distribution. The commutity of ruminant species from the Dacian Basin before the Pachycrocuta event is dominated by Pliocene holdovers: Pliotragus ardeus, Gazellospira torticornis, Rucervus radulescui, Metacervocerus rhenanus and Mitilanotherium inexspectatum. This assemblage also contains new forms for Western Eurasia as Dama eurygonos, Eucladoceros dicranios, Eucladoceros ctenoides, Alces sp. and Bison (Eobison) sp. This specific regional fauna of ruminants became extinct during the Pachycrocuta faunal turnover and was replaced by a more cold-adapted assemblage of ruminants (Megalovis latifrons, Eucladoceros sp., Dama sp., Praemegaceros obscurus and its specialised diminished form Praemegaceros cf. mosbachensis) that show a greater affinity with coeval Levant faunas. The revealed dynamics of paleobiogeographic zones from the Early Pleistocene of southeastern Europe supports the hypothesis of early hominin dispersals in Western Eurasia via Balkan-Anatolia path.
... However, we cannot know whether canines were lacking in the species (e. g., as in modern fallow deer) or were visible only in males (e. g., as in modern red deer). The facial portion can give information on the feeding habits of ruminants (Solounias and Dawson-Saunders 1988;Solounias and Moelleken 1993;Caloi and Palombo 1995;Janis 1995;Palombo 2005): -The portion of the maxillaries anterior to the mesial edge of P 2 , dorsally delimitated by the nasals and anteriorly by the praemaxillaries, is more elongated in modern deer species that feed mainly on leaves, so that its general outline is a trapezium, while it is more shortened, and thus more triangular, in mixed feeders and grazers (Solounias and Dawson-Saunders 1988;). The only specimen from Untermassfeld in which this area is present is female skull IQW 1992/23 910 (Mei. ...
... However, we cannot know whether canines were lacking in the species (e. g., as in modern fallow deer) or were visible only in males (e. g., as in modern red deer). The facial portion can give information on the feeding habits of ruminants (Solounias and Dawson-Saunders 1988;Solounias and Moelleken 1993;Caloi and Palombo 1995;Janis 1995;Palombo 2005): -The portion of the maxillaries anterior to the mesial edge of P 2 , dorsally delimitated by the nasals and anteriorly by the praemaxillaries, is more elongated in modern deer species that feed mainly on leaves, so that its general outline is a trapezium, while it is more shortened, and thus more triangular, in mixed feeders and grazers (Solounias and Dawson-Saunders 1988;). The only specimen from Untermassfeld in which this area is present is female skull IQW 1992/23 910 (Mei. ...
The deer remains from the Early Pleistocene site of Untermassfeld (Thuringia, Germany) are analysed in detail, including published and previously unpublished material. The analysis confirms the presence of four species previously recorded at the site: the roe deer Capreolus cusanoides, the moose Cervalces carnutorum, the fallow deer ›Pseudodama‹ vallonnetensis and the large comb-antlered deer Eucladoceros giulii.
The remains of roe deer and moose are extremely rare in general, so each new specimen has been described in detail and measurements have been provided. Their importance rests in Untermassfeld being the type locality for C. cusanoides and the earliest European occurrence of the genus, and for C. carnutorum providing the richest collection of this otherwise poorly recorded species. E. giulii and ›P.‹ vallonnetensis are very abundant at the site, thus only selected antler, cranial and dental remains have been described in detail.
In particular, their completeness and good state of preservation allows a detailed morphological analysis which, by comparison with modern deer taxa of known ecology, allows the reconstruction of their likely feeding habits. The Untermassfeld sample of E. giulii represents the species’ type-population and last known representative of the Eucladoceros lineage. ›P.‹ vallonnetensis is the richest published collection of this last
representative of the ›Pseudodama‹ lineage which likely includes the ancestor of the modern genus Dama.
Updated MNIs haves been calculated for the four species recorded.
... Over the last four decades, scientists have established many proxies for the ecology and evolution of diet in mammals. The most important ones are hypsodonty (Fortelius, 1985;Janis, 1988), low magnification microwear analysis (Solounias and Semprebon, 2002), mesowear analysis (Fortelius and Solounias, 2000;Kaiser and Solounias, 2003;Schubert, 2007), morphology of the masticatory apparatus (Solounias and Dawson-Saunders, 1988), morphology of the masseter muscle (Solounias et al., 1995), shape analysis of the ruminant premaxilla Solounias and Moelleken, 1993a;Solounias and Moelleken, 1993b), tooth wear gradients (Kaiser and Schulz, 2006;Kubo and Yamada, 2014), cranial foramina (Solounias and Moelleken, 1999) and dental allometry (Ungar, 2014). Dental microwear has proven to be the best-studied palaeodiet proxy so far. ...
... The artiodactyls used in this study include browsers (Giraffa camelopardalis, Litocranius walleri), grazers (Bos gaurus, Connochaetes taurinus, Kobus ellipsiprymnus), mixed feeders (Muntiacus muntjak, Rangifer tarandus), and one frugivore taxon (Tragulus javanicus). The feeding ecology of extinct and extant artiodactyls was investigated by the use of one or more of the above mentioned proxies Solounias and Dawson-Saunders, 1988;Moelleken, 1993a, 1993b;Solounias et al., 1995;Solounias et al., 2000;Solounias and Semprebon, 2002;Semprebon et al., 2004;Merceron et al., 2004;Merceron et al., 2005b;Franz-Odendaal and Solounias, 2004;Solounias et al., 2010;Harris et al., 2010;Fraser and Theodor, 2011;Tariq and Jahan, 2014). ...
Low magnification dental microwear constitutes one of the most important proxies on the ecology and evolution of diet in mammals. Numerous studies have been established on the reconstruction of dietary ecology of even-toed and carnivorous taxa. To date, these studies have used the second permanent molars or carnassials exclusively, for ungulates or carnivores, respectively. In this study, for the first time, premolars and non-carnassials of artiodactyls and carnivores, respectively, were used. Tooth samples from nine artiodactyl and eight carnivore taxa, covering the largest dietary spectrum possible, were evaluated and the microwear signal of the premolars/non-carnassials compared with known data of the second permanent molars and carnassials. The results reveal an almost identical statistical microwear signal of the premolars and molars for artiodactyls and the non-carnassials and carnassials for carnivores. Based on the even-toed taxa, new expanded morphospaces for both browsers and grazers are depicted on an average scratches versus average pits scatterplot. Additionally, dietary tendencies were delineated for the carnivore taxa with low or high counts of small and large pits for specific preferences such as a meat/bone, frugivorous or mixed carnivore diets in non-carnassial tooth positions as well. Lastly, low scratch and low large pit score plots for the ungulate and carnivore taxa, respectively, reveal an even clearer separation between the main dietary categories.
... La caracterización trófica de los mamíferos herbívoros más exhaustiva y de índole cuantitativa ha sido realizada fundamentalmente por Christine Janis , 1989, 1990a, 1990bJanis y Ehrhardt, 1988;Janis y Fortelius, 1988;Janis y Constable, 1993;Janis et al., 1995;Mendoza et al., 2002) y Nikos Solounias Solounias y Dawson-Saunders, 1988;Solounias y Hayek, 1993;Solounias y Moelleken, 1992a, 1992b con numerosas aplicaciones a especies fósiles y una abundante bibliografía. No obstante, otros investigadores también han analizado o destacado algunas diferencias morfológicas entre los distintos grupos tróficos de ungulados, especialmente pacedores frente a ramoneadores (Estes, 1974;Gosling, 1974;Sinclair y Gwynne, 1972;Vrba, 1980;Gordon e Illius, 1988;Bodmer, 1990;Spencer, 1995;Pérez-Barbería y Gordon, 1999). ...
... Caracterización multivariante. Janis y Solounias trataron finalmente de caracterizar distintas adaptaciones tróficas de los ungulados partiendo de un gran número de variables morfológicas (Solounias y Dawson-Saunders, 1988;Janis, 1990aJanis, , 1995Solounias y Moelleken, 1992b, pero siempre con un enfoque sencillo y univariante, en el que a lo sumo se utilizaban índices basados en dos variables, generalmente para relativizar las variables afectadas por una adaptación trófica frente al tamaño corporal. Janis (1990a), por ejemplo, efectuó regresiones entre la masa corporal y 24 variables craneodentales en 136 especies de ungulados y 52 especies de macropódidos, analizando la distribución de cada grupo trófico respecto a la recta de regresión mediante el test T de comparación de Wilcoxon. ...
... A number of aspects of craniodental morphology can be shown to correlate with dietary behavior (see Solounias and Dawson-Saunders 1988;Janis 1995;Mendoza et al. 2002). The features distinguishing grazers from browsers relate to the different physical demands of feeding on grass versus browse. ...
... In the early late Miocene of Eurasia there was a reduction in numbers of browsers such as suids, tapirs, tragulids, and brachydont rhinos, suggesting a loss of forest habitat, and a radiation of bovids, equids, and more hypsodont rhinos and giraffoids, suggesting the spread of more grass-dominated habitats (Barry 1995;Fortelius et al. 1996;Agustí and Antón 2002;Costeur et al. 2004.). However, these faunas contained few undoubted grazing species, the hypsodont forms most likely being mixed feeders, and the habitat of these localities appears to have been woodland and shrubland rather than open savanna (Solounias and Dawson-Saunders 1988;Prins 1998). Few highly hypsodont bovids were present in Africa: the late Miocene fauna contained a large diversity of suids, rhinos, proboscideans, and giraffids, as well as bovids and hipparionine equids, probably representing an assemblage of browsers and mixed feeders (Turner and Antón, 2004). ...
... La caracterización trófica de los mamíferos herbívoros más exhaustiva y de índole cuantitativa ha sido realizada fundamentalmente por Christine Janis , 1989, 1990a, 1990bJanis y Ehrhardt, 1988;Janis y Fortelius, 1988;Janis y Constable, 1993;Janis et al., 1995;Mendoza et al., 2002) y Nikos Solounias Solounias y Dawson-Saunders, 1988;Solounias y Hayek, 1993;Solounias y Moelleken, 1992a, 1992b con numerosas aplicaciones a especies fósiles y una abundante bibliografía. No obstante, otros investigadores también han analizado o destacado algunas diferencias morfológicas entre los distintos grupos tróficos de ungulados, especialmente pacedores frente a ramoneadores (Estes, 1974;Gosling, 1974;Sinclair y Gwynne, 1972;Vrba, 1980;Gordon e Illius, 1988;Bodmer, 1990;Spencer, 1995;Pérez-Barbería y Gordon, 1999). ...
... Caracterización multivariante. Janis y Solounias trataron finalmente de caracterizar distintas adaptaciones tróficas de los ungulados partiendo de un gran número de variables morfológicas (Solounias y Dawson-Saunders, 1988;Janis, 1990aJanis, , 1995Solounias y Moelleken, 1992b, pero siempre con un enfoque sencillo y univariante, en el que a lo sumo se utilizaban índices basados en dos variables, generalmente para relativizar las variables afectadas por una adaptación trófica frente al tamaño corporal. Janis (1990a), por ejemplo, efectuó regresiones entre la masa corporal y 24 variables craneodentales en 136 especies de ungulados y 52 especies de macropódidos, analizando la distribución de cada grupo trófico respecto a la recta de regresión mediante el test T de comparación de Wilcoxon. ...
A primary goal of palaeoecology is the reconstruction of the adaptations and modes of life of extinct species, or palaeoautoecological characterization. The behavioral attributes of the fossil species have been inferred in different ways, but the comparative method is probably the most widely used. For this, some authors have tried to carry out a cuantitative characterization of different ecological adaptations in ungulates, using the dental-skeleton morphology of the living species, which is known as ecomorphological characterization. At first, that characterization was only qualitative. Later on, some authors began to quantify the relationship between morphology and different ecological adaptations, but this quantification simply entailed finding significant statistic correlations between some measurements and the adaptation that was analized. However, the more and more frequent use of the multivariate statistics has allowed a more complex, precise and reliable morphological characterization of the ecological adaptations of ungulates. In this review, I analize the evolution that the undestanding of ungulate ecomorphology has undergone during the last decades, with special emphasis on the methodology and results of some recent papers which are characteristic of a complex approach.
... Dental microwear and mesowear results from the abrasion of teeth by food items consumed during the last days prior to the death of the animal (Solounias and Dawson-Saunders, 1988;Teaford and Oyen, 1989;Fortelius and Solounias, 2000). Dental microwear study is a very useful tool for characterizing the diet of ancient mammals and therefore helping in reconstructing past environments. ...
... According to the properties of food and related items, the microwear may vary. For example, the dental microwear pattern on shearing molar facets of browsing ungulates differs from that of grazers by a higher percentage of pits (Solounias and Dawson-Saunders, 1988;Solounias and Semprebon, 2002). Grasses and related plants leave numerous scratches on the teeth partly because of the high concentration of silica phytoliths in their cell walls (MacNaughton et al., 1985;Robert and Roland, 1998). ...
a b s t r a c t Late Miocene climate change particularly monsoon intensification brought about by tectonic upheavals changed significantly the regional vegetation scenario of the Indian Subcontinent. Siwalik Middle Miocene closed forests gave way to Late Miocene open and seasonal forests, which in turn were gradually replaced by latest Miocene and Early Pliocene wooded grasslands. Here, new stable isotope and microwear data of a Late Miocene rhizomyid rodent, and dental microwear analyses of selected Late Miocene, and Plio-Pleistocene large and small herbivorous mammals from India is presented. These findings are then integrated with the earlier palaeodiet and palaeohabitat data of Siwalik mammals, in order to get an overall picture of climate vis a vis dietary and habitat change between ~14 and ~1 Ma. The results indicate that the Late Miocene rhizomyid Rhizomyides sivalensis was primarily a C 3 grazer, but might have also consumed leaves, tubers, roots, and seeds. The Miocene murine rodents Antemus chinjiensis and Karnimata cf. K. intermedia most likely ate fruits and insects. Among the Plio-Pleistocene murines Cremnomys and Mus cf. Mus flynni were mixed feeders of seeds, grass and insects. Bandicota was a seasonal grazer and mixed feeder, whereas Golunda was primarily a grazer with some intake of seeds and insects. A late Miocene rhinocerotid from Haritalyangar was a C 3 grazer, whereas its Plio-Pleistocene relative was a C 4 grazer. The hippo Hexaprotodon was a C 4 grazer. Plio-Pleistocene giraffid Sivatherium giganteum was a C 4 grazer, whereas its close relative Bramatherium was a mixed feeder with its main diet being C 4 grass. Camelus was also a mixed feeder with a dominant C 4 grass diet. Damalops was a mixed feeder with C 4 grass being its principal diet, whereas Hemibos was a C 4 grazer. Plio-Pleistocene ele-phantids Stegodon and Elephas were C 3 browsers and C 4 grazers, respectively. The Miocene arboreal frugivore/browser primates (sivapithecenes and sivaladapids) that lived in tropical forests were replaced by immigrant terrestrial grazers (baboons) in the Plio-Pleistocene, when the landscape was dominated by savannah type grasslands. On similar lines, Late Miocene monsoon intensification was responsible for the disappearance of several lineages of frugivore/browser suids, tragulids and bovids, which then were later replaced by omnivores/mixed feeders/grazer immigrants. Lineages of rhinos, equids, giraffids, anthracotheres and murids that had a C 3 diet in the Late Miocene started adapting to a C 4 diet. However, among the proboscideans the Miocene dinotheres (C 3 browsers) were replaced by the Plio-Pleistocene elephantids, which were both browsers and grazers.
... These range from dense forest to wide-open secondary grasslands to steep mountainous terrain. Thus, the value of bovids as habitat indicators has long been well-recognized (Gentry, 1970;Scott, 1979;Greenacre and Vrba, 1984;Kappelman, 1984Kappelman, , 1986Kappelman, , 1988Kappelman, , 1991Scott, 1985;Shipman and Harris, 1988;Solounias and Dawson-Saunders, 1988;Kappelman et al., 1997). As bovids are often common at fossil sites of relevance to hominin evolution, they are an important part of hominin paleohabitat reconstructions. ...
... Thus, the presence of taxonomic groups whose extant representatives tend to be associated with a particular habitat was viewed as evidence for that habitat's existence in the past. This approach is limited by the possibility that fossil representatives of extant taxa may well have used different habitats (see Solounias and Dawson-Saunders, 1988). More recently, habitat preferences have been reconstructed based on ecomorphology (e.g., Kappelman, 1988;Scott et al., 1999;Kovarovic et al., 2002;Vrba, 2003, 2005;Plummer et al., 2008;Bishop et al., 2011;Curran, 2012;Meloro et al., 2013). ...
... Nevertheless, P. inexspectatus is rather hypsodont for a giraffid (Geraads 1998, Kostopoulos andAthanassiou 2005), hence a more grazing diet than other giraffids would be expected. Moreover, the low p/m index is considered by many scholars as an ecomorphological adaptation of ruminants to a diet with a greater grazing component (Solounias and Dawson-Saunders 1988, Greaves 1991, Codron et al. 2019. Previous mesowear and microwear studies confirm that premolar/molar row ratios are, to some degree, indicative of the diet of the giraffids (Solounias et al. 2010, although a recent microwear analysis showed that things might be more complex (Merceron et al. 2018). ...
Eurasian Giraffidae went through a drastic biodiversity decline after the Miocene–Pliocene boundary; scanty palaeotragine populations are likely to have survived in Central Asia, providing the necessary stock for a Late Pliocene–Early Pleistocene expansion from Central Asia to Spain and from the Mediterranean to southern Russia. Here, we describe new giraffid findings from the Greek middle Villafranchian faunas of Dafnero-3 and Volax and from the late Villafranchian faunas of Tsiotra Vryssi and Krimni-3, and we revise previous material from Dafnero-1. Our results
support the synonymy of almost all the Villafranchian Eurasian giraffids under a single species of Palaeotragus, i.e. Palaeotragus inexspectatus, and allow us to improve its diagnosis. The orientation of the ossicones and the relative shortening of the lower premolar row might indicate affinities to some Late Miocene–Pliocene Palaeotragus from China. Our study suggests that P. inexspectatus was equally abundant at MNQ17 and MNQ18 in the Eastern Mediterranean and that its extinction after MNQ19 was probably attributable to the combination of the climatic and environmental turnover at the Villafranchian–Epivillafranchian boundary, along with the competition with emerging ruminant groups, such as giant cervids. A preliminary analysis of its palaeoecology suggests a giraffid more involved in grazing than its Late Miocene relatives.
... Αlthough time-range distributions from both China and Anatolia support the possible co-occurrence of similar-sized 'ovibovin' bovids with highly specialized cranial features, the Çorakyerler record is in fact the first confirmed evidence of such a joint presence in the same assemblage (see also Shi et al., 2016). Their co-occurrence might have been possible due to distinct feeding preferences and ethology, as deduced from indirect evidence for C. argalioides from Samos and Kemiklitepe D and H.? inundata from Garkin, based on differences in relative hypsodonty, masticatory morphology, horn shape, and limb proportions (Bouvrain, 1994;Köhler, 1993;Solounias & Dawson-Saunders, 1988;Solounias et al., 2010). According to those data, C. argalioides might have been a herd-living mixed feeder to grazer, with moderate running abilities and a complex fighting behavior (possibly combining pushing/ramming with stabbing/wrestling). ...
Fossil ‘ovibovin’ bovids are described from the Upper Miocene of Çorakyerler (north-central Anatolia). Two taxa have been recognized: the predominant Criotherium argalioides, known by several craniodental remains, and the less common Hezhengia? cf. inundata, documented by a few dentitions. A review of C. argalioides records from Samos and Kemiklitepe D and a thorough comparison with the Çorakyerler sample provides new data on the morphological and metric variability of this taxon, as well as its chronological and geographic range. Hezhengia? cf. inundata from Çorakyerler reveals important similarities with the Garkin (Turkey) taxon, which, in contrast to previous studies, we find more similar to the Chinese Hezhengia than to the Chinese Plesiaddax as originally suggested. The same is true for “Plesiaddax” simplex from Kayadibi (Turkey), which is referred to as Hezhengia? simplex. The co-occurrence of two ‘ovibovin’ bovids of similar size in the same assemblage was previously suspected but never before documented.
... To some extent, the length of premolar row is related to the feeding habits and the feeding efficiency. In ruminants, shorter premolar rows are related to more hypsodont teeth, which appear to intake more grass and adapt to a rather open living habitat, whereas longer premolar rows seem to correspond to a more soft, leafy diet (Solounias and Dawson-Saunders 1988;Spencer 1997;Mendoza et al. 2002). Danowitz et al. (2016) predicted the diets of some extinct giraffids in China and Greece by combined-mesowear analysis, and demonstrated that almost all the P. rouenii in Greece are browsers. ...
Palaeotragus is a common genus of the Giraffidae that was widely distributed in the Neogene strata of Eurasia. Early studies usually placed small giraffids in the Palaeotragus clade, leading to considerable controversy over the validity and systematic classification of this genus. Palaeotragus microdon and Palaeotragus cf. coelophrys are two representative species of Palaeotragus in China, their dental specimens are mainly distinguished by size, causing problems in their actual identification. In this study, several mandibles of a small giraffid found in Linxia Basin, Gansu Province, China (MN 10–13), are studied and identified as Palaeotragus cf. coelophrys. Based on a detailed comparison, the size and morphological variability of the Chinese Palaeotragus dental material are discussed, as well as the validity of the two species and the validity of other Eurasian Palaeotragus. This study reveals that the Chinese Palaeotragus has a continuous sequence of size variability and premolar features, which is insufficient to separate the mandible of the two species. In addition, the type species P. rouenii can be clearly distinguished from the Chinese Palaeotragus by the relatively long lower premolar row, possibly due to the difference in feeding habits. The holotype of P. coelophrys is closely related to Chinese Palaeotragus, while the P. coelophrys mandible from Maragheh (MMTT37 2517) should belong to different species.
... Plutarch, the well-known ancient Greek writer, and politician (45-120 AD) observed some fossilized bones found in the village of Mytilinioi of Samos Iceland (Solounias & Dawson-Saunders, 1988), refers to the bones of the Amazons (mythological women fighters), while Pausanias, the ancient Greek traveler and geographer (110-180 AD) encounters in a sanctuary of ancient God Asclepius, some huge bones, which, according to him, belonged to giants who protected his mother, the goddess Rhea. In 675 BC, when the Greek traveler, Aristaeus, visited Scythian nomads in the Gobi deserts, the nomads told him about an area beyond Sidonia where griffins defended gold from the nomads. ...
