Dietary response of early Pleistocene ungulate communities to the climate oscillations of the Gelasian/Calabrian transition in Central Italy
Introduction
A gradual drop of the global temperatures characterized the late Pliocene, with seasonality and glacial activities increasing in the Northern Hemisphere (Flesche Kleiven et al., 2002). From this, the beginning of the Quaternary (around 2.6 Ma) corresponds to the initiation of the glacial/interglacial cycles marked by a 41 ka periodicity (Zubakov and Borzenkova, 1990; Lourens et al., 1996; Lisiecki and Raymo, 2005). These alternations exacerbated the climatic conditions with successive drop in temperature and humidity from the Gelasian (Kahlke et al., 2011; Head and Gibbard, 2015). In the Italian Peninsula, several cold and arid phases led to a type of vegetation (such as herbaceous plants and shrubs; mostly Artemisia and Ephedra) typical of open landscapes, while deciduous forests were developed during the more humid interglacial phases with a steady decline of subtropical taxa (Bertini, 2003; Fortelius et al., 2006; Kahlke et al., 2011; Bertini, 2013; Combourieu-Nebout et al., 2015).
Landscape alteration deeply influences mammal communities both in terms of composition (e.g., diversity) and ecology, having to develop new resource exploitation strategies to avoid extinction. This is especially true for large herbivores that, while having the ability to tolerate a range of unfavourable climate oscillations and palaeoenvironmental changes (e.g., from open landscapes to forests, from forests to wooded steppes), are highly susceptible to vegetation modification and their diets mirror the availability of plant resources (DeMiguel et al., 2010, DeMiguel et al., 2011). By investigating the feeding behaviour of large herbivores, it is thus possible to collect valuable information on how they adapted to, and cope with, alterations in habitat (Fortelius and Solounias, 2000; Solounias and Semprebon, 2002; Rivals and Athanassiou, 2008; Pushkina et al., 2014; DeMiguel, 2016; Bernor et al., 2017; Rivals et al., 2018).
According to the original definition of Azzaroli (1977), the Olivola Faunal Unit (FU) is the first one of the late Villafranchian placed at Gelasian/Calabrian boundary (at 1.8 Ma) (Torre et al., 1992) (Fig. 1A). As a result, this FU records the important changes in the large mammal faunal composition and diversity that occurred at the Gelasian/Calabrian transition and the passage from the middle to the late Villafranchian: the spread of large ungulates living in herds such as the leptobovine Leptobos etruscus, of two derived deer (Eucladoceros dicranios and Pseudodama nestii) and the appearance of the rare Caprinae Procamptoceras brivatense whose presence is registered only in the Olivola FU (Gliozzi et al., 1997; Rook and Martínez-Navarro, 2010). Among the carnivores, the first occurrence of the large hyena, Pachycrocuta brevirostris and the modern dog, Canis etruscus marks the spread of pack-hunter carnivores known as the “Wolf-event” (Azzaroli, 1983) or the “Pachycrocuta brevirostris event” due to the high-impact of this carrion eater on the early Pleistocene faunal assemblages (Sardella and Palombo, 2007; Palombo et al., 2008; Martínez-Navarro, 2010). Moreover, the early late Villafranchian also marks the arrival of the first Homo populations in the European continent, as testified by the exceptional hominid findings at Dmanisi (Georgia) (Gabunia and Vekua, 1995; Lordkipanidze et al., 2006 and references therein). The Olivola local fauna from northwest Tuscany (Aulla) is the most important site of the Olivola FU and, according to biochronological comparison with the nearby palaeomagnetically calibrated Matassino site, is probably located at the Olduvai Subchron (Gliozzi et al., 1997). Olivola faunal assemblage thus represents an excellent case study to investigate the effects of palaeoenvironmental alterations on large herbivores. By studying the dietary adaptations of the fossil ungulates of this site and by comparing the results with those obtained from the similar faunal assemblage of the middle Villafranchian locality of Coste San Giacomo (Anagni, Central Italy) (Strani et al., 2015, Strani et al., 2017), pivotal information can be gathered on the response of large ungulates in terms of resource exploitation to the climatic changes that occurred during the Gelasian/Calabrian transition.
Section snippets
Material and methods
The rich fossil fauna of Olivola comes from fluvial pelitic sandstone and conglomerate deposit (“Olivola Conglomerates”) underlying the Olivola village (Aulla, Tuscany) (Fig. 1B). The site is known from the 18th century and palaeontologists as Igino Cocchi and Giovanni Capellini, collected some specimens now stored in the palaeontological museums of the Universities of Pisa and Bologna. At the end of the 18th century the Olivola site was excavated by Forsyth Major (Forsyth Major, 1890)
Hypsodonty inference
The deer E. d. olivolanus display generally low molar crowns (Table 1) and has a brachydont dentition according to its HI for the lower third molar (HIm3 = 1.3) and a mesodont one for the upper or lower second molars (HIMm2 = 0.8). The other cervid of the assemblage, P. nestii, displays higher crowns and exhibits a mesodont dentition according to both indices (HIm3 = 1.6; HIMm2 = 0.8). The bovid L. etruscus also displays mesodont teeth (HIMm2 = 1.0) and S. etruscus has brachydont ones
Discussion
Hypsodonty is mostly associated to wear resistance with higher crowns indicating a higher level of abrasion in dry and open habitats (e.g. grasslands) (Janis, 1988; Eronen et al., 2010a, Eronen et al., 2010b), though it appears to be also linked to biomechanical effectiveness on consuming resistant items (DeMiguel et al., 2015). Considering these factors, the presence of one hypsodont, two mesodont, one brachydont and one brachy-mesodont ungulate seems to suggest a certain degree of aridity and
Conclusions
Analysis of the dental morphology and mesowear patterns of the Olivola ungulates allowed us to gather new information about the response of fossil ungulates to the Gelasian/Calabrian climatic changes, and on the habitats that characterized the region during this phase of significant environmental modifications. The occurrence of taxa with, mostly, medium-to-high molar crowns coupled with the lack of strict browsers, the abundance of mixed feeders (E. d. olivolanus, P. nestii, L. etruscus, P.
Acknowledgments
This work has been possible thanks to Fondi Dottorato di ricerca in Scienze della Terra – Sapienza, Università di Roma and to Ricerche Universitarie, Sapienza 2015 (ref. C26A153RNH to Raffaele Sardella). We also thank support from the AEI/FEDER EU (CGL2016-76431-P). Finally we'd like to acknowledge Elisabetta Cioppi (Natural History Museum of the University of Florence) for granting access to the Olivola collection.
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