Elsevier

Quaternary Science Reviews

Volume 110, 15 February 2015, Pages 131-151
Quaternary Science Reviews

The remarkable Panthera pardus (Felidae, Mammalia) record from Equi (Massa, Italy): taphonomy, morphology, and paleoecology

https://doi.org/10.1016/j.quascirev.2014.12.020Get rights and content

Highlights

  • Equi represents the most complete and well preserved leopard collection in Europe.

  • We compare Equi Panthera pardus with both European fossils and modern leopards.

  • We provide a revaluated taxonomy of the of European leopards.

  • Sexual dimorphism in leopards is appreciable thanks skull features.

  • Tooth morphology differentiate Middle and Late Pleistocene subspecies.

Abstract

Leopard remains are rare in the European fossil record, probably a consequence of its solitary and elusive habits. Equi, dating back to Marine Isotope Stage 3 (MIS3), represents a rich and outstanding exception. Historical excavations (from 1911 to about 1917) at Equi allowed more than 200 remains to be recovered of this large felid, a sample attesting the richest leopard-bearing site in the Pleistocene of Europe. The Equi site testifies the survival of a prosperous leopard population in northwestern Italy (the Apuan Alps area) during the pre Last Glacial Maximum Late Pleistocene.

Almost all skeletal elements are represented in the collection (except ribs, and most of axial skeleton). At Equi, leopard is represented by both young and adult animals and, up to now, it seems the first and unique Pleistocene evidence of cave usage for cubs raised in Europe.

This study focuses on the whole record of leopard from Equi, housed in the Museum of Natural History of Florence, and is aimed to analyze the taphonomical assemblage, to provide an anatomical description of the remains and the comparison to the European evidences. Moreover, we infer the paleoecology of leopards, and their relationships with other large carnivores from the Equi guild (Ursus spelaeus, Canis lupus, and Panthera leo spelaea). The site was probably an occasional large carnivores den, used over time by different large carnivore taxa, perhaps in different seasons or times (mutual avoiding).

Our morphological analysis allows us to recognize sexual dimorphic characters in the cranium, and a large morphological variability in body size.

Section snippets

Foreword

Panthera pardus, the largest spotted cat in Africa and Asia, with several recognized subspecies, is a quite common, solitary, and opportunistic predator in the wild (Miththapala et al., 1996, Uphyrkina et al., 2001, Sunquist and Sunquist, 2002). It shows a great size variability within its range: generally speaking, in open country the animals are larger than in the forest, from 20 to 30 kg up to 90 Kg, with a large variability in food habits (Hayward et al., 2006). It can live in a wide range

The site of Equi

Equi is a small town on the northern side of the Apuane Alps (Fig. 1A–B). The topography of the area is characterized by dried environment and V-shaped fluvial valleys, even though some areas were interested by Late Pleistocene glaciation events. Evidence of small glaciers is spread most on the north-eastern side of the Apuane Alps (Braschi et al., 1986, Giraudi, 2004, Giraudi, 2011).

The Equi paleontological site is a cave and shelter located just out and above of the town, on the left side of

Materials and methods

This study focuses on the Equi leopard fossils recovered during the fieldwork by De Stefani and his team (1911–1917) and housed in the Natural History Museum, Geology and Paleontology Section, of the University of Florence (IGF), firstly described by Del Campana (1954) as P. pardus antiqua.

The Equi P. pardus collection consists in more than 200 bones (some of which are exposed in the Florence Natural History Museum permanent exhibition). Most of cranial and postcranial bones are almost

Taphonomic remarks

With a total number of 224 determined finds, Equi currently represents the most abundant collection of leopards across Europe, far exceeding the Late Pleistocene sites of Zafarraya (Barroso et al., 2003), Wildkirchli Cave (Switzerland), Baumann's Cave (Germany), the assemblage from Cueva de Los Rincones (Spain) and Caune de l’Arago and Hortus (France) (Testu, 2006, Testu et al., 2011, Diedrich, 2013, Saoqué et al., 2014a).

Adults are dominant (60% of the whole collection), followed by young

Systematic paleontology

  • Order Carnivora Bowdich, 1821.

  • Family Felidae Fischer Von Waldheim, 1817.

  • Subfamily Felinae Fischer Von Waldheim, 1817.

  • Genus Panthera Oken, 1816

  • P. pardus Linnæus, 1758

General overview

The site of Equi is a very rich deposit. The site has been known in literature for almost one century, but fossils and the available information about the original excavations were disconnected and never considered together. Until a recent re-evaluation analysis of the historical collection (Ghezzo et al., 2014b), the context was virtually ignored and a comparison with the new research (started since 2009) could not have been possible. On the basis of the available information from original

Conclusions

Leopards combine the power and strength of the larger cats with the grace and versatility of the smaller felinae.

For the first time from its discovery at the beginning of 19th century, the study of leopard collection from Equi allows for the definition of the morphological characterization, of a quite contemporaneous population (mid to late MIS3) within a singular context. Surprisingly, leopards from Equi were characterized by a great size variability both for cranial and limbs, maintaining

Acknowledgments

For the access to paleontological and osteological collections we are very grateful to the curators of the Natural History Museum of the University of Florence, Dr. Elisabetta Cioppi (Geological and Paleontological section) and Dr. Paolo Agnelli (Zoology section “La Specola”), the Natural History Museum of Berlin, Dr. Oliver Hampe (Vertebrate Paleontology section) and the Hungarian Natural History Museum, Prof. Mihály Gasparik (Department of Palaeontology and Geology), as well as to to Dr. P.

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