Felines of the World: Discoveries in Taxonomic Classification and History

By Giovanni G. Bellani

Felines of the World: Discoveries in Taxonomic Classification and History provides the most recent taxonomic, paleontological, phylogenetic and DNA advances of wild felid and domestic cat species following guidelines dictated by the IUCN SSC Cat Specialist Group. It highlights the importance of felines and their role as predators in maintaining the ecological biome balance in which they have evolved. The book delves into the anatomical, evolutionary and zoogeographic features of fossil and current felid species. Each species is described in detail, detailing its classification, habitat and biological habits. This book also presents the most updated threat and conservation status of each species.

This book is an ideal resource for zoologists and paleontologists, primarily those interested in the evolution and features of extinct and extant felines.

• Details the lineage, features and habits of over 40 felid species
• Covers all species within the Felidae family, including lions, lynxes, pumas and domestic cats
• Features detailed and colorful illustrations, diagrams and species location maps
• Informs readers on endangered species, current threats and conservation efforts

• Language: English
• Publisher: Academic Press
• Release date: Sep 19, 2019
• ISBN: 9780128172773

Giovanni G. Bellani

Dr. Giovanni Giuseppe Bellani is a zoology consultant for some of the most prestigious natural history museums in Italy, including his recent past affiliation with the Natural History Museum “Faraggiana Ferrandi” of Novara, where he organized the classification of the collections and the restructuring, planning, and following production of the new permanent exhibitions. He received his Ph.D. in natural sciences from the University of Pavia. He is a member of the Scientific Committee for the Design of Watercourses and participates in an exhibition of the Ecomuseum of the Sesia River Valley, which is part of the Piedmontese ecomuseum project. Dr. Bellani has published four books on natural sciences and has collaborated with two other books and on numerous scientific journal publications on zoology, museology, conservation and environmental sciences.

Book preview

Felines of the World

Discoveries in Taxonomic Classification and History

Giovanni Giuseppe Bellani

Zoology Consultant and Museologist, Natural History Museums, Italy

Table of Contents

Cover image

Title page

Copyright

Acknowledgments

Chapter 1. Order of carnivores (Carnivora)

Abstract

Introduction

Man and carnivores

Chapter 1.1 Anatomical features common to order Carnivora

Chapter 1.2 Features of Caniformia and Feliformia

Carnivora denture or dentition

Chapter 1.3 Evolution, phylogeny, and classification

Chapter 2. Family Felidae

Abstract

Chapter 2.1 Classification

Classification of the Felidae family

Chapter 2.2 Evolution

Chapter 2.2.1 The origins: the subfamily Proailurinae

Chapter 2.3 Anatomical features of current Felids

Skeleton and muscles

The skeleton (skeletal system)

The legs and retractable claws

The skull

The function of the teeth

Musculature (muscular system)

Cats always land on their feet

The fur

Furs: patterns, colors, and habitat

Fur as a protection from cold

Chapter 3. Zoogeography of felids

Abstract

3.1 The zoogeographical regions

South-Eastern Asia: a biodiversity hot spot for safeguarding some rare species of felids

The Felids of the Indochinese Subregion and Sundaic Subregion

New species in the Sunda Archipelago

The great eruption of the Toba volcano in Sumatra

3.2 The origins of the current distribution of felids around the world

Part 1: Gallery of All Species of Felidae

Chapter 4. Subfamily Pantherinae

Abstract

Chapter 4.1 Evolutionary line of the Pantherinae subfamily and lineage Panthera

Biology

Biology

Biology

Biology

Biology

Biology

Biology

Chapter 5. Subfamily Felinae

Subfamily Felinae

Chapter 5.1. Bay cat lineage

Chapter 5.1.1 Borneo bay cat (Catopuma badia) Gray, 1874

Chapter 5.1.2 Asian golden cat or Temminck’s cat (Catopuma temminckii) Vigors and Horsfield, 1827

Chapter 5.1.3 Marbled cat (Pardofelis marmorata) Martin, 1836

Chapter 5.2. Caracal lineage

Chapter 5.2.1 Serval (Caracal=Leptailurus serval) (Schreber, 1776)

Chapter 5.2.2 Caracal (Caracal caracal) (Schreber, 1776)

Chapter 5.2.3 African golden cat (Caracal aurata) (Temminck, 1827)

Chapter 5.3. Ocelot lineage

Chapter 5.3.1 Ocelot (Leopardus pardalis) Linnaeus 1728

Chapter 5.3.2 Margay Leopardus wiedii (Schinz, 1821)

Chapter (5.3.3/5.3.4) Oncilla or tiger cat or tigrina (superspecies) Leopardus tigrinus, Leopardus guttulus (and proposed Leopardus emilae)

Chapter 5.3.3 Northern oncilla or northern tigrina Leopardus tigrinus (Schreber, 1775)

Chapter 5.3.4 Southern oncilla or southern tigrina Leopardus guttulus (Hensel, 1872)

Chapter 5.3.5 Geoffroy’s cat Leopardus geoffroyi (d’Orbigny and Gervais, 1844)

Chapter 5.3.6 Guigna or Guiña Leopardus guigna (Molina, 1782)

Chapter 5.3.7 Colocola or pampas cat Leopardus colocola (Molina, 1782)

Chapter 5.3.8 Andean cat Leopardus jacobita

Chapter 5.4. Lynx lineage

Chapter 5.4.1 Eurasian lynx (Lynx lynx) (Linnaeus, 1758)

Chapter 5.4.2 Canadian lynx (Lynx canadensis) (Kerr, 1792)

Chapter 5.4.3 Iberian lynx (Lynx pardinus) (Temminck, 1827)

Chapter 5.4.4 Bobcat or red lynx (Lynx rufus) (Schreber, 1777)

Chapter 5.5. Puma lineage

Chapter 5.5.1 Jaguarundi or eyra Puma (Herpailurus) yagouaroundi (É. Geoffroy Saint-Hilaire, 1803)

Chapter 5.5.2 Puma, mountain lion, or cougar Puma concolor (Linnaeus, 1771)

Chapter 5.5.3 Cheetah or hunting leopard Acinonyx jubatus (Schreber, 1775)

Chapter 5.6. Asian leopard cat lineage

Chapter 5.6.1 Asian leopard cat (Prionailurus bengalensis) (Kerr, 1792)

Chapter 5.6.2 Sunda leopard cat Prionailurus javanensis (Desmarest, 1816)

Chapter 5.6.3 Fishing Cat (Prionailurus viverrinus) (Bennett, 1833)

Chapter 5.6.4 Flat-headed cat (Prionailurus planiceps) (Vigors and Horsfield, 1827)

Chapter 5.6.5 Rusty-spotted cat (Prionailurus rubiginosus) (I. Geoffroy Saint-Hilaire, 1831)

Chapter 5.6.6 Manul or Pallas cat (Otocolobus manul) Pallas, 1776

Chapter 5.7. Domestic cat (Felis) lineage

Chapter 5.7.1 European wildcat (Felis silvestris) Schreber, 1777

Chapter 5.7.2 African and Asian Wildcat (Felis lybica) Forster, 1780

Chapter 5.7.3 Chinese mountain cat or Chinese alpine steppe cat (Felis bieti) Milne-Edwards, 1892

Chapter 5.7.4 Sand cat or Sand Dune cat (Felis margarita) Loche, 1858

Chapter 5.7.5 Black-footed cat (Felis nigripes) Burchell, 1824

Chapter 5.7.6 Jungle cat or swamp cat (Felis chaus) Schreber, 1777

Chapter 5.7.7 Domestic cat (Felis catus) Linnaeus 1728

Chapter 6. Ecology and predation

Abstract

Chapter 6.1 Population ecology

The role of feline predation in the ecosystem

Ecological or food pyramid (biomass)

Predators maintain the plant–herbivore balance

Puma and mule deer in Kaibab’s highland (Arizona)

Felines and the discovery of the dynamic ecological balance: numerical fluctuations

The case of the Canadian lynx and the snowshoe hare

Chapter 6.2 Predation: hunting methods and techniques

Felids: the most skilled world’s predatory carnivores

Relationship between dimensions of the predator and the prey

The fatal bite

Felines: not only solitary ambushes

Serval (Leptailurus serval)

Caracal (Caracal caracal)

Chapter 7. Ethology

Abstract

Chapter 7.1. Intraspecific communication

Chapter 7.1.1 The organs of senses

The view: the eyes

Night vision

The tapetum lucidum

Vertical pupil and round pupil

The binocular or stereoscopic vision

Sound

The ears: the sense of hearing

Smell: the nose

Vomeronasal or Jacobson’s organ

The tongue: taste

Chapter 7.1.2 Communication signals: visual, olfactory, auditory

Visual communication

Olfactory communication and scent markings

Auditory communication

Chapter 7.1.3 Pantherinae and Felinae: the roar and the purr

The latest discoveries on the anatomy of the vocal system of Felidae

What causes the roar

The purr

Chapter 7.2. Social and territorial behavior

Territorial range

Chapter 7.2.1 Reproductive behavior

Chapter 7.2.2 Parental care

Chapter 8. Genetics

Abstract

Genetic variability

Chapter 8.1 DNA, chromosomes, genes, genotypes, and phenotypes

Heredity traits

Dominant and recessive Alleles

Genetic mutations

Molecular clock

Melanism in the Jaguar

Chapter 8.1.2 Genetics of some colors of the domestic cat

Spontaneous mutations in the color of the hair: albinism

The white lions of Timbavati and Kruger

Chapter 8.1.3 Crossing between the species of great felines (Pantherinae)

Chapter 9. Felidae and man

Abstract

Chapter 9.1. Felids in human culture

Chapter 9.1.1 Felidae in art

The rocky art of the Paleolithic

The first Mediterranean civilizations

Egyptian civilization

Assyrian-Babylonian civilization

Greek and Roman art

Animals of the mosaics of Villa Del Casale Di Piazza Armerina (Enna, Sicily, Italy)

The Middle Ages

Medieval bestiaries (books of beasts)

The Renaissance

The birth of Naturalism in art

The cat and other felides in the art of the 20th century

Franz Marc (1880–1916)

The Novecento painters and their cats

Antonio Ligabue (Antonio Laccabue) (1899–1965)

The great illustrators of the end 20th and start of the 21st centuries

Robert Bateman—Toronto, 1930

Walty (Walty Dudok Van Heel) Hilversum—Holland, 1959–2009

ROBERT DALLET (1923, près d'Etretat, Normandy—2006)

Chapter 9.1.2 Felidae in literature: the Lonza the mysterious feline of Dante Alighieri in the Divine Comedy

Chapter 9.1.3 Felidae in the cinema and cartoons

Films of other production companies, cartoons, and strips

Films

Chapter 9.1.4 Joy and George Adamson: the History of the lioness Elsa

Chapter 9.2. History of taming of the felidae and domestication of the cat

The ancestor of the domestic cat

Chapter 9.2.1 The domestic cat in Ancient Egypt: the goddess Bastet and cat mummies

Chapter 10. Conservation of felidae

Abstract

10.1 Situation of felid species in the wildlife

The causes of the decline to stop

Other causes of the rarity of many species of felids

Bibliography

Periodic scientific journal

Book

Index

Copyright

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Acknowledgments

I would like to thank all the people who provided a kind help, answering my questions, giving me valuable advice, and giving permission to publish diagrams, graphs, maps, and drawings of their scientific publications or the logos of their associations.

✝Groves P. Colin (1942–2017) Prof., Emeritus School of Archeology and Anthropology, Australian National University, Acton, Australia

Anton Mauricio Prof., Curator of Paleontology, Natural History Museum of Madrid

Barbuti Roberto Prof., Direttore del Museo di Storia Naturale dell’ Universita di Pisa (Calci)

Barnett Ross, Centre For Geogenetics, København Universitet, The Natural History Museum, Øster Voldgade, Copenhagen

Batemann Robert Mr., Naturalist and Wildlife Artist

Brotchie Kate Ms., Assistant to Robert Bateman

Boulton Mark Dr., Elsa Conservation Trust, Naivasha Kenya

Brown Jeffrey Dr., Anatomic and Clinical Pathology, Independent Researcher, New York, United States

Campos Trigo Tatiane Prof., Biologa – Setor de Mastozoologia Museu de Ciencias Naturais Fundacao Zoobotanica do Rio Grande do Sul

Cherin Marco Prof. Paleontologia dei Vertebrati Universita’ di Perugia

Chimento R. Nicolas Dr., Laboratorio de Anatomía Comparada y Evolución de Los Vertebrados, Museo Argentino de Ciencias Naturales Bernardino Rivadavia, Buenos Aires, Argentina

Cooper Alan Prof., School of Earth and Environmental Sciences, University of Adelaide, Adelaide, Australia

Cooper David Prof. PHD, The University of Edinburgh, School of Geosciences

Dondas Alejandro Dr., Museo Municipal de Ciencias Naturales, Lorenzo Scaglia Mar del Plata, Argentina

Eizirik Eduardo Prof., Coordenador–PPG de Faculdade de Biociencias, PUCRS

Pontifícia Universidade Católica do Rio Grande do Sul Porto Alegre, Brazil

Farina Simone Dr., Curator, Natural History Museum of Pisa University (Calci Pisa)

Feijò Anderson Dr., Universidade Federal de Paraiba – Fiel Museum of Natural History, Integrative Research Center, Chicago

Fiechter Matt Dr., Snow Leopard Trust www.snowleopard.org

Fondazione Archivio Ligabue–Parma - Italy

Gandois Hélène Ms., CITES Secretariat Programme and Documentation Officer

Gerard Patrice Mr., Amis- Assistant-Engineer Archaeological Field Techniques CNRS

Gerngross Peter Dr., Biogeomaps

Göhlich Ursula Prof., Curator of Vertebrate Paleontology of Natural History Museum, Wien

Guilaine Jean Prof., Musee National D’histoire Naturelle- Paris

Malcolm H. Hast Prof. Emeritus, The Feinberg School of Medicine Northwestern University

Hyward W. Mattew Prof. PhD, University of New South Wales, Australia

Von Hoenning-O’Carroll Betty Dr., President of Cheetah Conservation Foundation, Italiy

Holubiev Artem, Drawing of Homotherium letidens

Kitchener Andrew Dr., Principal Curator of Vertebrates in the Department of Natural Sciences of National Museum of Scotland, Edinburgh

Jacobson Andrew Prof., Nicholas School of the Environment, Duke University, Durham, United States

Mallon David Dr., Co-chair of the IUCN/SSC Antelope Specialist Group, Associate Lecturer in the Division of Biology and Conservation Ecology at Manchester Metropolitan University, United Kingdom

Marker Laurie DPhil, Founder and Executive Director of Cheetah Conservation Fund

Martelli Paolo Dr., Veterinarian at Ocean Park of Hong Kong

Mcknight Lidija Prof., Manchester University

Murphy J. William Prof., Veterinary Integrative Biosciences Faculty of Genetics, Faculty of Ecology and Evolutionary Biology, Texas A&M University

Naturaliter SNC (Catia Morucci, Alessio Salvadori, Gianluca Salvadori) Museo di Storia Naturale di Capannoli, Pisa, Italy

Oliveira do Nascimento Fabio Dr., Mastozoologia, Museu de Zoologia da Universidade de San Paulo, Brasil (MZUPS)

Palacio Rocio Dr., Coordinadora General A.G.A. Alianza Gato Andino

Pimm Stuart Prof., Nicholas School of the Environment, Duke University, Durham, United States

Price Campbell Dr., Curator Of Egypt and Sudan - Manchester Museum

Protopopov Albert Prof. Academy of Sciences of Republic of Sakha

Richard Thomas Dr., Global Communication Coordinator of Traffic: Wildlife Trade Specialists

Salesa J. Manuel Dr., Curator of Paleontology, Natural History Museum of Madrid

Sanderson Jim Prof., Global Wildlife Conservation, Austin, United States

Small Cats Conservation Foundation

Simeonowski Velizar, Wildlife artist and zoologist, Field Museum of Natural History, Chicago

Trompert ‘Beer’ Frans Mr., For the Works of Walty - Holland

Vigne Jean Denis Prof., Musee National D'histoire Naturelle- Paris

Von Hoenning O'carroll Betty Prof., Language Teacher and President Cheetah Conservation Fund Italia

Werdelin Lars Prof., Senior Curator, Fossil Vertebrate, Natural History Museum, Stockholm

Whitlatch Terryl, Illustrator, Creative Talent Network for Drawings

Wilting Andreas Dr., IZW Leibniz-Institut for Zoo and Wildlife Research/Sabag Forestry Departments

Thank you to all those who gave me permission to publish their photographic images, some very rare and obtained with photographic traps, and those who have gave them specifically to publish in my book.

Arias Madariaga Emiliano Andrés Mr., South American Photographer

Martegani Vincenzo Dr., Italian photographer

Nunez-Lemos Antonio, South American photographer

Sliwa Alexander Dr., Researcher and Curator of Cologne Zoo, Germany

Gil Paulo, Photographer, Sector de Cinematografia (DED)- Zoo de San Paulo do Brasil

Koeberl Christian Prof., Director of Natural History Museum, Wien

Wilting Andreas Dr., IZW Leibniz-Institut for Zoo and Wildlife Research/Sabag Forestry Departments

Rivas Antonio, photographer, Programa de Conservacion ex situ del Lince Iberico

Lynx ex situ, Programa de Conservacion ex situ del Lince Iberico

Monticelli Cauê, Chefe do Centro de Conservacao da Fauna, San Paulo Zoo

Kirilyuk Vadim Dr., Managing Director of Daursky State Biosphere Reserve, Russia

Korinek Milan, photographer

Klatil Lubomir, photographer

Rudloff Klaus, photographer

Madaravally Ashok, photographer

Magalaes Bressan Paulo Dr., President de San Paulo Zoo Foundation

Veronesi Federico Dr., Italian photographer

Pfleiderer Johannes Dr., Head of Zoology at Duisburg Zoo, Germany

Shanshui Conservation Center, Peking University

Schumacher Alice, photographer of Natural History Museum, Wien

Townshend Terry Dr., Birding Beijing Web, Director of Eco-Action

Finally, I would like (last but not least) to thank, for their magnificent work on the revised taxonomy of the Felidae family, all the components of the Cat Classification Task Force of the IUCN/SSC Cat Specialist Group. Their work led to the realization of the Scientific Journal, (Editors, Christine Breitenmoser-Würsten and Urs Breitenmoser):

Kitchener A. C., Breitenmoser-Würsten Ch., Eizirik E., Gentry A., Werdelin L., Wilting A., Yamaguchi N., Abramov A. V., Christiansen P., Driscoll C., Duckworth J. W., Johnson W., Luo S.-J., Meijaard E., O’Donoghue P., Sanderson J., Seymour K., Bruford M., Groves C., Hoffmann M., Nowell K., Timmons Z. & Tobe S. 2017.

A revised taxonomy of the Felidae. The final report of the Cat Classification Task Force of the IUCN/SSC Cat Specialist Group.

Cat News Special Issue 11, 80 pp.

This innovative text is the one I followed for the taxonomic classification of the felids in my book.

Thanks are therefore extended to the authors of this scientific publication:

Andrew Kitchener (Chair CCTF), National Museums Scotland, United Kingdom

Christine Breitenmoser-Würsten, Co-Chair IUCN/SSC Cat Specialist Group, Switzerland

Eduardo Eizirik, PUCRS, Porto Alegre, Brazil

Anthea Gentry, The Natural History Museum, London, United Kingdom

Lars Werdelin, Naturhistoriska riksmuseet, Stockholm, Sweden

Andreras Wilting, Leibniz Institute for Zoo and Wildlife Research, Berlin, Germany

Nobuyuki Yamaguchi, Qatar University, Doha, Qatar Expert Group

Alexei Abramov, Zoological Institute, Russian Academy of Sciences, Saint-Petersburg, Russia

Per Christiansen, Zoologisk Museum Statens Naturhistoriske Museum Kobenhavns Universitet

Carlos Driscoll, Wildlife Institute of India, Dehradun, India

Will Duckworth, IUCN/SSC Cat Specialist Group, United Kingdom

Warren Johnson, Smithsonian Institution, United States

Shu-Jin Luo, Peking University, China

Erik Meijaard, Borneo Futures, Jakarta, Indonesia

Paul O’Donoghue, University of Chester, United Kingdom

Jim Sanderson, Small Wild Cat Conservation Foundation, United States

Kevin Seymour, Royal Ontario Museum, Canada Review Group

Mike Bruford, University of Cardiff, United Kingdom

Colin Groves, Australian National University, Acton, Australia

Mike Hoffmann, Zoological Society London, United Kingdom

Kristin Nowell, IUCN/SSC Cat Specialist Group, United States

Zena Timmons, National Museums Scotland, United Kingdom

Shanan Tobe, Arcadia University, United States

As for the editorial preparation of this book, the publisher Charlotte Cockle; the acquisition editor Ana Valutkevich, who also believed in my project; the editorial project manager Devlin Person; the production project manager Sreejith Viswanathan; the senior copyright coordinators Regina Lavanya Remigius and Praveen Kumar; and the cover designer Mark Rogers gave indispensable help.

To everyone, I extend my heartfelt thanks.

Chapter 1

Order of carnivores (Carnivora)

Abstract

Many animals have a diet based on meat (animal protein): from sharks to crocodiles, and from snakes to birds of prey; the mammal order also includes bats (Chiroptera) and insectivores (Insectivora), monkeys (chimpanzees and baboons), and prosimians.

However, this chapter deals with the carnivores (order Carnivora), that from a taxonomic point of view belongs to a precise order of mammals and that share a common phylogenetic history together with particular anatomical features.

They belong to the order of carnivores, which is divided into 16 families, for example, dogs and wolves (family Canidae), cats and big cats (Felidae), bears (Ursidae), raccoons (Procyonidae), weasels (Mustelidae), genets (Viverridae), sea lions, walruses, seals, and sea elephants (superfamily Pinnipedia), etc.

All Carnivora have specialized teeth for their meat-based diet and tusk-shaped canines with which they kill their prey. The bones of their front legs have paw bones (carpal bones) called scaphoid and lunar (semilunar), which are fused together in a single bone called the scapholunar.

Keywords

Order Carnivora; meat diet; carnassial teeth; carpal bones; scaphoid; lunar; scapholunar; evolution; Miacidae

Chapter Outline

Introduction 1

The carnivorous animals 1

Man and carnivores 2

Chapter 1.1 Anatomical features common to order Carnivora 3

Chapter 1.2 Features of Caniformia and Feliformia 4

Carnivora denture or dentition 6

Chapter 1.3 Evolution, phylogeny, and classification 7

African lion (Panthera leo leo), order Carnivora, certainly the best known representative of this group of animals, as all members of the family Felidae, is a skilled predator and has a hypercarnivorous diet.

Introduction

The carnivorous animals

Many animals can be defined as carnivores from the point of view of their diet: for example, sharks, among the fish, crocodiles and snakes, of current reptiles as well as many dinosaurs from the past; and eagles, hawks, and owls among the birds.

Many mammals are carnivores, examples include the marsupial carnivores such as the Dasiuridae family or quoll (Dasyurus), the Tasmanian devil (Sarcophilus harrisii), the famous Tasmanian wolf or thylacine (Thylacinus cynocephalus) (relatively recently extinct), and the marsupial lion (Thylacoleo) of the Australian Pleistocene. Other orders of mammals present with totally or mostly carnivorous diets including bats, other insectivores, and some monkeys (chimpanzees, baboons) and prosimians. However, those we will deal with are the carnivores (order Carnivora) that, from a taxonomic point of view, belong to the precise order of mammals and share a common phylogenetic history together with particular anatomical features.

The order of Carnivora includes dogs and wolves, cats and big cats, bears, raccoons, weasels, genets, seals, etc.

(A) Eurasian lynx (Lynx lynx linx) order Carnivora, family Felidae. (B) Siberian weasel (Mustela sibirica) order Carnivora, family Mustelidae. (C) Maned wolf (Chrysocyon brachyurus) order Carnivora, family Canidae. This South American species has an omnivorous diet and is particularly greedy for a certain type of wild tomato (Solanum lycocarpum).

Man and carnivores

Within the order of Carnivora there are two species of mammals which are the most popular of human pets: the cat and the dog. These are bred everywhere that man has settled; with cats being the most common pet with about 500 million specimens around the world. Dogs, often called man’s best friend, is also used as a working animal (for hunting, towing sleds, guarding flocks, etc.) or as guide dogs for the blind, sniffer dogs (police dogs), and for carrying out rescues after avalanches and storms.

Their relatives and wild ancestors, namely wolves and the big wild cats, are admired as great hunters, but the relationship with the Carnivora also creates a series of conflicts and a complicated outlook, with opinions that can be very different or even conflicting, in different cultures and in various historical periods. This simple admiration for their strength and beauty has occasionally escalated to a veneration or viewing them as sacred as in many ancient cultural and religious traditions. However, carnivores have also always been considered as rivals and dangerous competitors, especially when man hunted prey for food; they were and can still be a serious danger to farmers and their livestock, and there is also a conflict over space due to natural and wild environments giving way to agriculture.

The invention of hunting as a sport as a test of courage or to obtain their precious furs, or for other multiple reasons whether selfish, utilitarian, or merely as a convenient petty economy, has reduced many species of carnivores to the point of extinction, especially the larger ones (tiger, lion, bears, wolves, etc.) which need large spaces in which to live together with their prey, with which they play a primary role as predators and/or superpredators, which is an indispensable role in the maintenance of the ecological balance of the natural environment. Many species have been eradicated from most of their natural habitats and relegated to live in restricted protected areas, which are not always sufficient to safeguard their future.

(A) Red tabby (long haired) and (B) Caucasian shepherd dog (a female with four puppies). These are breeds domesticated and bred by man, but which derive from wild carnivores.

Chapter 1.1 Anatomical features common to order Carnivora

The Carnivora are an order of mammals to which belong animals that are almost all adapted to a predominantly carnivorous diet (i.e., they need high percentages of animal proteins obtained from meat and fish meat), and for this reason, their anatomical structure is specialized for predation and in the digestion of this type of food; their small intestine is short (three to six times the length of their body) and their digestive enzymes include urate oxidase or uricase, which allows them to digest the uric acid contained in meat. