Beatriz Azanza

Ancestral feeding state of ruminants reconsidered: earliest grazing adaptation claims a mixed condition for Cervidae.

BackgroundSpecialised leaf-eating is almost universally regarded as the ancestral state of all ruminants, yet little evidence can be cited in support of this assumption, apart from the fact that all early ruminants had low crowned cheek teeth. Instead, recent years have seen the emergence evidence contradicting the conventional view that low tooth crowns always indicate leaf-eating and high tooth crowns grass-eating.ResultsHere we report the results of two independent palaeodietary reconstructions for one of the earliest deer, Procervulus ginsburgi from the Early Miocene of Spain, suggesting that despite having lower tooth crowns than any living ruminant, this species included a significant proportion of grass in its diet.ConclusionThe phylogenetic distribution of feeding styles strongly supports that leaf-grass mixed feeding was the original feeding style of deer, and that later dietary specialization on leaves or grass occurred independently in several lineages. Evidence for other ruminant clades suggests that facultative mixed feeding may in fact have been the primitive dietary state of the Ruminantia, which would have been morphologically expressed only under specific environmental factors.

Vertebrate taphonomy in circum-lake environments: three cases in the Guadix-Baza Basin (Granada, Spain)

Abstract This work analyses the origin and taphonomy of three Plio-Pleistocene mammal assemblages from the Guadix-Baza basin (Granada, Spain): Huelago; Huescar-1; and Cullar de Baza-1. Similarities and differences in the taphonomic features are evaluated with regard to marginal lacustrine sedimentary processes involved in the accumulation of these mammal assemblages. Selective preservation of skeletal-part and taxa in relation to facies have been explored to determine their suitability for palaeoecological analysis. The mortality in Cullar de Baza-1 and Huelago-1 corresponds to a catastrophic pattern, but other taphonomic features of the bones indicate that accumulation processes were diachronic as occurred in Huescar-1. This could be explained by selective-predation and/or seasonal accumulation of bones. Hydraulic sorting and transportation greatly controlled the size and shape of accumulated skeletal elements, and the different preservational circumstances of each environment largely constrained the taxonomic composition of the fossil bone assemblages. Autochthonous fauna only occurs in Huescar-1 corresponding to subaquatic environments. The absence of carnivores among the identifiable remains from Huelago-1 has no plausible taphonomic or sedimentological explanation. Nevertheless, after taking into account the taphonomic factors, the differences in taxonomic composition between these three mammal assemblages reflect differences in the surrounding habitats. The great diversity of ruminant artiodactyls, in particular browsers, is indicative of wooded or bushy areas in the surroundings of Huelago-1 although not in the immediate area of the lake margin. This diversity coincides with that observed in other Villafranchian localities and represents the greatest diversity of artiodactyls known from the Neogene and Quaternary record of Western Europe. In turn, the high diversity of sub-aquatic forms and a smaller variety of artiodactyl species in Huescar-1, reflects an open, sparsely forested environment susceptible to seasonal drought. A similar sparse forested environment has been deduced for Cullar de Baza-1. The climate and environmental conditions in Huescar-1 and Cullar de Baza-1 are consistent with a climatic deterioration at the beginning of the Middle Pleistocene (‘Glacial Pleistocene’).

A stratigraphical framework for Miocene (MN4-MN13) continental sediments of Central Spain

New bio- and magnetostratigraphic data from the Miocene continental sediments of Central Spain are used to update the existing stratigraphical framework. Our revised record is based on the study of more than two hundred mammal faunas, ranging from the Late Ramblian (ca 18 Ma) to the Late Turolian (ca 6 Ma).

Dietary innovations spurred the diversification of ruminants during the Caenozoic

Global climate shifts and ecological flexibility are two major factors that may affect rates of speciation and extinction across clades. Here, we connect past climate to changes in diet and diversification dynamics of ruminant mammals. Using novel versions of Multi-State Speciation and Extinction models, we explore the most likely scenarios for evolutionary transitions among diets in this clade and ask whether ruminant lineages with different feeding styles (browsing, grazing and mixed feeding) underwent differential rates of diversification concomitant with global temperature change. The best model of trait change had transitions from browsers to grazers via mixed feeding, with appreciable rates of transition to and from grazing and mixed feeding. Diversification rates in mixed-feeder and grazer lineages tracked the palaeotemperature curve, exhibiting higher rates during the Miocene thermal maxima. The origination of facultative mixed diet and grazing states may have triggered two adaptive radiations—one during the Oligocene–Miocene transition and the other during Middle-to-Late Miocene. Our estimate of mixed diets for basal lineages of both bovids and cervids is congruent with fossil evidence, while the reconstruction of browser ancestors for some impoverished clades—Giraffidae and Tragulidae—is not. Our results offer model-based neontological support to previous palaeontological findings and fossil-based hypothesis highlighting the importance of dietary innovations—especially mixed feeding—in the success of ruminants during the Neogene.

The Middle Miocene mammalian siteof Belometchetskaya, North Caucasus: An important biostratigraphic link between Europe and China

Abstract The Middle Miocene fauna from Belometchetskaya, North Caucasus, provides an important biostratigraphiclink that permits refinement of correlations between the European land mammal zones (MN zones) and the faunal ages of China. This study concentrates on a review of the mammalian fauna from Belometchetskaya because a certain degree of confusion has entered the literature regarding its correlation to the MN sequence (it has been variously correlated to MN 4, MN 5 or MN 6) as well as its position within the Miocene (Early Miocene or Middle Miocene). It is concluded that the faunal elements hitherto considered to be of Oligocene age are in fact typical of the Middle Miocene and that the best correlation of the fauna to the European sequence is to MN 6. This means that Belometchetskaya is close in age to Sansan (France), Pasalar and Candir (Turkey), Koujiacun (China), Chinji (Pakistan) and Fort Ternan (Kenya).

Diversification of mammals from the Miocene of Spain

Abstract The mammalian fossil record of Spain is long and taxonomically well resolved, offering the most complete record of faunal change for the Neogene of Europe. We evaluated changes in diversification, composition, trophic structure, and size structure of large mammals over the middle and late Miocene with methods applied to this record for the first time, including ordination of fossil localities to improve temporal resolution and estimation of confidence intervals on taxa temporal ranges. By contrast, analysis within the traditional Mammal Neogene (MN) biochronology obscures important aspects of diversification. We used inferred temporal ranges of species and evaluated per capita rates of origination, extinction, diversification, and turnover over 0.5-Myr time intervals. Three periods of significant faunal change occurred between 12.0 and 5.5 Ma: (1) From 12.0 to 10.5 Ma, elevated origination rates led to an increase in diversity without significant change in ecological structure. Immigrants and geographic-range shifts of species to lower latitudes during an interval of global cooling contributed to these faunal changes. (2) From 9.5 to 7.5 Ma, high extinction rates followed by high origination rates coincided with significant changes in taxonomic composition and ecological structure. These changes represent the Vallesian Crisis, with replacement of a fauna of forest affinities (with frugivores and browsers) by a fauna of open woodlands (with grazers and mixed feeders). (3) From 6.5 to 5.5 Ma, high extinction rates reduced diversity without substantial changes in ecological structure, and large mammal faunas became highly endemic across the northern Mediterranean region. This interval includes the Messinian Salinity Crisis, the desiccation of the Mediterranean basin. Extinction may have been caused by geographic isolation and aridification, with evolution of endemic lineages giving rise to new species in the early Pliocene. These distinct macroevolutionary patterns of faunal change correspond to different geographic scales of inferred climatic and tectonic drivers.

The early Turolian (late Miocene) Cervidae (Artiodactyla, Mammalia) from the fossil site of Dorn-Dürkheim 1 (Germany) and implications on the origin of crown cervids

Dental and cranial appendage remains of Cervidae from the fossil site of Dorn-Dürkheim are studied in detail. The material mainly includes isolated teeth, isolated pedicles and antler pieces. Neither tooth rows nor complete appendages are recorded. Comparative morphology and statistics of morphometrics (principal component analysis and discriminant analysis) allow for the classification of small and large dentitions, small cranial appendages, two morphotypes of large pedicles and two morphotypes of large antlers. Possible combinations of the classified units document the sympatric occurrence of three species, namely, Procapreolus sp., Muntiacinae gen. and sp. indet., cf. Cervavitulus mimus, but the fragmentary condition of the material leads to ambiguity regarding their composition and, consequently, to a certain extent regarding the taxonomic identification. However, these remains indicate the contemporaneous occurrence of early Turolian members of the crown cervids Muntiacinae and Capreolinae and close a previous spatiotemporal gap in the European cervid record. In addition, their presence proves the progressive turnover from dichotomous-antlered muntiacines to early monopodial-antlered crown cervids from NE to SW Europe in the late Miocene. The taxonomical assignment challenges the recent hypothesis on the origin of crown Cervidae around the middle/late Miocene border since Dorn-Dürkheim cervids provide further evidence for the successive achievement of derived characters in cranial appendages of crown cervids (mediopostorbital position and backwards orientation of pedicles, coronet development, shaft development/elongation, beam development and increase in number of antler tines) in the lineage of crown cervids, which originated during the middle Miocene.

Key innovations in ruminant evolution: a paleontological perspective

Key innovations are newly acquired structures that permit the performance of a new function and open new adaptive zones, and are, therefore, of paramount significance for understanding the history of the Ruminantia, particularly its diversification through the Miocene. Here we review and discuss what is known about these evolutionary novelties, with special emphasis on the appearance and evolution of cranial appendages and high-crowned (or hypsodont) teeth. Cranial appendages probably favored the diversification of pecorans by being structures strongly related to sexual selection, whereas the acquisition of hypsodont teeth could have expanded potential dietary breadth and allowed species to extend diets into the grazing range without eliminating browsing as a potential diet. When analyzed in conjunction with patterns of faunal diversity and in the context of climatic changes, it seems that the overall view that ruminant cranial appendages and hypsodonty may have started responding to increased patterns of seasonality and the opening-up of ecosystems is roughly valid. Instead, they occurred through several distinct pulses and varied widely among continents. This review is, to our knowledge, the first to highlight that the evolution of these innovations has been far from constant and uniform through time. Furthermore, we identify that both a first attempt to increase hypsodonty starting in the early Miocene and a first evolution of antlers and pronghorns were interrupted as a consequence of wide climatic fluctuations in the early-middle Miocene transition.

Origin of an assemblage massively dominated by carnivorans from the miocene of Spain.

Carnivoran-dominated fossil sites provide precious insights into the diversity and ecology of species rarely recovered in the fossil record. The lower level assemblage of Batallones-1 fossil site (Late Miocene; Madrid Basin, Spain) has yielded one of the most abundant and diversified carnivoran assemblage ever known from the Cenozoic record of mammals. A comprehensive taphonomic study is carried out here in order to constrain the concentration mode of this remarkable assemblage. Another distinctive feature of Batallones-1 is that the accumulation of carnivoran remains took place in the context of a geomorphological landform (cavity formation through a piping process) practically unknown in the generation of fossil sites. Two characteristics of the assemblage highly restrict the probable causes for the accumulation of the remains: (1) the overwhelming number of carnivorans individuals; and (2) the mortality profiles estimated for the four most abundant taxa do not correspond to the classic mortality types but rather were the consequence of the behavior of the taxa. This evidence together with other taphonomic data supports the hypothesis that carnivoran individuals actively entered the cavity searching for resources (food or water) and were unable to exit. The scarcity of herbivores implies that the shaft was well visible and avoided by these taxa. Fossil bones exhibit a very good preservation state as a consequence of their deposition in the restricted and protective environment of the chamber. Batallones-1 had another assemblage (upper level assemblage) that was dominated by herbivore remains and that potentially corresponded to the final stages of the cavity filling.

Dietary behaviour and competition for vegetal resources in two Early Miocene pecoran ruminants from Central Spain

ABSTRACT Dietary behaviour and competition for resources are investigated for the small-sized ruminants Andegameryx Ginsburg, 1971 and Procervulus Gaudry, 1877 representatives of two largely distinct states of diversification of pecorans. Results obtained from dental microwear and mesowear methodologies are concordant with a mixed feeder strategy for the taxa from the Early Miocene environments of the Iberian Chain (Central Spain). Further, the Spanish taxa investigated had less abrasive diets than their relatives from others similarly aged localities in Europe. This fact raises an important evolutionary uncertainty concerning the traditional characterization of first pecorans as specialized browsers. Instead, data strongly corroborate the recently proposed notion that some Pecora ruminant lineages were able to consume a mixture of browse and grass, and that were originally facultative mixed feeders. However, there is a large degree of variation of their dental wear features. Species were not dependent on a limited type of vegetation and, consequently, were able to exploit different food resources. Dental wear data seem to indicate that these ruminants did not compete for vegetation. In addition, last Andegameryx and earliest Procervulus apparently had a sufficiently different body size to partition the available vegetation in terms of height above ground level to elude competition. Body size difference coupled with an important degree of dietary opportunism may have been factors necessary for their coexistence.