Investigating the dietary niches of fossil Plio-Pleistocene European macaques: The case of Macaca majori Azzaroli, 1946 from Sardinia.

The genus Macaca includes medium- to large-bodied monkeys and represents one of the most diverse primate genera, also having a very large geographic range. Nowadays, wild macaque populations are found in Asia and Africa, inhabiting a wide array of habitats. Fossil macaques were also present in Europe from the Late Miocene until the Late Pleistocene. Macaques are considered ecologically flexible monkeys that exhibit highly opportunistic dietary strategies, which may have been critical to their evolutionary success. Nevertheless, available ecological information regarding fossil European species is very sparse, limiting our knowledge of their evolutionary history in this geographic area. To further our understanding of fossil European macaque ecology, we investigated the dietary ecology of Macaca majori, an insular endemic species from Sardinia. In particular, we characterized the dental capabilities and potential dietary adaptations of M. majori through dental topographic and enamel thickness analyses of two M2s from the Early Pleistocene site of Capo Figari (1.8 Ma). We also assessed its diet through dental microwear texture analysis, while the microwear texture of M. majori was also compared with microwear textures from other European fossil macaques from mainland Europe. The dental topographic and enamel thickness analyses suggest that M. majori frequently consumes hard/mechanically challenging and/or abrasive foods. The results of the dental microwear analysis are consistent with this interpretation and further suggest that M. majori probably exhibited more durophagous dietary habits than mainland Plio-Pleistocene macaques. Overall, our results indicate that M. majori probably occupied a different dietary niche compared to its mainland fossil relatives, which suggests that they may have inhabited different paleoenvironments.

Morphology of the Bony Labyrinth Supports the Affinities of Paradolichopithecus with the Papionina.

Discoveries in recent decades indicate that the large papionin monkeys Paradolipopithecus and Procynocephalus are key members of the Late Pliocene - Early Pleistocene mammalian faunas of Eurasia. However, their taxonomical status, phylogenetic relationships, and ecological profile remain unclear. Here we investigate the two latter aspects through the study of the inner ear anatomy, as revealed by applying micro-CT scan imaging techniques on the cranium LGPUT DFN3-150 of Paradolichopithecus from the lower Pleistocene (2.3 Ma) fossil site Dafnero-3 in Northwestern Greece. Using geometric morphometric methods, we quantified shape variation and the allometric and phylogenetic signals in extant cercopithecines (n = 80), and explored the morphological affinities of the fossil specimen with extant taxa. LGPUT DFN3-150 has a large centroid size similar to that of baboons and their relatives. It shares several shape features with Macacina and Cercopithecini, which we interpret as probable retention of a primitive morphology. Overall, its inner ear morphology is more consistent with a stem Papionini more closely related to Papionina than Macacina, or to a basal crown Papionina. Our results, along with morphometrical and ecological features from previous studies, call into question the traditional hypothesis of a Paradolichopithecus-Macacina clade, and provide alternative perspectives in the study of Eurasian primate evolution during the late Neogene-Quaternary. Supplementary Information: The online version contains supplementary material available at 10.1007/s10764-022-00329-4.

Feeding ecology of the last European colobine monkey, Dolichopithecus ruscinensis.

Currently, very little is known about the ecology of extinct Eurasian cercopithecids. Dietary information is crucial in understanding the ecological adaptations and diversity of extinct cercopithecids and the evolution of this family. For example, the colobine genus Dolichopithecus is represented by multiple large-bodied species that inhabited Eurasia during the Pliocene-Early Pleistocene. The available evidence, though limited, suggests semiterrestrial locomotion, which contrasts with most extant African and Asian colobines that exhibit morphological and physiological adaptations for arboreality and folivory. These differences raise questions regarding the dietary specialization of early colobine taxa and how/if that influenced their dispersion out of Africa and into Eurasia. To further our understanding of the ecology of Plio-Pleistocene cercopithecids, we characterized the dental capabilities and potential dietary adaptations of Dolichopithecus ruscinensis through dental topographic and enamel thickness analyses on an M1 from the locality of Serrat d'en Vacquer, Perpignan (France). We also assessed the feeding behavior of D. ruscinensis through dental microwear texture analysis on a broad sample of fossil molars from fossil sites in France, Greece, Bulgaria, and Romania. Dental topographic and enamel thickness analyses suggest that D. ruscinensis could efficiently process a wide range of foods. Results of the dental microwear texture analysis suggest that its diet ranged from folivory to the consumption of more mechanically challenging foods. Collectively, this suggests a more opportunistic feeding behavior for Dolichopithecus than characteristic of most extant colobines.

The earliest known crown-Testudo tortoise from the late Miocene (Vallesian, 9 Ma) of Greece.

We here report on fossil remains of the earliest known crown-Testudo, an extant clade of Mediterranean testudinid tortoises from the late Miocene (Vallesian, MN 10) from the hominoid locality Ravin de la Pluie (RPl) in Greece. The material studied is a small, nearly complete carapace with a clearly distinct hypo-xiphiplastral hinge. This supports the sensu stricto generic assignment. This new terrestrial testudinid specimen is characterized by a possible tectiform, narrow, elongated shell with a pentagonal pygal and a long, posteriorly elevated, lenticular and rounded dorsal epiplastral lip. These unique features differ from those of other known Mediterranean hinged forms and allow the erection of the new species Testudo hellenica sp. nov. This taxon is phylogenetically close to two Greek species, the extant T. marginata and the fossil T. marmorum (Turolian, around 7.3 Ma). This record provides evidence for the first appearance of the genus Testudo sensu stricto at a minimum age of 9 Ma.

A 2Ma old baboon-like monkey from Northern Greece and new evidence to support the Paradolichopithecus - Procynocephalus synonymy (Primates: Cercopithecidae).

A new fossil cranium of a large papionin monkey from the Lower Pleistocene site of Dafnero-3 in Western Macedonia, Greece, is described by means of outer and inner morphological and metric traits using high-resolution micro-computed tomography. Comparisons with modern cercopithecids and contemporaneous Eurasian fossil taxa suggest that the new cranium could equally be ascribed to either the Eurasian Paradolichopithecus or to the East Asian Procynocephalus. The combination of the available direct and indirect fossil evidence, including the new cranium from Dafnero, revives an earlier hypothesis that considers these two sparsely documented genera as synonyms. The timing and possible causes of the rise and demise of Paradolichopithecus - Procynocephalus are discussed.

Stable isotope ecology of Miocene bovids from Northern Greece and the ape/monkey turnover in the Balkans.

Eurasia was home to a great radiation of hominoid primates during the Miocene. All were extinct by the end of the Miocene in Western Eurasia. Here, we investigate the hypothesis of climate and vegetation changes at a local scale when the cercopithecoid Mesopithecus replaced the hominoid Ouranopithecus along the Axios River, Greece. Because they are herbivorous and were much more abundant than primates, bovids are preferred to primates to study climate change in the Balkans as a cause of hominoid extinction. By measuring carbon stable isotope ratios of bovid enamel, we conclude that Ouranopithecus and Mesopithecus both evolved in pure C3 environments. However, the large range of δ(13)C values of apatite carbonate from bovids combined with their molar microwear and mesowear patterns preclude the presence of dense forested landscapes in northern Greece. Instead, these bovids evolved in rather open landscapes with abundant grasses in the herbaceous layer. Coldest monthly estimated temperatures were below 10°C and warmest monthly temperatures rose close to or above 20°C for the two time intervals. Oxygen isotope compositions of phosphate from bulk samples did not show significant differences between sites but did show between-species variation within each site. Different factors influence oxygen isotope composition in this context, including water provenience, feeding ecology, body mass, and rate of amelogenesis. We discuss this latter factor in regard to the high intra-tooth variations in δ(18)Op reflecting important amplitudes of seasonal variations in temperature. These estimations fit with paleobotanical data and differ slightly from estimations based on climate models. This study found no significant change in climate before and after the extinction of Ouranopithecus along the Axios River. However, strong seasonal variations with relatively cold winters were indicated, conditions quite usual for extant monkeys but unusual for great apes distributed today in inter-tropical regions.

Ruminant diets and the Miocene extinction of European great apes.

The successful evolutionary radiations of European hominoids and pliopithecoids came to an end during the Late Miocene. Using ruminant diets as environmental proxies, it becomes possible to detect variations in vegetation over time with the potential to explain fluctuations in primate diversity along a NW-SE European transect. Analysis shows that ruminants had diverse diets when primate diversity reached its peak, with more grazers in eastern Europe and more browsers farther west. After the drop in primate diversity, grazers accounted for a greater part of western and central European communities. Eastwards, the converse trend was evident with more browsing ruminants. These opposite trends indicate habitat loss and an increase in environmental uniformity that may have severely favoured the decline of primate diversity.