The present work is an attempt at a bibliographic overview in the field of paleontology and specifically in the
field of fossils regarding their value and connection with history and mythology, and how it has been used to
teach theory of evolution through natural selection (TENS) in a university course. To make our case, we use as
our paradigm two well-known historical locations of Greece, namely Thermopiles and Marathon. The area of
Marathon includes the location of Pikermi, which is very well known for its fossils that historically have been one
of the first locations rich in fossils that have been studied so extensively. Before we make a short tour of other
similar locations of the Greek peninsula that are good cases for teaching evolution using paleontology, we present
an example of how fossils can be formed via sedimentation. For that, we are using the case of Thermopile, while
in antiquity was chosen as the field of the famous battle as it was a very narrow strip of land between the mountain
and the sea, nowadays, due to sedimentary alluvial deposits, has become a rather large field. To make the reader
familiar with the fossils found around Greece and their topology, we present a short tour and some history about
the fossils found throughout its territory. And finally, we argue about how the fossils and the museum education
could be used to prepare the pupils for first contact with TENS.
Keywords: Pikermi fossils, museum education, paleontology, evolution teaching
... Several early studies found correlates of particular skull traits or metrics with such environmental variables (e.g., refs. 8,9 ), but these approaches did not take phylogeny into account, so their results are difficult to interpret. More recent work that incorporated phylogeny found support for a relationship between dietary ecology and traits such as hypsodonty, facial elongation, and mandibular shape 10,11 . ...
The role of environmental selection in generating novel morphology is often taken for granted, and morphology is generally assumed to be adaptive. Bovids (antelopes and relatives) are widely differentiated in their dietary and climatic preferences, and presumably their cranial morphologies are the result of adaptation to different environmental pressures. In order to test these ideas, we performed 3D geometric morphometric analyses on 141 crania representing 96 bovid species in order to assess the influence of both extrinsic (e.g. diet, habitat) and intrinsic (size, modularity) factors on cranial shape. Surprisingly, we find that bovid crania are highly clumped in morphospace, with a large number of ecologically disparate species occupying a very similar range of morphology clustered around the mean shape. Differences in shape among dietary, habitat, and net primary productivity categories are largely non-significant, but we found a strong interaction between size and diet in explaining shape. We furthermore found no evidence for modularity having played a role in the generation of cranial differences across the bovid tree. Rather, the distribution of bovid cranial morphospace appears to be mainly the result of constraints imposed by a deeply conserved size-shape allometry, and dietary diversification the result of adaptation of existing allometric pathways.
... Taxon-based methods, such as taxonomic uniformitarianism (Andrews, 1995;Faith and Lyman, 2019) or taxonomic analogy (Reed, 1998) draw ecological correlations between fossil taxa and their extant counterparts (be they the same or very closely related genera and species) and utilize all available paleoecological data on extinct taxa to reconstruct a species' ecological adaptations. Using this approach, one assumes niche stasis across space and time, an assumption that has been shown to be invalid in some taxa (Solounias and Dawson-Saunders, 1988;Curran, 2015). However, this method is used widely and provides a good starting point for paleoenvironmental reconstructions. ...
The Early Pleistocene is recognized as a time of major global climatic and environmental change, including increasing aridity, significant spread of grasslands, and substantial faunal turnovers and dispersals. Importantly, this is the first time hominins are found in Eurasia. Reconstructing the types of environments that existed during this time is imperative for understanding mammalian, including hominin, dispersal patterns relative to climatic change. One proposed dispersal corridor across Europe is the Danube River. Here we characterize the 2.2 to ~1.1 million years ago (Ma) paleoenvironments surrounding one of the tributaries to the Danube, the Olteţ River, in southern Romania using a multiproxy approach, including taxonomic uniformitarianism, dental mesowear, dental microwear, enamel stable isotope (carbon and oxygen), and coprolite/palynology analyses, and compare our results to other penecontemporaneous Eurasian sites. Older sites from this region, Grăunceanu and La Pietriş, both dating to 2.2–1.9 Ma, are reconstructed as being primarily open, though with some nearby woodlands and significant water resources. Fântâna lui Mitilan, which is younger (1.8–1.1 Ma), is reconstructed as slightly more closed, though still relatively open in nature. These results are similar to reconstructions for other Early Pleistocene Eurasian sites, including ones with and without hominins, suggesting that hominins were likely not inhibited from dispersing across Eurasia due to environmental constraints at this time.
... While extant Perissodactyla possess an enlarged and elongated premolar tooth row (P2-P4), ruminants often have shortened premolar tooth rows (Mendoza et al. 2002). These specific dental traits are most pronounced between specialized perissodactyl grazers like extant equids, in which the premolar tooth row exceeds the molar tooth row (M1-M3) in length, and grazing ruminant Cetartiodactyla and camelids which have elongated the third molar and reduced the premolar tooth row (Janis 1988, 1990a,b, Solounias & Dawson-Saunders 1988. The role of premolars and molars in the masticatory process is hence less comparable between Perissodactyla and Cetartiodactyla and likely such morphologies are also a result of dietary trait evolution. ...
Teeth are the hardest and most durable parts of the mammalian body. Even after millions of years they provide a wealth of paleobiologic information. Teeth reflect the interaction of mammals with the environment, as evident from their shape and traces of mastication on their surface. Teeth of fossil and extant mammals provide information on the diet, mode of food processing, biomechanics of mastication, and energy gain. Modern analytical and visualization techniques such as micro-computed tomography, high-resolution surface analysis, and 3D imaging have greatly boosted the research on dental function in recent years. The book “Mammalian Teeth – Form and Function” offers a comprehensive synopsis of the latest advances in the field of dental function research. It will be of interest not only to paleontologists and biologists, but also to students and scholars in archeology, animal nutrition, and dentistry.
... Helladotherium duvernoyi and Samotherium major have very similar body masses and sizes, justifying that they could reach the same vegetation heights (Merceron et al. 2018). Previous work has shown that even though H. duvernoyi bears some mixed-feeding features (Solounias and Dawson-Saunders 1988;Solounias et al. 1999), it is generally accepted that the taxon was a large browsing sivatheriine (Solounias et al. 2000;Solounias et al. 2010;Solounias et al. 2013;Solounias and Danowitz 2016a). Samotherium major was primarily considered a grazer (Solounias and Moelleken 1993;Solounias et al. , 1999Solounias et al. , 2000. ...
Kemiklitepe is a well-known locality with four recognised fossiliferous horizons, KTA to KTD, which have yielded a plethora of mammalian remains. Previous taxonomic studies indicate the presence of three giraffid taxa: Samotherium major and Palaeotragus rouenii from the uppermost three horizons, KTA, KTB and KTC, as well as Palaeotragus rouenii and Samotherium? sp. from the lowermost KTD horizon. In this study a new locality, Kemiklitepe-E, is presented for the first time. Kemiklitepe-E is located approximately 350 m NW of the classic Kemiklitepe locality. The fossiliferous sedimentary rocks at Kemiklitepe-E occur at the same stratigraphic level as localities KTA, KTB and KTC. The preliminary faunal list includes representatives of Proboscidea, Chalicotheriidae, Equidae, Bovidae and Giraffidae. Comprehensive descriptions and comparisons of the Kemiklitepe-E Giraffidae specimens suggest the co-occurrence of two large giraffids: Samotherium major and Helladotherium duvernoyi. Samotherium major, previously documented from this region, is the most common taxon at Kemiklitepe. Helladotherium duvernoyi is rare at Kemiklitepe and here reported for the first time. The two taxa coexisted during the middle Turolian in Greece and Western Anatolia. In addition, it is suggested that specimens of Samotherium? sp described from KTD possibly belong to Samotherium neumayri. Based on the stratigraphic position of fossiliferous rocks, as well as the faunal data presented herein, the newly discovered locality is considered to be of middle Turolian (MN12) age.
... The explicit, quantitative modelling of the relationships between functional traits of organisms and their environments in a more fundamentally Humboldtian spirit has recently been called ecometrics (Eronen, Polly, et al., 2010;Polly et al., 2011;Vermillion, Polly, Head, Eronen, & Lawing, 2018); for an early example, see Solounias and Dawson-Saunders (1988). Organisms interact with their biotic and abiotic environments via traits. ...
Aim
The links between geo‐ and biodiversity, postulated by Humboldt, can now be made quantitative. Species are adapted to their environments and interact with their environments by having pertinent functional traits. We aim to improve global ecometric models using functional traits for estimating palaeoclimate and apply models to Pleistocene fauna for palaeoclimate interpretation.
Location
Global at present day, Pleistocene of Europe for fossil data analysis.
Taxa
Artiodactyla, Perissodactyla, Proboscidea and Primates.
Methods
We quantify functional traits of large mammal communities and develop statistical models linking trait distributions to local climate at present day. We apply these models to the fossil record, survey functional traits, and quantitatively estimate climates of the past. This approach to analysing functional relationships between faunal communities and their environments is called ecometrics.
Results and main conclusions
Here, we present new global ecometric models for estimating mean annual and minimum temperature from dental traits of present day mammalian communities. We also present refined models for predicting net primary productivity. Using dental ecometric models, we produce palaeoclimate estimates for 50 Pleistocene fossil localities in Europe and show that the estimates are consistent with trends derived from other proxies, especially for minimum temperatures, which we hypothesize to be ecologically limiting. Our new temperature models allow us to trace the distribution of freezing and non‐freezing ecosystems in the recent past, opening new perspectives on the evolution of cold‐adaptive biota as the Pleistocene cooling progressed.
... These two species could reach the same foliage heights. Based on tooth morphology, we would have expected more grazing habits for the large palaeotragine compared to the sivatheriine, because small premolars compared to molars are usually taken as indicating more grazing habits in ruminants (Solounias and Dawson-Saunders 1988), but no significant difference in the present study seems to distinguish the species with the larger premolars (H. duvernoyi) from the similar-sized species with the smaller premolars (S. major; Table 4, Figs. 5 and 6). ...
Today, the family Giraffidae is restricted to two genera endemic to the African continent, Okapia and Giraffa, but, with over ten genera and dozens of species, it was far more diverse in the Old World during the late Miocene. We attempt to describe here how several species may have shared feeding resources in the Eastern Mediterranean. Dietary preferences were explored by means of Dental Microwear Textural Analysis in combination with estimation of body mass and the maximum height at which the various species were able to browse. One of our main results concerns the modern okapi, Okapia johnstoni. It is a forest dweller usually regarded as a browser, but we show that it might also forage on tough plants, possibly herbaceous monocots. Such feeding habits including portions of herbaceous monocotyledons were also found for some extinct species, especially the genera Samotherium and Palaeotragus. Palaeogiraffa shows a contrasted pattern: the specimens of P. pamiri from a site in Thrace were leaf-dominant browsers whereas those belonging to P. major and P. macedoniae from the Axios valley in Greece ingested herbaceous monocotyledons. Helladotherium duvernoyi, the only sivatheriine analyzed here is described as a leaf-dominant browser. The giraffine Bohlinia attica also falls within the leaf-dominant browser category but could browse on higher foliages than H. duvernoyi. On the whole, the reconstructed diets confirm the relationship between more grazing habits and smaller premolars, but not with higher dental crown height.
... Unlike the cervid (Lucentia aff. piernesis) and equid (Hippotherium intrans), the elevated Sr/Ca ratios of the bovid Miotragocerus sp. are incongruent with morphological and stable isotope data, which suggest a diet dominated by leaves with a small fruit component (Solounias and Dawson-Saunders, 1988;Spassov and Geraads, 2004;Merceron et al., 2007;Merceron, 2009;Eastham et al., 2016). This type of diet would typically be associated with relatively low Sr/Ca ratios (Burton et al., 1999;Lee-Thorp et al., 2003;Sponheimer et al., 2005;Drouet and Herbauts, 2008). ...
The early Late Miocene vertebrate locality of Rudabánya II (R. II) in northeastern Hungary preserves an
abundance of forest-adapted ungulate species. To better understand the ecological relationships within this ancient
ecosystem, we used analysis of enamel strontium/calcium (Sr/Ca) ratios to infer dietary preferences. The goals of
the analysis were to: i) determine whether these ungulate species specialized in specific plants or plant parts; ii)
discern whether the Sr/Ca ratios support what was previously suggested about the ecology of these species and iii) evaluate the factors that may have acted to promote coexistence within this diverse community of predominantly browsing herbivores. Results show significant differences in the diets of the sampled species. The highest Sr/Ca ratios were displayed by the suids Parachleuastochoerus kretzoii (fortelius et al., 2005) and Propotamochoerus palaeochoerus (pilgrim, 1926) implying a preference for Sr-rich underground plant parts. Elevated Sr/Ca ratios yielded by the cervid Lucentia aff. pierensis (thomas, 1951) and equid Hippotherium intrans (kretzoi, 1983) are indicative of intermediate feeding. The bovid Miotragocerus sp. (stromer, 1928) showed higher Sr/Ca ratios than
the gomphothere Tetralophodon longirostris (kaup, 1832), which is incongruent with morphological and stable
isotope data, and suggested browsing by both taxa. This finding is likely the result of a difference in digestive
physiology (ruminant vs. monogastric) rather than a difference in dietary behaviour. The lowest Sr/Ca ratios were displayed by the traguild Dorcatherium naui (kaup and scholl, 1834) and moschid Micromeryx flourensianus (lartet, 1851) suggesting a preference for Sr-poor fruits. Resource specialization and partitioning within the local environment likely acted to decrease interspecific competition and promote coexistence within the diverse ungulate community at R. II.
... Eremotherium Spillman, 1948 from northern South America and North America, and other Megatherium species from north central and north western South America). We cannot determine the degree to which higher hypsodonty values in megatheriids and mylodontids correspond to feeding on abrasive grasses rather than browsing on foliage, as has been done for living ungulates ( Janis, 1988;Solounias and Dawson-Saunders, 1988), simply because we cannot know the proportion of grass in their diet. However, one obvious factor in explaining differences in hypsodonty in ground sloths is the increased presence of grit caused by environmental differences resulting from geographic distribution, or environmental change over time, or particular habits. ...
... suggest a traditional style of browsing, with a small component of fruit and seed feeding (Merceron et al., 2007). Higher δ 13 C E and δ 18 O E values could indicate the seasonal consumption of fruits, and these data support those of Merceron et al. (2007), as well as several other studies of Miotragocerus (Solounias and Dawson-Saunders, 1988;Spassov and Geraads, 2004;Merceron et al., 2006;Merceron, 2009). ...
Examining how species use and partition resources within an environment can lead to a better understanding of community assembly and diversity. The rich early Late Miocene (early Vallesian) deposits at Rudabánya II (R. II) in northern central Hungary preserve an abundance of forest dwelling taxa, including the hominoid Rudapithecus hungaricus. Here we use the carbon and oxygen stable isotope compositions of tooth enamel carbonate from 10 genera of medium to large-bodied mammals to evaluate resource use and partitioning among the herbivore community, and to reconstruct the paleoenvironment of Rudapithecus. The range of stable carbon and oxygen isotope values (δ 13 C E and δ 18 O E) displayed by the R. II fauna indicates a variable forest environment, which included both open and closed canopy habitats. The relatively low δ 13 C E and δ 18 O E values found in all sampled taxa are consistent with high levels of precipitation and humidity. Significant differences in stable isotope values were observed among the sampled fauna, supporting the interpretation of resource specialization and partitioning. Higher δ 13 C E values found in Aceratherium incisivum (Rhinocerotidae), Lucentia aff. pierensis (Cervidae), Hippotherium intrans (Equidae), Tetralophodon longirostris (Gomphotheriidae), Propotamochoerus palaeochoerus and Parachleuastochoerus kretzoii (Suidae) suggest foraging in more open canopy habitats, while lower δ 13 C E values found in Miotragocerus sp. (Bovidae), Dorcatherium naui (Tragulidae), and Micromeryx flourensianus (Moschidae) imply a preference for more densely canopied habitats. Several of the sampled taxa yielded relatively higher δ 18 O E and δ 13 C E values indicative of fruit consumption, including the small ruminants, cervid, and bovid. The analyzed isotope values reflect a moderate degree of dietary niche overlap between taxa. An abundance of plant resources likely allowed for the coexistence of this diverse community of predominantly browsing herbivores. Within the gradient of more open to closed canopy forest, it is likely that Rudapithecus occupied dense closed canopy habitats where access to fruit was relatively continuous. The progressive fragmentation and replacement of humid forests by more open and seasonal woodlands during the late Vallesian would have had a significant influence on the extinction of this fossil ape.
... Well-developed lips are a common feature of ungulates due to their importance in food manipulation. Browsers in particular have well developed lips and lip musculature as compared with grazers and this can be seen osteologically in the relatively high facial region and well developed muscle scars where these muscles originate (Solounias and Dawson-Sanders, 1988;Janis, 1995). The ridges on the maxilla and premaxilla of Antilohyrax are likely due to bone remodeling resulting from the presence of well developed buccinator and maxillolabialis muscles. ...
The hyracoid Antilohyrax pectidens Rasmussen and Simons, 2000 from quarry L-41 in the Fayum, Egypt displays many interesting features, including a comb-like, pectinate lower first incisor similar to that of the dermopteran Cynocephalus. Antilohyrax was originally described as lacking upper incisors, and having retracted nasal bones and selenodont cheek teeth, functionally resembling characters found in bovid artiodactyls. Analysis of cranial and postcranial material led to the hypothesis that Antilohyrax was a cursorial browser.
Recent expeditions have recovered additional material that contributes greater detail about cranial and dental morphology and allows for the reassessment of characters previously unknown or misinterpreted. Among these is the presence of at least two pairs of upper incisors, the central pair of which form tusks as in all other hyracoids, and a slender nasal bone which projects anteriorly to the level of the premaxilla. There is evidence of a pad on the premaxilla that occludes with the pectinate lower incisors. In the mandible, a second pair of incisors has been recovered that are sickle shaped, lack pectinations, and occlude with the upper tusks. Other notable characters preserved in the new specimen include the presence of a complete postorbital bar, a large, round, and blunt postorbital boss, a lateral flange on the zygomatic arch, a deep antorbital groove of the frontal bone, a lambdoid crest, a long paroccipital process, and a unique nuchal region. Comparisons with newly recovered and as yet undescribed cranial material of Titanohyrax reveal more characters shared by these genera, strengthening the case for their inclusion as sister taxa within the Titanohyracinae. Examination of characters relevant to paenungulate phylogenetics confirm earlier observations that extant hyracoids have changes in cranial proportions that result in some character states not representative of early hyracoids. Functional inferences lend further support to the suggestion that Antilohyrax was a folivorous browser.
... Bovids have occupied a variety of habitats including closed forests, a variety of woodland biomes, and open plains. The bovids as habitat indicators have been widely studied and their value as habitat indicators in ancient ecosystems has been well-established (Gentry, 1970(Gentry, , 1980Thomas, 1979;Vrba, 1980Vrba, , 1995Scott, 1985;Solounias & Dawson-Saunders, 1988;Plummer & Bishop, 1994;Kappelman et al., 1997;Scott et al., 1999). The bovids are among the most abundant taxa at the Dhok Pathan Formation of Chakwal. ...
The new Late Miocene boselaphine remains have been recovered from the outcrops of the Padhri village, which belongs to the Middle Siwalik Subgroup. The specimens comprise mandible fragments and isolated dentitions. The material has been assigned to Pachyportax latidens, P. nagrii, Selenoportax vexillarius and Tragoportax punjabicus. Their morphometric features are discussed and compared to the material from the Siwalik Group.
... suggest a traditional style of browsing, with a small component of fruit and seed feeding (Merceron et al., 2007). Higher δ 13 C E and δ 18 O E values could indicate the seasonal consumption of fruits, and these data support those of Merceron et al. (2007), as well as several other studies of Miotragocerus (Solounias and Dawson-Saunders, 1988;Spassov and Geraads, 2004;Merceron et al., 2006;Merceron, 2009). ...
Examining how species use and partition resources within an environment can lead to a better understanding of community assembly and diversity. The rich early Late Miocene (early Vallesian) deposits at Rudabánya II (R. II) in northern central Hungary preserve an abundance of forest dwelling taxa, including the hominoid Rudapithecus hungaricus. Here we use the carbon and oxygen stable isotope compositions of tooth enamel carbonate from 10 genera of medium to large-bodied mammals to evaluate resource use and partitioning among the herbivore community, and to reconstruct the paleoenvironment of Rudapithecus. The range of stable carbon and oxygen isotope values (δ13CE and δ18OE) displayed by the R. II fauna indicates a variable forest environment, which included both open and closed canopy habitats. The relatively low δ13CE and δ18OE values found in all sampled taxa are consistent with high levels of precipitation and humidity. Significant differences in stable isotope values were observed among the sampled fauna, supporting the interpretation of resource specialization and partitioning. Higher δ13CE values found in Aceratherium incisivum (Rhinocerotidae), Lucentia aff. pierensis (Cervidae), Hippotherium intrans (Equidae), Tetralophodon longirostris (Gomphotheriidae), Propotamochoerus palaeochoerus and Parachleuastochoerus kretzoii (Suidae) suggest foraging in more open canopy habitats, while lower δ13CE values found in Miotragocerus sp. (Bovidae), Dorcatherium naui (Tragulidae), and Micromeryx flourensianus (Moschidae) imply a preference for more densely canopied habitats. Several of the sampled taxa yielded relatively higher δ18OE and δ13CE values indicative of fruit consumption, including the small ruminants, cervid, and bovid. The analyzed isotope values reflect a moderate degree of dietary niche overlap between taxa. An abundance of plant resources likely allowed for the coexistence of this diverse community of predominantly browsing herbivores. Within the gradient of more open to closed canopy forest, it is likely that Rudapithecus occupied dense closed canopy habitats where access to fruit was relatively continuous. The progressive fragmentation and replacement of humid forests by more open and seasonal woodlands during the late Vallesian would have had a significant influence on the extinction of this fossil ape.
... Previous microwear and mesowear investigations of giraffid paleodiets from Samos and Pikermi concluded that the majority of Samos and Pikermi giraffids were not browsers, but incorporated grass into their diets (Solounias and Dawson-Saunders, 1988;Solounias et al., , 2000Solounias et al., , 2010Solounias et al., , 2012. The dietary habits of the giraffids found in North China have never been studied or reported. ...
... Hypsodonty was used to assess diet in extinct species. High hypsodonty indices are correlated with diets high in abrasive materials (Janis 1988, Solounias and Dawson-Saunders 1988, Pérez-Barbería et al. 2001). Greater crown height helps teeth withstand abrasion by phytoliths and grit (McNaughton et al. 1985, Janis 1988, MacFadden, 1997 et al. (2002) and Mendoza and Palmqvist (2008). ...
... As first suggested by , the shape of the premaxillary tip is strongly dependent on the diet of the animal, being pointed in specialized browsers (to enable a better selection of the most palatable leaves and buds from the twigs), squared in specialized grazers (to allow the grasping of a larger amount of grass with each single bite) and intermediate in mixed feeders. Within the deer family, composed of more or less specialized browsers and mixed feeders, the premaxillary tip is a useful tool to infer the feeding behaviour of extinct species (Solounias and Dawson-Saunders, 1988;Solounias and Moelleken, 1993;Caloi and Palombo, 1995;Janis, 1995); according to Valli and Palombo (2005), it is the most useful among the morphological characters of the skull with functional meaning. Unfortunately the records are scarce. ...
The deer remains from the renowned Palaeolithic site of Isernia la Pineta (southern Italy) are revised. Both material already published, and new specimens from the last 15 years of excavation, is taken into account and some identifications are revised. The published assemblage consists of Megaceroides solilhacus, Cervus sp. cf. C. elaphus acoronatus, Dama sp. cf. D. clactoniana and Capreolus sp. The literature dealing with these species is discussed and the problems still unresolved are outlined. The giant deer, now assigned by most authors to the genus Praemegaceros as P. solilhacus, is the largest and most common deer species at the site, represented by numerous antler fragments plus a few isolated teeth and fragmentary postcranial elements. The retrieval of new, well-preserved specimens, namely one nearly complete antler, plus a distal portion of palmation, two neural skulls and one premaxillary bone, allows a better characterization of this otherwise still poorly understood species. The sub-specific attribution of the red deer, in the lack of more complete antler specimens, is still dubious. The fallow deer remains probably belong to Dama roberti, a recently described species from the early Middle Pleistocene of Western Europe. The morphology of the lower portion of the only preserved antler fits better with this species rather than with D. clactoniana, to which it was previously tentatively assigned. At last, the specific identity of the few roe deer remains from Isernia is discussed, but the problem remains unanswered because of lack of agreement, in the literature, about the taxonomy of the Pleistocene species. Copyright © 2015 John Wiley & Sons, Ltd.
... Masseter origination A suite of morphological characters has been associated with grazers and browsers. Janis (1995) summarized two studies, Solounias and Dawson-Saunders (1988) and Janis (1990), that described cranial, mandibular, and dental differences between grazers and browsers, with mixed feeders often having intermediate states. Mendoza et al. (2002) provided a few more character differences between grazers and browsers. ...
... These authors used this type of measurement, along with the palatal length, to define a 'relative muzzle width ratio', which they used to represent the ratio between body size and the oral aperture, as well as possibly representing oral intake and processing rate. Ratios are poor shape descriptors since all a ratio can represent adequately is an ellipse, if the two measurements represent orthogonal axes, as in the method used by Solounias and Dawson-Saunders [89]. This approach may be sufficient for partially representing extremes of the browser end of the shape spectrum, but can just as easily describe a typically blunt grazing form. ...
Snout shape is a prominent aspect of herbivore feeding ecology, interacting with both forage selectivity and intake rate. Previous investigations have suggested ruminant feeding styles can be discriminated via snout shape, with grazing and browsing species characterised by 'blunt' and 'pointed' snouts respectively, often with specification of an 'intermediate' sub-grouping to represent ambiguous feeding styles and/or morphologies. Snout shape morphology is analysed here using a geometric morphometric approach to compare the two-dimensional profiles of the premaxilla in ventral aspect for a large sample of modern ruminant species, for which feeding modes are known from secondary criteria. Results suggest that, when browsing and grazing ruminants are classified ecologically based on a range of feeding style indicators, they cannot be discriminated unambiguously on the basis of snout profile shape alone. Profile shapes in our sample form a continuum with substantial overlap between groupings and a diverse range of morphologies. Nevertheless, we obtained an 83.8 percent ratio of correct post hoc feeding style categorisations based on the proximity of projected profile shapes to group centroids in the discriminant space. Accordingly, this procedure for identifying species whose feeding strategy is 'unknown' can be used with a reasonable degree of confidence, especially if backed-up by additional information. Based on these results we also refine the definitions of snout shape varieties, taking advantage of the descriptive power that geometric morphometrics offers to characterize the morphological disparities observed. The shape variance exhibited by both browsing and grazing ruminants corresponds strongly to body mass, providing further evidence for an interaction between snout shape, feeding style, and body size evolution. Finally, by exploring the role of phylogenetic similarity in snout shape, we find a slight increase in successful categorisation when repeating the analysis with phylogenetic control on the geometric profiles.
... Understanding the evolutionary history of the herbivore niche within African bovids has traditionally relied on examining anatomical adaptations to diet, particularly those related to digestive strategy (Clauss et al. 2003;Codron et al. 2008a). From a palaeontological point of view, morphological characters discernible in the fossil record, such as anterior dentary shape, mandible shape and size and hypsodonty index, have been used to inform on the dietary preferences of both extant and extinct species (e.g. Gordon and Illius 1988;Solounias and Dawson-Saunders 1988;Solounias et al. , 1995Janis 1990Janis , 1995Janis , 2008Solounias and Moelleken 1993;Gordon 1999, 2001;MacFadden 2000;Williams and Kay 2001;Mendoza et al. 2002;Solounias and Semprebon 2002;Feranec 2007;Mendoza and Palmqvist 2007;Codron et al. 2008b;Faith et al. 2011Faith et al. , 2012Fraser and Theodor 2011). However, the significance of some of these traits in identifying dietary preferences has been re-examined in the light of the potentially confounding effects of phylogenetic affinity (P erez-Barberia and Gordon 1999Gordon , 2001Clauss et al. 2008;Raia et al. 2010). ...
Understanding the evolutionary history of the herbivore niche within African bovids has traditionally relied on examining anatomical adaptations to diet, particularly those related to digestive strategy. More recently, mesowear and stable isotope analyses have been used to great effect to reconstruct dietary preferences. We use these dietary proxies to construct a morphology-free dietary ecospace and examine the topology of the phylogenetic rela-tionships of African bovids mapped onto this ecospace. The reconstructed dietary ecospace provides evidence for four distinct dietary classes: species with C3-or C4-dominated diets that produce low or high occlusal relief, likely related to diets high or low in abrasives, respectively. We detected no evidence for a discrete mixed feeder category; the species often categorized as such represent the end members of groups of species with either C3-or C4-dominated diets. Our analysis reveals high variability within the C4 grazing ecospace, and phylogenetic evidence indicates at least two pathways to grazing, likely related to the abrasive qualities of ingested food, which may be determined by the moisture content or the height of consumed grasses. These different pathways probably contribute to the high diversity of African grazers, both today and in the fossil record. C3 browsers (non-frugivores) also display a high degree of variation, but there are no species associated with highly abrasive diets and there is evidence for only a single evolutionary pathway. We find evidence for only one evolutionary route towards frugivory, which includes species with diets that produce both high and low occlusal reliefs. The cause of abrasive wear in frugivores may be related to grit and/or the hard parts of fruits, but this requires further examination.
... Additional support for such consumption can be derived from the cranial functional morphology and the dentition of Myotragus. While up to now only the abrasive character of the diet of Myotragus has been inferred (Alcover et al., 1981, Marcus, in press), in recent years a number of workers have established a series of criteria to evaluate the palaeodiets of ruminant extinct species (Solounias & Dawson-Saunders, 1988;Solounias et al., 1995;Dompierre & Churcher, 1996). Analysis of the cranial morphology of Myotragus balearicus, including the morphology of the premaxillaries and the location and depth of the masseteric muscle, suggest an essentially browsing habit. ...
Myotragus balearicus Bate 1909 is an artiodactyl Caprinae endemic to the Balearic Islands which became extinct more than 4000 years ago. Coprolites produced by this species have been collected from the excavation of Holocene cave sediments in Cova Estreta (Serra de Tramuntana, Mallorca). The pollen content of several samples of coprolites has been studied in order to determine the diet of Myotragus. Myotragus balearicus from Cova Estreta was a browser, and consumed huge amounts of box, Buxus balearica, a plant known for its high content of steroidal alkaloids. The coprolites are very fine textured, probably due to the result of a very efficient digestive process.
... Eremotherium Spillman, 1948 from northern South America and North America, and other Megatherium species from north central and north western South America). We cannot determine the degree to which higher hypsodonty values in megatheriids and mylodontids correspond to feeding on abrasive grasses rather than browsing on foliage, as has been done for living ungulates (Janis, 1988;Solounias and Dawson-Saunders, 1988), simply because we cannot know the proportion of grass in their diet. However, one obvious factor in explaining differences in hypsodonty in ground sloths is the increased presence of grit caused by environmental differences resulting from geographic distribution, or environmental change over time, or particular habits. ...
The fossil xenarthrans include giant forms, the ground sloths (Tardigrada), characteristic of the mammal fauna of the Pleistocene of South America. Although most authors agree in considering them as herbivorous, these forms have not been studied in terms of detailed morpho-functional analyses of their masticatory apparatuses. The aim of this work is the study the masticatory apparatus of the large Pleistocene ground sloths Glossotherium robustum, Lestodon armatus, Mylodon darwini and Scelidotherium leptocephalum (Mylodontidae) applying biomecanichal and morphogeometrical methods, and to compare with the information obtained for Megatherium americanum (Megatheriidae). The results are integrated with recent ecomorphological analyses that include three variables (hypsodonty index, dental occlusal surface area and relative width and shape of the muzzle) providing useful information for the inference of dietary habits and to propose a niche partitioning among these species. Glossotherium robustum and Lestodon armatus, the wide-muzzled sloths, were mostly bulk-feeders (i.e. ingest great amounts of food with each bite; probably grass and herbaceous plants). Mylodon darwini and Scelidotherium leptocephalum, the narrow-muzzled sloths, were mixed or selective-feeders (i.e. select plants or plant parts; grass and/or tree and shrubs foliage). The tooth design of mylodontids indicates that teeth were used mainly for crushing and grinding turgid and fibrous items respectively. Megatherium americanum was probably the most selective feeder among these sloths, and selectively fed on particular plants (shrubs) or plant parts (leaves, twigs, fruits). Its dentition was designed mostly for cutting soft but tough items which might include flesh, leaving open the possibility of an omnivorous diet.
One of iconic Africa's Big Five, the African buffalo is the largest African bovine or antelope that occurs throughout most of sub-Sahara and in a wide range of ecosystems from savanna to rainforest. The African buffalo is also one of the most successful large African mammals in terms of abundance and biomass. This species thus represents a powerful model to enhance our understanding of African biogeography and wildlife conservation, ecology and management. Edited by four researchers experienced in different aspects of the African buffalo's biology, this volume provides an exhaustive compilation of knowledge on an emblematic species that stands out as an important component of African natural and human ecosystems. It delivers a global view of the African buffalo and all known aspects of its ecology and management. This book will appeal to students, scholars, scientists and wildlife managers as well as those enthusiastic about the charismatic species. This title is also available as Open Access on Cambridge Core.
One of iconic Africa's Big Five, the African buffalo is the largest African bovine or antelope that occurs throughout most of sub-Sahara and in a wide range of ecosystems from savanna to rainforest. The African buffalo is also one of the most successful large African mammals in terms of abundance and biomass. This species thus represents a powerful model to enhance our understanding of African biogeography and wildlife conservation, ecology and management. Edited by four researchers experienced in different aspects of the African buffalo's biology, this volume provides an exhaustive compilation of knowledge on an emblematic species that stands out as an important component of African natural and human ecosystems. It delivers a global view of the African buffalo and all known aspects of its ecology and management. This book will appeal to students, scholars, scientists and wildlife managers as well as those enthusiastic about the charismatic species. This title is also available as Open Access on Cambridge Core.
Reconstruction of palaeobiomes, ancient communities that exhibit a physiognomic and functional structure controlled by their environment, depends on proxies from different disciplines. Based on terrestrial mammal fossils, the late Miocene vegetation from China to the eastern Mediterranean and East Africa has been reconstructed as a single cohesive biome with increasingly arid conditions, with modern African savannahs the surviving remnant. Here, we test this reconstruction using plant fossils
spanning 14–4 million years ago from sites in Greece, Bulgaria, Turkey, the Tian Shan Mountains and Baode County in China, and East Africa. The western Eurasian sites had a continuous forest cover of deciduous or evergreen angiosperms and gymnosperms, with 15% of 1,602 fossil occurrences representing conifers, which were present at all but one of the sites. Raup–Crick analyses reveal high floristic similarity between coeval eastern Mediterranean and Chinese sites, and low similarity between
Eurasian and African sites. The disagreement between plant-based reconstructions, which imply that late Miocene western Eurasia was covered by evergreen needleleaf forests and mixed forests, and mammal-based reconstructions, which imply a savannah biome, throws into doubt the approach of inferring Miocene precipitation and open savannah habitats solely from mammalian dental traits. Organismal communities are constantly changing in their species composition, and neither animal nor
plant traits by themselves are sufficient to infer entire ancient biomes. The plant fossil record, however, unambiguously rejects the existence of a cohesive savannah biome from eastern Asia to northeast Africa.
The Late Miocene fossiliferous locality of Kerassia is located in northwestern Euboea, near the homonymous village. In the current undergraduate thesis material from all the excavations that have taken place in the area was studied. The material consists of bones from the most common mammal families that are represented in the Turolian faunas of the Balkan-Iranian Province. Perissodactyla (Equidae, Rhinocerotidae, Chalicotheriidae) and artiodactyla (Suidae, Giraffidae, Bovidae) are the most common taxa in the fauna of Kerassia. Other important taxa that were identified in the area are Aves, Proboscidea and Carnivora, which are considered rare findings. In the locality of Kerassia, two fossiliferous horizons have been recognized, as mentioned in previous works. The lower one includes the sites K2, K3 and K4, while the upper one includes the sites K1 and K6. The palaeoenvironment is characterized as mosaic savanna-woodland, but the species distribution in the two horizons indicates a transition into more forested conditions from the lower to the upper horizon.
Giraffids include the only living giraffomorph ruminants and are diagnosed by the presence of bi-lobed canines and a special type of epiphyseal cranial appendages called ossicones. The family Giraffidae ranges from the latest early Miocene until today. However they are currently extant relics with only two living representatives, the African genera Okapia and Giraffa. Giraffids were much more diverse and widespread in the past, with more than 30 fossil species described. For the past decades a number of studies intended to resolve the phylogenetic relationships of the family, but due to the lack of really good cranial material no clear consensus was reached regarding the phylogenetic relationships amongst the different members of the group. The exceptionally complete remains of a new large giraffid from the late Miocene of Spain, Decennatherium rex sp. nov., allows us to improve and reassess giraffid systematics, offering a lot of new data, both anatomic and phylogenetic, on the large late Miocene giraffids of Eurasia. The results of our cladistic analysis show Decennatherium as a basal offshoot of a clade containing the gigantic samotheres and sivatheres, characterized by the presence of a Sivatherium-like ossicone-plan among other features. Decennatherium thus offers the most ancient evidence of this Sivatherium-plan and firmly establishes the early morphological patterns of evolution of a sivathere / samothere-clade that is defined as the less inclusive clade that contains Decennatherium and Sivatherium. Finally, this large group of four-ossiconed giraffids evolutionarily links Miocene Europe and Africa indicating vicariance / migration processes among the giraffid genetic pools separated by the Mediterranean Sea.
http://onlinelibrary.wiley.com/doi/10.1111/ahe.12257/abstract
The aims of this study were to describe the anatomy of the mouth and
pharynx of the pampas deer, and to consider its evolutionary feeding niche
according to those characteristics. Gross dissections of the mouth and
pharynx were performed in 15 animals, 10 adult females and 5 young
animals under one year (3 males and 2 females), all dead by causes
unrelated to this anatomical region. The upper lip entered in the
constitution of a pigmented nasolabial plane. The masseter muscles
weighed 43.8 ± 3.5 g and represented 0.23% of body weight, which
corresponds to ruminants of feeders intermediate to grazers and browsers.
Parotid glands represented 0.08% of the body weight, characteristic that
also categorize the pampas deer as belonging to the intermediate feeding
group. The dental formula was the same of the domestic ruminants. The
upper incisors and canines were absent and instead of them there was a
dental pad (Pulvinus dentalis). The upper canine teeth were present only in
the deciduous dentition. The existence of a brachydont dentition turns
Ozotoceros very vulnerable to continuous use as there is no compensatory
teeth growth. The particular anatomy of the mouth and lips of this animal
was adapted to a very selective feeding, taking highly nutritious sprouts
beyond plant category. In conclusion and in addition to previous studies of
anatomy of the digestive organs in this species, pampas deer may be
categorized as belonging to the intermediate type of feeding.
The determination of the palaeoecological conditions of the Perivolaki fauna is given in the present article using various methods. The faunal diversity of Perivolaki is studied using several indices (Simpson, Shannon-Wiener, Whittaker) and indicates a homogeneous and equilibrated fauna with normal taxa distribution. The dental microwear analysis of the ungulates (Bovidae, Equidae) provided data about the feeding preferences of the identified taxa and in comparison to those from other localities a bushy-shruby-woody palaeoenvironment is possible for Perivolaki. The faunal composition of Perivolaki fauna suggests the dominance of the bovids and equids, indicating a relatively open environment. The comparison of the faunal composition of the Perivolaki fauna with those from various European localities, as well as with the recent ones from certain environments, using multivariate analysis suggests that the Perivolaki fauna matches to the recent open and dry ones. The faunal similarity is also studied using the Simpson's index, indicating close relations of the Perivolaki fauna to those of Axios valley than to those of Southern Greece (Pikermi, Halmyropotamos), Eastern Aegean Sea (Samos) and Asia Minor. All the available results from this study suggest for Perivolaki an open bushy-woody environment with grass undergrowth.
This chapter explains how understanding of tectonic events can aid the understanding of patterns of the climatic change that influenced Cenozoic mammal evolution on the Earth. The chapter shows that the reciprocal illumination is possible in the understanding of mammal evolution in the context of climatic changes-that is, climatic changes can help understand events in the mammalian fossil record and the patterns in the record as well. The late Eocene marked the time of great diversification of ungulates with teeth indicative of a leaf-eating diet with vegetation being more available for consumption by mammals in a more seasonal environment. Recurrence of a warming trend in the Miocene possibly combined with drying resulted in the spread of grasslands and a rebound in mammalian diversity from an Oligocene low. However, mammalian diversity declined in the later Miocene in combination with declining temperatures and possibly also with changing atmospheric conditions. Studies of the diets of ungulates reveal a diversity of browsers in the mid-Miocene unlike that in any present-day environment and may be reflective of non-analogous vegetational habitats under conditions of greater atmospheric carbon dioxide.
A new approach of reconstructing ungulate diets, the mesowear method, was introduced by Fortelius, Solounias (2000). Mesowear is based on facet development on the occlusal surfaces of the teeth. Restricting mesowear investigation on the M2 as has previously been suggested would limit application of the mesowear methodology to large ungulate assemblages. Most of the fossil, subfossil and recent ungulate assemblages that have been assigned to a single taxon have a smaller number of individuals. This results in the demand to extend the mesowear method to further tooth positions in order to obtain stable dietary classifications of fossil taxa. The focus of this paper is to test, if a consistent mesowear classification is obtainable for the remaining positions of the upper cheek tooth dentition (P2, P3, P4, M1 and M3) and for combinations of these tooth positions. For statistical testing, large assemblages of isolated cheek teeth of the Vallesian hipparionine horse Hippotherium primigenium Meyer, 1829 and of two populations of the recent zebra Equus burchelli Gray, 1824 are employed as models. Subsequently, all single cheek tooth positions and all possible combinations of these tooth positions are tested for their consistency in classification of the mesowear variables compared to the M2, the model tooth of Fortelius, Solounias (2000). As the most consistent model for the proposed "extended" mesowear method, the combination of four tooth positions P4, M1, M2, and M3 is identified, which allows to include the largest number of isolated tooth specimens from a given assemblage, and fulfills the demand of being consistent in the dietary mesowear classification with the "original" mesowear method. We propose the "extended" mesowear method to be particularly well suited for the reconstruction of paleodiets in hypsodont equids. © Publications Scientifiques du Muséum national d'Histoire naturelle, Paris.
Bovids are often used as paleoenvironmental proxies because they are among the most commonly recovered large mammals at many fossil hominin sites and because modern African bovids occupy a wide range of dietary and environmental niches. This study uses dental microwear texture analysis to examine 25 species of extant African bovids, representing six dietary categories and with an emphasis on various levels of mixed feeding. The results show significant differences among the dietary classifications and confirm previous work suggesting that grazing taxa have less complex, more anisotropic surfaces with smaller features than browsing taxa. The results also indicate that dental microwear texture analysis can distinguish beyond the classic grazer-browser-mixed feeder trichotomy and accurately separate variable grazers, generalists, browser-grazer intermediates and frugivores from obligate grazers and browsers, as well as from one another. Some differences among taxa within dietary categories were also found, probably reflecting seasonal and/or geographic differences in diet. In addition to demonstrating the effectiveness of the technique at differentiating between different levels of mixed feeding in bovids, this study also provides a comprehensive comparative dataset of extant bovid microwear textures that can be applied to fossil taxa from sites and time periods across Africa.
Myotragus balearicusBate 1909 is an artiodactyl Caprinae endemic to the Balearic Islands which became extinct more than 4000 years ago. Coprolites produced by this species have been collected from the excavation of Holocene cave sediments in Cova Estreta (Serra de Tramuntana, Mallorca). The pollen content of several samples of coprolites has been studied in order to determine the diet ofMyotragus.Myotragus balearicusfrom Cova Estreta was a browser, and consumed huge amounts of box,Buxus balearica, a plant known for its high content of steroidal alkaloids. The coprolites are very fine textured, probably due to the result of a very efficient digestive process.
Approximately one-third of the Earth's vegetative cover comprises savannas, grasslands, and other grass-dominated ecosystems. Paleobotanical, paleofaunal, and stable carbon isotope records suggest five major phases in the origin of grass-dominated ecosystems: (1) the late Maastrichtian (or Paleocene) origin of Poaceae; (2) the opening of Paleocene and Eocene forested environments in the early to middle Tertiary; (3) an increase in the abundance of C3 grasses during the middle Tertiary; (4) the origin of C4 grasses in the middle Miocene; and (5) the spread of C4 grass-dominated ecosystems at the expense of C3 vegetation in the late Miocene. Grasses are known from all continents except Antarctica between the early Paleocene and middle Eocene. Herbivore morphology indicative of grazing, and therefore suggestive of grass-dominated ecosystems, appears in South America by the Eocene-Oligocene boundary, prior to the occurrence of grazing morphology elsewhere, and persists throughout the Cenozoic. Clear vertebrate and paleobotanical evidence of widespread grass-dominated ecosystems in northern continents does not occur until the early to middle Miocene. C4 grasses are present from approximately 15 Ma and undergo a dramatic expansion in the lower latitudes of North America, South America, East Africa, and Pakistan between 9 and 4 Ma. The expansion may have taken place in a shorter interval in some regions. C4 grasses are characteristic of seasonal, arid, and warm environments and are more tolerant of lower atmospheric CO2 (< 400 ppmv) than C3 plants. C4 grass distribution, therefore, is climatically controlled. The late Miocene spread of C4 grasses possibly involved a decrease in atmospheric CO2 and heralded the establishment of modern seasonality and rainfall patterns.
Members of the African Bovidae exhibit dietary resource partitioning, which presumably allows coexistence of many species with herbivorous diets. Levels of resource partitioning based on diet include primary food preference, habitat preference, and feeding-height preference. Morphological correlates of these levels of resource partitioning were sought in the skull and vertebral column of 33 bovid species. A quantitative morphometric study of the mandible, skull, and thoracic vertebrae in a large sample of bovids (n > 700) explores these correlates using video-image analysis. Results of this study indicate that many significant morphological differences exist among bovids that have a diet of either grass, dicots, or some combination of these two primary resources. In addition, some variables significantly distinguished bovids with different habitat and feeding-height preferences. These morphological correlates crossed taxonomic boundaries and, in many cases, were related to the structural properties of the herbivorous diet.
Les fouilles récentes dans plusieurs gisements de Vertébrés «pontiens» de basse Macédoine ont livré d'abondants restes (principalement crâniens) de Paléotraginés, sous-famille qui n'avait jamais été signalée dans cette région. On peut distinguer trois espèces, appartenant aux genres Palaeotragus et Samotherium.
The savanna ungulate faunas of the North American Miocene were broadly similar to those of present-day East Africa in terms of overall morphological and taxomic diversity. However, the predominant ungulates of the African faunas are bovids, which possess bony horns that are primitively sexually dimorphic in their occurrence. The predominant ungulates of the North American Tertiary were equids, camelids and oreodonts, which all lacked horns. The limited number of horned ruminants were largely Miocene immigrants from Eurasia. Horns were also absent from the large-bodied herbivores in the endemic faunas of South America and Australia. Studies on living ungulates show that a strong correlation exists between habitat type, feeding behaviour, social behaviour and morphology. The importance of the post-Eocene climatic changes to the history of mammalian evolution is stressed. The primitive condition in eupecorans and protoceratids is the absence of horns. The first horned members of these divisions had horns in the males only. Small present-day antelope, where horns may also be present in the females of the species, are probably secondarily small. Horns were acquired independently in ruminant artiodactyls at least 3 times; a maximum number of 7 times is not unlikely. In each case, horns first appeared at a critical body weight of about 18kg, and in correlation with a change in habitat from closed to open woodland. Horns in living ruminants are associated with territorial defence by males holding exclusive feeding and reproductive territories in woodland habitats. Such behaviour in present-day antelope is correlated with a body size of greater than 15 kg and a folivorous diet. Perissodactyls never evolved sexually dimorphic bony horns of the type seen in ruminant artiodactyls because their foraging and digestive strategies necessitate a larger daily intake of food. Study of the morphology and paleoecology of oreodonts suggest that they were woodland herd-forming browsers with exclusively folivorous diets. Studies of the behaviour and morphology of of living members of the Ruminantia, and of the morphology and paleoecology of their fossil ancestors, suggest that they were primitively tree browsers living in closed woodland habitats. The radiation of the Bovidae into open grassy habitats in the Pliocene may have been dependent on the immigration of grazing equids into the Old World. During the Tertiary, the food resources in North America were more widely dispersed; this may have been the result of the trees being more widely spaced. A possible causal mechanism for this was the stable land mass of North American continent during the Tertiary resulting in a more continental climate, with more severe effect of the post-Eocene seasonality on the vegetation. Thus most endemic North American ruminants did not evolve horns because, at the critical combination of body size and diet seen in the evolution of horns in the Old World ruminants, the dispersal of the food resources within the vegetation was too great for an effective home range to be maintained as an exclusive territory. -from Author
Bovid (essentially antelope) biostratigraphy and phylogeny indicate a widespread environmental change in Africa near 2.5 Myr. Morphological and ecological analyses of the taxa lead to the inference that across sub-Saharan Africa a spreading of open grassland, probably caused by global reduction in temperature and associated changes in rainfall, occurred at around this time. -from Author
Broadleaved evergreen sclerophyllous taxa occupied a subhumid belt across much of North America-Eurasia by the middle Eocene. They originated from alliances in older laurophyllous forests that adapted to spreading dry climate. Since the continued trend to aridity finally restricted sclerophyllous vegetation to subhumid areas separated by drier tracts, it now occurs in areas with summer rain as well as in summer-dry mediterranean climates. Taxa of chaparral and macchia habit are common undershrubs in sclerophyll woodlands, to which they are seral. Shrublands spread only recently, though the adaptive structural features of the taxa are ancient and probably not pyrogenic. The history of Madrean-Tethyan sclerophyll vegetation illuminates three biogeographic problems. First, related taxa that link the Mediterranean-California areas are part of the larger problem of ties between these areas and those of summer rainfall, of taxa now in summer-rain areas that were in presently summer-dry areas into the early Pleistocene, and of the more numerous taxa that linked sclerophyllous vegetation of the Madrean-Tethyan regions during the Tertiary. The ties between summer-dry and summer-wet areas are relicts of the Neogene; taxa now in mediterranean-climate areas adapted functionally to these new climates during the Pleistocene; and most trans-Atlantic links owe to migration across a narrower ocean with more numerous islands, to a broader zone of subhumid climate, and to a more easterly trending Appalachian axis with numerous dry edaphic sites. Second, by the mid-Oligocene spreading dry climate had confined a formerly continuous temperate rainforest to southern Mexico, the West coast and the Appalachian area. Winter cold and summer drought exterminated it in the West, whereas in the East winter cold eliminated most evergreen dicots, leaving a dominantly deciduous hardwood forest there. The temperate "Appalachian disjuncts" in southern Mexico are therefore ancient, and did not migrate south to enter a forest previously without deciduous hardwoods, as others maintain. Third, the Canarian laurel forest derived its taxa from those in laurophyllous forests that covered northern Africa into the middle Miocene, not by southward migration from southern Europe in the Pliocene. Since many shrubs in the surviving laurel forest also contribute to macchia on bordering slopes, the ancient origin of their typical adaptive structural features is clearly implied.
Revision of the stratigraphy and depositional environmentsin the Eastern Neogene basin of Samos. Mapping, structural analysis together with new K/Ar dates provide the following general pattern: two fluvial-lacustrine cycles separated by a disconformity. Basal cycle is Astaracian-Vallesian, the disconformity occured between 9 and 8.5 MY, upper cycle is Turolian and has yielded the famous «Samos mammal fauna wich is between 8.5 and 7.0 MY old.RésuméRévision de la stratigraphie et des milieux de dépôtdu Néogène du bassin oriental de Samos. La cartographie, l'analyse structurale et de nouvelles datations isotopiques mettent en évidence deux cycles sédimentaires fluvio-lacustres séparés par une discordance. Le premier cycle est d'âge astaracien-vallésien, la discordance se situe entre 9 et 8,5 MA, le deuxième cycle est d'âge turolien et comprend la riche faune de mammifères, dite «de Samos, dont l'âge se situe entre 8,5 et 7,0 MA.
Specimens of Climacoceras africanus are described from Maboko, Kenya.
The new species Climacoceras gentryi is established on the basis of
ossicones, mandibles, and upper and lower dentitions from Fort Ternan
and Baringo, Kenya. By interpretation of its lower canines Climacoceras
is identified as a giraffoid and is placed in the new family
Climacoceridae. Canthumeryx sirtensis is identified from Muruarot and
Rusinga, Kenya. A dentition and associated partial skeleton of this
species are described. The teeth agree closely with specimens of the
same species from Gebel Zelten, Libya. Zarafa zelteni from Gebel Zelten
is synonymized with Canthumeryx sirtensis. Again on the basis of its
lower canines Canthumeryx is identified as a giraffoid and is placed in
the new family Canthumerycidae. Specimens of Palaeotragus primaevus are
described from Baringo, Kenya. This material includes a cranium with the
ossicones, skull roof, occipital and basicranial regions preserved.
Palaeotragus primaevus specimens from Fort Ternan are used in this
description and some of these are redescribed. The relations of the
giraffoids are assessed by methods of phylogenetic systematics.
Palaeomeryx, Prolibytherium and Propalaeoryx are excluded from the
Giraffoidea as their lower canines are not known. The Palaeotraginae is
shown to be an invalid polyphyletic grouping and the genus Palaeotragus
is also shown to be polyphyletic. Palaeotragus microdon is probably
synonymous with Palaeotragus rouenii and the three species Palaeotragus
rouenii (P. microdon), Palaeotragus coelophrys and Palaeotragus
quadricornis are retained in the genus Palaeotragus. It is suggested
that 'Palaeotragus' expectans and 'Palaeotragus' decipiens are closely
related to Samotherium. Palaeotragus primaevus is probably synonymous
with Palaeotragus tungurensis and this species is closely related to the
giraffines. With slight changes the subfamilies Sivatheriinae and
Giraffinae are valid monophyletic groups. Hydaspitherium is synonymized
with Bramatherium and the Sivatheriinae includes the genera
Giraffokeryx, Birgerbohlinia, Bramatherium and Sivatherium while the
Giraffinae includes the genera Honanotherium, Bohlinia and Giraffa and
the species 'Palaeotragus' tungurensis (P. primaevus). Okapia is
identified as the sister-group of the other giraffids. Triceromeryx is
the sister-group of the Giraffidae. Canthumeryx is the sister-group of
Triceromeryx plus the Giraffidae while Climacoceras is the sister-group
of the other giraffoids.
The evolution and distribution of flora that flourished during the time of the middle Miocene to Pleistocene Hipparion fauna is outlined. Owing to the vast geographic range of the fauna, extendfrom Spain to the Pacific and from southwestern Siberia and Transbaylcalia to central Africa, summary of the flora involves the discussion of several different fossil assemblages. The variety of assemblages serves to illustrate the migration of the various flora as climatic changes took place. It also illustrates the gradual development of the open-terrain to which the fauna was adapted. Grasslands undoubtedly more wooded than the modern steppes appeared; however, intrazonal communities of mesophyll forests also existed along rivers, lakes, valleys and in the mountains. The author proposes that migration of the true Hipparion fauna to America across land bridges from Asia did not take place because of the predominance of forests and taiga-Siberian formations in those northern areas. --F. M. Hueber.
1) The savanna ungulate faunas of the North American Miocene were broadly similar to those of present‐day East Africa in terms of overall morphological and taxonomic diversity. However, the predominant ungulates of the African faunas are bovids, which possess bony horns that are primitively sexually dimorphic in their occurrence. The predominant ungulates of the North American Tertiary were equids, camelids and oreodonts, which all lacked horns. A limited number of horned ruminants were present, but these were largely Miocene immigrants from Eurasia. Horns were also absent from the large‐bodied herbivores in the endemic faunas of South America and Australia.
(2) The absence of horns in equids and tylopod artiodactyls is unlikely to be due to genetic insufficiency. Bony horns were present in brontotheres, which were closely related to equids, and in protoceratids, which were closely related to camelids. Nasal horns were present in one oreodont genus.
(3) Studies on living ungulates show that a strong correlation exists between habitat type, feeding behaviour, social behaviour and morphology. It is possible to use the morphological remains of extinct ungulates to reconstruct the types of feeding and social behaviour, and to use the distribution of morphologies and body sizes in a community of mammals, in conjunction with geological and paleobotanical evidence, to reconstruct the type of habitat.
(4) The importance of the post‐Eocene climatic changes to the history of mammalian evolution is stressed. Continents at higher latitudes have become increasingly seasonal in terms of temperature and rainfall since the equable global conditions of the early Tertiary. Savanna mosaic were the predominant biome in North America by the early Miocene, and in Eurasia by the middle Miocene. Living temperate‐latitude species of ungulates may not be a reliable guide for the assessment of the interrelationship between behaviour and morphology in an evolutionary perspective, as their behaviour may have been recently adapted to a habitat type that has only been in existence since the Pleistocene.
(5) The primitive condition in eupecorans and protoceratids is the absence of horns, with the presence of large sabre‐like canines in the males. The first horned members of these divisions had horns in the males only. Small present‐day antelope, where horns may also be present in the females of the species, are probably secondarily small.
(6) Horns were acquired independently in ruminant artiodactyls at least three times, and a maximum number of seven times is not unlikely. In each case, horns first appeared at a critical body weight of about 18 kg, and in correlation with a change in habitat from closed to open woodland.
(7) Horns in living ruminants are associated with territorial defence by males holding exclusive feeding and reproductive territories in woodland habitats. Such behaviour in present‐day antelope is correlated with a body size of greater than 15 kg and a folivorous diet. It is argued that horns evolved in ruminant artiodactyls on the adoption of this type of territorial behaviour once the critical combination of body size, diet and habitat type had been attained in their evolution from small, essentially frugivorous, forest‐dwelling animals.
(8) Perissodactyls never evolved sexually dimorphic bony horns of the type seen in ruminant artiodactyls. This is because their foraging and digestive strategies necessitate a larger daily intake of food. In a woodland habitat they were never able to adopt a feeding area small enough to make exclusive territory maintenance an economical proposition. Territory holding in male perissodactyls is seen, but under the opposite conditions of habitat to territorial behaviour in ruminant artiodactyls.
(9) Study of the morphology and paleoecology of oreodonts suggests that they were woodland herd‐forming browsers with exclusively folivorous diets. They probably had some forestomach fermentation, but did not chew the cud. Similar studies of Tertiary camelids suggest that they were predominantly selective browsers eating herbage at a low level in open country and formed mixed‐sex feeding groups. These combinations of feeding and social behaviour suggest a more open structure of the mid‐Tertiary habitat in North America than in Eurasia.
(10) Studies of the behaviour and morphology of living members of the Ruminantia, and of the morphology and paleoecology of their fossil ancestors, suggest that they were primitively tree browsers living in closed woodland habitats. Such habitats were abundant in the Old World, but in limited supply in North America during the Oligocene, where the protoceratids were the only ungulates to parallel the eupecoran type of feeding and social behaviour. South America appears to have had an even more open habitat in the Oligocene than North America, and no parallel to the eupecorans was seen amongst the indigenous ungulates. The radiation of the Bovidae into open grassy habitats in the Pliocene may have been dependent on the immigration of grazing equids into the Old World.
(11) I conclude that there was a difference in habitat structure between North America and the Old World during the Tertiary. The food resources in North America were more widely dispersed, and this may have been the result of the trees being more widely spaced. A possible causal mechanism for this was the stable land mass of the North American continent during the Tertiary, resulting in a more continental climate, with a more severe effect of the post‐Eocene seasonality on the vegetation. The faunal record of the two continents also implied a greater density of trees in the Old World.
(12) Thus most endemic North American ruminants did not evolve horns because, at the critical combination of body size and diet seen in the evolution of horns in the Old World ruminants, the dispersal of the food resources within the vegetation was too great for an effective home range to be maintained as an exclusive territory.
(13) Attention is drawn to the dangers of constructing evolutionary stories about living animals without primary reference to the fossil record to see if the hypotheses are upheld, and of assuming that fossil animal communities can be made to fit models of existing communities.
The types of social organisation displayed by the African antelope species have been assigned in this paper to five classes, distinguished largely by the strategies used by the reproductively active males in securing mating rights, and the effects of those strategies on other social castes. The paper attempts to show that these strategies are appropriate to each class because of the effects of other, ecological, aspects of their ways of life. The paper describes different feeding styles among antelope, in terms of selection of food items and coverage of home ranges. It argues that these feeding styles bear a relationship to maximum group size of feeding animals through the influence of dispersion of food items upon group cohesion. The feeding styles also bear a relationship to body size and to habitat choice, both of which influence the antelope species' antipredator behaviour. Thus feeding style is related to anti-predator behaviour which, in many species, influences minimum group size. Group size and the pattern of movement over the annual home range affect the likelihood of females being found in a given place at a given time, and it is this likelihood which, to a large extent, determines the kind of strategy a male must employ to achieve mating rights. The effects of the different strategies employed by males can be seen in such aspects of each species' biology as sexual dimorphism, adult sex ratio, and differential distribution of the sexes.
The abstract for this document is available on CSA Illumina.To view the Abstract, click the Abstract button above the document title.
Supported by the study of Miocene bovids, mainly from the Indian Subcontinent, the Arabian Peninsula and Africa, a provisional synthesis of the bovid biogeographical history is provided. Different aspects are examined successively: the initial bovid radiation, the evolution and biogeography of the Boselaphini, the intercontinental faunal exchanges, the biostratigraphic correlations, the emergence of the Ethiopian fauna and some of the major characteristics of the evolution of the palaeoenvironments.
The People's Republic of China has designated a system of reserves in the mountains that rim the Sichuan Basin as areas to be managed specifically for the conservation of the giant panda Ailuropoda melanoleuca. We were invited to visit three of these reserves in April and May 1981 as guests of the China Association for Science and Technology and the Ministry of Forestry. Two reserves, Wanglang (27,000 ha) and Tangjiahe (40,000 ha) are located in the headwaters of the northern tributtaries of the Chang Jiang (Yangtze) in the Min Shan, 400 km north of Chengdu. The Fengtongzhai (40,000 ha), type locality of Ailuropoda, is in the Qionglai Shah, 250 km west of Chengdu. We describe the physiography, faunistic and floristic position, and conservation management of these areas. Major issues in the conservation of Ailuropoda are discussed.
The Deer and the Tiger is Schaller's detailed account of the ecology and behavior of Bengal tigers and four species of the hoofed mammals on which they prey, based on his observations in India's Kanha National Park. "This book is a treasure house of biological information and it is also a delight to read. . . . Excellent phoographs accompany the text."âRobert K. Enders, American Scientist "The one book that has been my greatest source of inspiration is The Deer and the Tiger by George Schaller, based on the first ever scientific field study of the tiger. . . . This book is written by a scientist, but speaks from the heart. . . . It reveals startling information on feeding habitats, territorial behaviour, and the nuances that make up the language of the forest; you become totally immersed in the world of the tiger. . . . For all of us who work in tiger conservation, this book is the bible."âValmik Thapar, BBC Wildlife
Thesis (Ph.D.)--University of Chicago, Department of Paleozoology. Includes bibliographical references.
Masticatory responses of pygmy goats (Capra hircus) to different foods
- De Vree
De Vree, F. and Gans, C., 1973. Masticatory responses of pygmy goats (Capra hircus) to different foods. Ann. Soc. R. Zool. Belg., 105 (34): 255-306.
Animaux Fossiles et Géologie de l'Attique
- Gaudry
Gaudry, A., 1862-1867. Animaux Fossiles et G~ologie de l'Attique. Martinet, Paris, 476 pp.
Late Cenozoic palynology Aspects of Palyno-logy
- E Leopold
Leopold, E., 1969. Late Cenozoic palynology. In: R.S. Tschudy and R.A. Scott (Editors), Aspects of Palyno-logy. Wiley Interscience, New York, N.Y., pp. 377-438.
The Ruminant Stomach (East Afr
- R R Hofmann
Hofmann, R. R., 1973. The Ruminant Stomach (East Afr. Monogr. Biol.). East African Literature Bureau, Nai-robi, 2, pp. 1-354.
East African Mammals. 3B. Large Mammals
- Kingdon
Kingdon, J., 1979. East African Mammals. 3B. Large Mammals. Academic Press, London, 436 pp.
Co-evolution of the Hipparion fauna and vegetation in the Paretethys region
- Gabunia
Gabunia, L. K. and Chochieva, K. I., 1982. Co-evolution of the Hipparion fauna and vegetation in the Paratethys region. Evol. Theory, 6: 1-13.
Adaptation Conceptual Issues in Evolutionary Biology
- R C Lewontin
Lewontin, R. C., 1984. Adaptation. In: E. Sober (Editor), Conceptual Issues in Evolutionary Biology. MIT Press, Cambridge, pp. 235 251.
Erste Ergebnisse einer palynologischen Untersuchung der Lignite von Pikermi/Attica
- Von Orgetta
Von Orgetta, M., 1979. Erste Ergebnisse einer palynologis-chen Untersuchung der Lignite von Pikermi/Attica. In: VIIth Int. Congr. Medit. Neogene, Athens, 1979. Ann. Geol. Pays Hell., 1: 909-921.
East African Mammals. 3D. Bovids
- Kingdon
Kingdon, J., 1982b. East African Mammals. 3D. Bovids. Academic Press, London, pp. 394 746.
The Mammals of the Southern African Subregion. Univ. Pretoria, CTP Printers Cape, Pretoria, 736 pp. Solounias, N., 1981a. Mammalian fossils of Samos and Pikermi. Part 2: Resurrection of a classic Turolian Fauna
- R H N Smithers
Smithers, R. H. N., 1983. The Mammals of the Southern African Subregion. Univ. Pretoria, CTP Printers Cape, Pretoria, 736 pp. Solounias, N., 1981a. Mammalian fossils of Samos and Pikermi. Part 2: Resurrection of a classic Turolian Fauna. Ann. Carnegie Mus., 50: 231-270.
Die Vorgeschichte der Einhufer. Sauge-tierkd
- E Thenius
Thenius, E., 1966. Die Vorgeschichte der Einhufer. Sauge-tierkd., 31 (2): 150-171.