Primates and Lagomorpha 3030340422, 9783030340421

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Handbook of the Mammals of Europe Series Editors: Klaus Hackländer · Frank E. Zachos

Klaus Hackländer · Paulo C. Alves  Editors

Primates and Lagomorpha

Handbook of the Mammals of Europe Series Editors Klaus Hackländer, University of Natural Resources and Life Sciences, Vienna, Department of Integrative Biology and Biodiversity Research, Institute of Wildlife Biology and Game Management, Vienna, Austria Deutsche Wildtier Stiftung (German Wildlife Foundation), Hamburg, Germany Frank E. Zachos, Natural History Museum Vienna, Mammal Collection, Vienna, Austria University of Vienna, Department of Evolutionary Biology, Vienna, Austria University of the Free State, Department of Genetics, Bloemfontein, South Africa

This handbook offers a unique collection of information on all wild mammal species living in Europe and will serve as a standard reference guide for all mammalogists and readers interested in research on mammals. The introductory volume provides an overview of mammalian phylogeny, the history and current status of European mammals and their management as well as their habitats and the history of mammalogy in Europe. The remaining volumes present comprehensive species-specific chapters covering all aspects of mammalian biology, including palaeontology, physiology, genetics, reproduction and development, ecology, habitat, diet, mortality and behaviour. The economic significance and management of mammals and future challenges for research and conservation are addressed as well. Each chapter includes a distribution map, a photograph of the animal, and a list of key literature. This authoritative handbook provides current and detailed descriptions of all European mammals; it will appeal to academics and students in mammal research alike, as well as to practitioners whose work involves mammal management, control, use and conservation.

Klaus Hackla¨nder • Paulo C. Alves Editors

Primates and Lagomorpha With 40 Figures and 17 Tables

Editors Klaus Hackländer University of Natural Resources and Life Sciences, Vienna, Department of Integrative Biology and Biodiversity Research, Institute of Wildlife Biology and Game Management Vienna, Austria

Paulo C. Alves Faculty of Sciences CIBIO-InBIO/BIOPOLIS University of Porto Vairão, Portugal

ISSN 2730-7387 ISSN 2730-7395 (electronic) Handbook of the Mammals of Europe ISBN 978-3-030-34042-1 ISBN 978-3-030-34043-8 (eBook) https://doi.org/10.1007/978-3-030-34043-8 © Springer Nature Switzerland AG 2023 This work is subject to copyright. All rights are reserved by the Publisher, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfilms or in any other physical way, and transmission or information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology now known or hereafter developed. The use of general descriptive names, registered names, trademarks, service marks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. The publisher, the authors, and the editors are safe to assume that the advice and information in this book are believed to be true and accurate at the date of publication. Neither the publisher nor the authors or the editors give a warranty, expressed or implied, with respect to the material contained herein or for any errors or omissions that may have been made. The publisher remains neutral with regard to jurisdictional claims in published maps and institutional affiliations. Cover photos with kind permission from © Ingo Arndt (young European hares), Bonaventura Majolo (Barbary macaque), Rolf Giger (European wild rabbit) This Springer imprint is published by the registered company Springer Nature Switzerland AG. The registered company address is: Gewerbestrasse 11, 6330 Cham, Switzerland

Acknowledgments

We are grateful to the editorial team of Springer Nature, especially to Barbara Wolf, Veronika Mang, Verena Penning, and Lars Koerner. In addition, we thank our volume editors and all species chapter authors for their support and contributions. Klaus Hackländer Frank Zachos Series Editors

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Contents

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Barbary Macaque Macaca sylvanus (Linnaeus, 1758) Bonaventura Majolo and Laëtitia Maréchal

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European Rabbit Oryctolagus cuniculus (Linnaeus, 1758) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Miguel Delibes-Mateos, Heiko G. Rödel, Carlos Rouco, Paulo C. Alves, Miguel Carneiro, and Rafael Villafuerte

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Eastern Cottontail Sylvilagus floridanus (J. A. Allen, 1890) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sandro Bertolino, David E. Brown, Jacopo Cerri, and John L. Koprowski

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Cape Hare Lepus capensis Linnaeus, 1758 . . . . . . . . . . . . . . . Massimo Scandura, Anna Maria De Marinis, and Antonio Canu

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Broom Hare Lepus castroviejoi Palacios, 1977 . . . . . . . . . . . . Fernando Ballesteros and Paulo C. Alves

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Corsican Hare Lepus corsicanus de Winton, 1898 . . . . . . . . . 111 Francesco Maria Angelici

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European Hare Lepus europaeus Pallas, 1778 . . . . . . . . . . . . 131 Klaus Hackländer

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Iberian Hare Lepus granatensis Rosenhauer, 1856 . . . . . . . . 167 Paulo C. Alves, Pelayo Acevedo, and José Melo-Ferreira

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Mountain Hare Lepus timidus Linnaeus, 1758 . . . . . . . . . . . . 191 Anders Angerbjörn and Stéphanie Claire Schai-Braun

Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 221

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About the Series Editors

Klaus Hackländer studied Zoology at PhilippsUniversity Marburg/Lahn and the University of Vienna. He is currently a Full Professor of Wildlife Biology and Wildlife Management at the University of Natural Resources and Life Sciences, Vienna (BOKU), where he also coordinates the master program on Wildlife Ecology and Wildlife Management. Prof. Hackländer is a member of numerous mammalogical societies and served as President of the German Society for Mammalian Biology from 2009 to 2016. He was Editor-inChief of the journal Mammal Review from 2009 to 2015. Prof. Hackländer’s research covers all aspects of mammalian biology, including ecology, behavior, physiology, diseases, and management. Since 2021, he is Head of the German Wildlife Foundation (Deutsche Wildtier Stiftung), besides being Head of the Institute of Wildlife Biology and Game Management at BOKU.

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About the Series Editors

Frank E. Zachos is Head of the Mammal Collection at the Natural History Museum in Vienna, Austria, an Affiliated Lecturer at the University of Vienna, and an Affiliated Professor at the Department of Genetics, University of the Free State in Bloemfontein, South Africa. He studied biology, the history of science, and philosophy in Kiel and Jena, Germany, and received his Ph.D. in Zoology in 2005. In 2009, he completed his habilitation (postdoctoral degree) in Zoology and Evolutionary Biology. His main research interests are intraspecific biodiversity, phylogeography, population genetics, and conservation of mammals and birds, with a particular focus on red deer and other ungulates. He also has a long-standing interest in species concepts and the species problem, as well as other theoretical and philosophical issues in evolutionary biology and systematics. Until 2021, he was Editor-in-Chief of the journal Mammalian Biology, and he is currently the president of the German Society for Mammalian Biology and the Editor of the Mammalia series within the Handbook of Zoology.

About the Volume Editors

Klaus Hackländer’s favorite mammalian taxon is Lagomorpha. He studied leporids in Europe, North America, and Africa. Together with the co-editor of this volume, he founded the World Lagomorph Society. Prof. Hackländer serves in the Lagomorph Specialist Group of the IUCN Species Survival Commission and contributed to the global Red List Assessment of Lepus europaeus. In addition, he is responsible for the European Red List Assessment of leporids. His publications include the leporid chapter of the Handbook of Mammals of the World, two books on lagomorphs, and numerous original work, mainly on L. europaeus and L. timidus. Paulo Célio Alves is an Associate Professor of Biology at the Faculty of Sciences, University of Porto and the Head of the Conservation Genetics and Wildlife Management research group at CIBIO (Research Center in Biodiversity and Genetics Resources/BIOPOLIS). Editor of the European Journal of Wildlife Research, author of more than 200 publications, including papers, chapters, and books, namely two co-edited books on lagomorphs. Cofounder and current President of the World Lagomorph Society. Currently is the Director of recently created Biological Station of Mértola, Portugal. His main research topics cover animal ecology, wildlife management, conservation biology, and conservation genetics.

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Contributors

Pelayo Acevedo Instituto de Investigación en Recursos Cinegéticos IREC (CSIC-UCLM-JCCM), Ciudad Real, Spain Paulo C. Alves CIBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, InBIO, Universidade do Porto, Vairão, Portugal BIOPOLIS Program in Genomics, Biodiversity and Land Planning, CIBIO, Vairão, Portugal Departamento de Biologia, Faculdade de Ciências, Universidade do Porto, Porto, Portugal Estação Biológica de Mértola (EBM), Mértola, Portugal Francesco Maria Angelici Fondazione Italiana per la Zoologia dei Vertebrati (FIZV), Rome, Italy Anders Angerbjörn Department of Zoology, Stockholm University, Stockholm, Sweden Fernando Ballesteros Fundación Oso Pardo (Brown Bear Foundation), Santander, Spain Sandro Bertolino Department of Life Sciences and Systems Biology, University of Turin, Torino, Italy David E. Brown Natural History Collections, Arizona State University, Tempe, AZ, USA Antonio Canu Department of Veterinary Medicine, University of Sassari, Sassari, Italy Miguel Carneiro CIBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, InBIO, Universidade do Porto, Vairão, Portugal BIOPOLIS Program in Genomics, Biodiversity and Land Planning, CIBIO, Vairão, Portugal Jacopo Cerri Institute for Environmental Protection and Research, Livorno, Italy

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Anna Maria De Marinis Italian Institute for Environmental Protection and Research (ISPRA), Ozzano dell’Emilia, Italy Miguel Delibes-Mateos Instituto de Estudios Sociales Avanzados (IESACSIC), Córdoba, Spain Klaus Hackländer University of Natural Resources and Life Sciences, Vienna, Department of Integrative Biology and Biodiversity Research, Institute of Wildlife Biology and Game Management, Vienna, Austria Deutsche Wildtier Stiftung (German Wildlife Foundation), Hamburg, Germany John L. Koprowski Haub School of Environment & Natural Resources, University of Wyoming, Laramie, WY, USA Bonaventura Majolo School of Psychology, University of Lincoln, Lincoln, UK Laëtitia Maréchal School of Psychology, University of Lincoln, Lincoln, UK José Melo-Ferreira Departamento de Biologia, Faculdade de Ciências da Universidade do Porto, Porto, Portugal CIBIO/InBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, Universidade do Porto, Vila do Conde, Portugal BIOPOLIS Program in Genomics, Biodiversity and Land Planning, CIBIO, Vairão, Portugal Heiko G. Rödel Laboratoire d’Ethologie Expérimentale et Comparée (LEEC), UR4443, Université Sorbonne Paris Nord, Villetaneuse, France Carlos Rouco Área de Ecología, Departamento de Botánica, Ecología y Fisiología Vegetal, Universidad de Córdoba, Córdoba, Spain Sociedad, Ecología y Gestión del Medio Ambiente, UCO-IESA, Unidad Asociada al CSIC, Córdoba, Spain Massimo Scandura Department of Veterinary Medicine, University of Sassari, Sassari, Italy Stéphanie Claire Schai-Braun Department of Integrative Biology and Biodiversity Research, Institute of Wildlife Biology and Game Management, University of Natural Resources and Life Sciences, Vienna, Vienna, Austria Rafael Villafuerte Instituto de Estudios Sociales Avanzados (IESA-CSIC), Córdoba, Spain

Contributors

Barbary Macaque Macaca sylvanus (Linnaeus, 1758)

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Bonaventura Majolo and Lae¨titia Mare´chal

Contents Common Names . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

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Taxonomy and Systematics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

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Paleontology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

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Current Distribution . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

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Description . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

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Physiology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

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Genetics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

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Life History . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

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Habitat and Diet . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

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Behavior . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 Parasites and Diseases . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 Population Ecology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 Conservation Status . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 Management . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 Future Challenges for Research and Management . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22

B. Majolo (*) · L. Maréchal School of Psychology, University of Lincoln, Lincoln, UK e-mail: [email protected]; [email protected] © Springer Nature Switzerland AG 2023 K. Hackländer, P. C. Alves (eds.), Primates and Lagomorpha, Handbook of the Mammals of Europe, https://doi.org/10.1007/978-3-030-34043-8_15

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B. Majolo and L. Mare´chal

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Common Names English German French Spanish Italian Russian

Barbary macaque (also wrongly called Barbary ape) Berberaffe, Magot Magot, singe magot, macaque Berbère Mona de Berberíc, mona/mono de Gibraltar Bertuccia, macaco Berbero Берберийская макака

Taxonomy and Systematics The Barbary macaque belongs to the genus Macaca comprising approximately 25 species (Li et al. 2009). It is the only nonhuman primate in Europe, the northernmost African nonhuman primate and the only extant species of the genus Macaca outside East-Asia. The genus Macaca probably originated in North Africa around 5.5 million years ago (Modolo et al. 2005), and then radiated into Europe and Asia (Li et al. 2009). However, extant macaque species all live in East Asia, with the exception of the Barbary macaque, between Afghanistan and Pakistan to the East and Japan to the West, and between China to the North and Indonesia to the South. By the early Pleistocene the Barbary macaque lineage disappeared in most of Europe and in the Middle East and as a result became separated from the other macaque lineages. The reduction in the distribution of the Barbary macaque was probably due to climate change and associated changes in vegetation and availability of resources (Elton and O’Regan 2014). Due to the geographic distance between the Barbary macaque and other macaque species, there is no record of hybridization with related species in the wild. The Barbary macaque population living in Gibraltar has both Algerian and Moroccan origins (Modolo et al. 2005). The origins of the Gibraltar population are unclear: it may have been originated from a remnant European population or it may have been established through the translocation of some animals from Africa (Modolo et al. 2005). In any case, the current Gibraltar

population might not include any remnants of the original European population, since there are no fossil records of Barbary macaques in Gibraltar after the last glaciation (Shaw and Cortes 2006).

Paleontology The fossil record and the modeled reconstruction of their geographic distribution suggest that the Barbary macaque inhabited vast parts of Europe, North-Africa, and the Middle East (Syria and the Caucasian region) between the Miocene and Pleistocene (Fig. 1; Elton and O’Regan 2014). Humans have probably affected the geographic range of the Barbary macaques since at least the last 4,000 years (Modolo et al. 2005). In historic times, the Barbary macaque faced a dramatic decline in its distribution. In Europe, this species was restricted to Spain by the end of the nineteenth century and it was only found in Gibraltar by the beginning of the twentieth century (Taub 1984).

Current Distribution The current distribution of the Barbary macaque is limited to Gibraltar (36°09’N, 05°21’W) and to small relic patches of forest in Morocco and Algeria (Fig. 2). In Gibraltar, around six groups of macaques, for a total of approximately 200 individuals, inhabit the Upper Rock Nature Reserve; some of these groups rely extensively of food provisioning from humans.

Description The Barbary macaque is often wrongly called the tailless monkey or Barbary ape. The Barbary macaque is not an ape (i.e., anthropoid primate) and it has a vestigial tail, which varies in length from 4 mm to 22 mm (Fooden 2007). Sexual dimorphism can be observed in relation to body length (males: 550–600 mm, females: 450 mm)

Barbary Macaque Macaca sylvanus (Linnaeus, 1758)

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Fig. 1 Location of Macaca fossils in Europe, North Africa and Caucasus from (a) the Miocene to the EarlyMiddle Pleistocene boundary and (b) in the Middle and

Late Pleistocene. Note that not all Miocene Macaca sites in North Africa are shown in the figure. (Redrawn from Elton and O’Regan (2014) with permission from Elsevier)

and weight (males: 15–17 kg, females: 10–11 kg; Fa 1989). The body size and morphology of Barbary macaques varies depending on age and on whether measurements are taken from provisioned or non-provisioned individuals (Fa 1984; Fooden 2007; Borg et al. 2014; Table 1 and Fig. 3). Provisioned macaques have a significantly larger body size than non-provisioned animals (Borg et al. 2014; Maréchal et al. 2016a). The dental formula of a Barbary macaque is I2/2, C1/1, P2/2, M3/3. Sub-adult and adult males

have long canine teeth, surpassing their incisors, which play an important role in dominancerelated behaviors, while females do not display long canine teeth. During the mating period (see section “Genetics” below), female sexual displays are characterized by a sexual swelling (Fig. 4). This swelling varies during the reproductive cycle of a female (Brauch et al. 2007; Young et al. 2013a) but, in provisioned conditions, a swelling might be still visible outside of the mating season despite the

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B. Majolo and L. Mare´chal

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Map template: © Getty Images/iStockphoto

Fig. 2 Current geographic distribution of the Barbary macaque. The distribution is based on the IUCN Red List of Threatened Species. Version 2020-2 (Wallis et al. 2020;

redrawn with permission from IUCN). (Map template: © Getty Images/iStockphoto)

female not being fertile at the time. There is no sexual dimorphism in the pelage color in the Barbary macaque. In adult and sub-adult Barbary macaques, the pelage coloration on the dorsal surface from the top of the head to the hindquarter is highly variable from light gray, beige, light brown, golden brown, burnt orange, to dark (Fig. 3; Fooden 2007). The ventral fur is generally lighter than the dorsal fur, and it is also not as thick, leaving ventral body parts often visible. To cope with extreme climatic conditions, with hot and dry summer (up to +40 °C) and cold winter (up to
20 °C), Barbary macaques have a thicker fur during autumn and winter, which molts during spring and summer (Fig. 3). Age determination is difficult in the Barbary macaques, especially at older age, because the

different traits used to age a macaque (e.g., dentition, pelage, locomotion, social behavior; Table 2) may significantly vary across individuals of the same age due to factors like dominance rank, diet, and feeding efficiency. In captivity or under provisioned conditions, Barbary macaques may live up to around 20–28 years; the maximum recorded lifespan of a Barbary macaque is 28 years, in captivity (Sue Wiper pers. comm.; Fooden 2007). Macaques are unlikely to live beyond 20 years in the wild in non-provisioned conditions. New noninvasive markers for immune function, urinary neopterin levels, have been recently found to significantly increase with age (Müller et al. 2017), which may provide a useful additional tool to assess age in wild individuals in the future.

Table 1 Body measurements of Barbary macaque by age/sex class Age/sex class Infants 5 hares/km2. In the hunting season 2012/2013, a total harvest of 4890 hares was reported by hunters in Sardinia (91% of data coverage). With the aim to increase local densities, areas are specifically set to favor the natural reproduction of Cape hare, wild rabbit, and barbary partridge (named “Zone Temporanee di Ripopolamento e Cattura”). In these areas, hunting is forbidden and annual activities (population counts, habitat improvements, etc.) are carried out to increase the productivity and natural dispersal to surrounding areas and to define a number of animals that could be captured and transferred to external depleted areas. Unfortunately, though existing in good numbers in Sardinia, these areas are seldom managed in a proper way and the conditions for restocking with wild-captured heads are persistently lacking. Cape hares are bred in captivity in a very limited number of public and private farms.

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Breeding protocols used for the L. europaeus (L. Mandas, pers. comm.) were initially applied but resulted ineffective and were progressively adapted to the biological requirements of the Cape hare. Nonetheless, a few releases of captive-reared hares have taken place in the past. The interest towards this species makes it a valuable resource. Actually, possible economic implications in Sardinia are constrained by a number of factors, namely: (1) poor management of the species, (2) prohibition of selling the meat of hunted hares, (3) prohibition of exporting Cape hares to the mainland, (4) low number of breeding farms, and (5) restrictions to the release of captivereared hares. Therefore, the only economic repercussions are limited to those linked to a naturalistic enjoyment or to the hunting activity (weapons, equipment, hunting dogs, hunting tourism). The species in Sardinia has a relatively low impact on human activities. Damages to crops are negligible, as well as those caused by traffic accidents. Nevertheless, some economically important diseases can be transmitted to livestock by tick infesting hares (see section “Parasites and Diseases”).

Future Challenges for Research and Management As remarked by many authors (e.g., Smith et al. 2018), knowledge on the Cape hare is largely incomplete. Notwithstanding the insights provided by recent studies, taxonomic uncertainties remain that would take advantage from new comprehensive studies on the phylogeography of the L. capensis complex. They would deserve a targeted sampling in some areas that have not been sufficiently considered in previous investigations (e.g., sub-Saharan and East African populations). As concerns the European population of Cape hares, researches on its ecology, behavior, and population dynamics urge, so as to assist a conservative management of the Sardinian population. In particular, it will be crucial to focus on:

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• Factors that are responsible of the low densities observed in many areas of Sardinia (even in presence of a high habitat suitability) • Relative contribution of the different mortality factors in free-living populations • Spatial behavior (especially habitat selection and dispersal patterns) • Food habits in different ecological conditions (agricultural landscape, Mediterranean shrubland, high-elevation pastures) Hunting needs to be reorganized to make possible the definition of sustainable annual quotas, on the basis of local abundances and recruitment rates estimated annually on the basis of standardized protocols. Hunting bag data must be regularly collected and analyzed, in order to provide information on relevant population parameters (sex and juvenile/adult ratios). Measures should be undertaken to favor natural reproduction and dispersal (e.g., by an efficient management of protected areas and the “Zone Temporanee di Ripopolamento e Cattura”), yet short-range (e.g., 37% of fecal pellets). In the other seasons, hares also included fruits, e.g., Prunus spinosa, Pyrus piraster, Malus sylvestris), which, in autumn were detectable in >27% of the analyzed droppings. There were significant differences among seasons. The smallest values of richness and diversity were observed in spring. Dietary overlap was low between spring and the other seasons; conversely, there was substantial overlap (>70% of fecal pellets) in the diets during the other seasons with a more pronounced similarity between summer and autumn (Freschi et al. 2014b). Finally Freschi et al. (2015) analyzed in detail the seasonal variation of the Apennine hare diet. In this study, the seasonal pattern of diet composition of a population of L. corsicanus occupying a semi-natural landscape was estimated by using the micro-histological technique of fecal analysis. The results showed that hares

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had a diversified diet, consuming plant parts from over 70 species (Freschi et al. 2015). Like other Lepus species, the Apennine hare consumed a large amount of herbaceous plants, e.g., Brachypodium sylvaticum, Trifolium pratense, Allium subhirsutum, and Festuca arundinacea, although it complemented its diet seasonally with fruits of Prunus spinosa, Pyrus piraster, and Malus sylvestris. Analysis of similarities (ANOSIM) evidenced significant differences among seasons, as a consequence of the seasonal occurrence of the various food items. Spring and autumn, as well as spring and winter, showed low diet similarities; these results were supported by similarity percentage analysis (> 71% between spring and autumn; > 69% between spring and winter) with taxa like P. spinosa, Cirsium strictus, T. pratense, and Rosa canina making the greatest contributions to these differences. Higher similarities were instead found when comparing other seasons. A study on feeding habits was carried out in Corsica (Rizzardini et al. 2019). This study is the first to consider the food selection of L. corsicanus. Grasses represented the basis of the diet, with frequencies around 50%, followed by non-leguminous forbes with an incidence of 29 to 31%. Leguminous shrubs complemented its diet. Poaceae were found to be the most preferred and selected family in the diet. A total of 79 species were detected in the diet, but only some of them had percentages higher than 5%. The most used species in the diet were Brachypodium sylvaticum, Briza max, Trifolium angustifolium, Digitaria sanguinalis, and Daucus carota. This study showed that in the areas considered, characterized also in the dry period by wide plant diversity, the Apennine hare used plants as a generalist (Rizzardini et al. 2019). Buglione et al. (2015) showed, in a mountain environment in Campania (Southern Italy), that there is a 26% food overlap (competition) between L. corsicanus and L. europaeus on the used plant species. These authors also showed how this mechanism can be negative and dangerous for the survival of L. corsicanus following recent climate changes, due to the lower food adaptability. That is, with the gradual

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increase in temperature, some taxa of mountain plants, many of which are specifically preferred by the Apennine hare, could likely face extinction. This could be strongly negative for L. corsicanus, a species already stressed and relegated to marginal environments (Buglione et al. 2015). Buglione et al. (2018) presented a non-invasive pilot study set to analyze the diet composition of the Apennine hare from Southern Italy, starting from fecal pellets, using, for the first time on this species, DNA metabarcoding and next generation sequencing to identify the plants. The findings of this study indicated that this genetic approach provides reliable qualitative and semi-quantitative information, allowing the characterization of the hare diet and its seasonal variation using 22 fecal samples. In a single experiment, through time and cost-effective screening of multiple DNA metabarcodes, Buglione et al. (2018) detected a broad diversity of plants (99 taxa). Unlike traditional methods, this approach can identify items that leave no solid remains or that simply are lacking in diagnostic taxonomic features.

Behavior As for the use of space and territoriality, the species seems to be rather sedentary with home ranges of limited size; from sunset and for most of the night it frequents the same pasture areas, which are located close to the night shelters (M. Lo Valvo, pers. comm.). In a study in Sicily conducted by Lo Valvo (2007), it was able to ascertain how the annual home ranges detected on six individuals vary from 0.7 to 267.7 ha, using the method of the Convex Minimum Polygon (MCP). Some preliminary data collected on a small sample of Apennine hare in Latium (Central Italy), using the 95% Kernel method, have determined annual home range extension data in two females ranging from 8 to 28 ha, while in one male the home range varied from 59 to 73 ha (Di Luzio and Barone 2012). Some Apennine hares reintroduced on the Island of Elba (Scarselli et al. 2016; Lari 2019) and monitored with GPS

F. M. Angelici

technology have occupied an average home range varying between 87.33 and 97.41 ha. Mori et al. (2020) in a multi-year study in Central Italy have shown that the home range (calculated both with the 95% MCP and with the 95% Kernel) is larger, both in males and females, during the warm months compared to winter. Furthermore, home ranges were larger during the night (hares more active for foraging) compared to daylight. In particular, in the warmer months the home ranges varied from about 40 to 70 ha, to decrease to a range of around 27 to 45 ha in the colder months; there were no significant differences between the two sexes (Mori et al. 2020). The spatial behavior of seven male and five female Apennine hares was studied in Central Italy, through radiotracking data (Lovari et al. 2020). The median percentage of home range overlap was about five times lower in females (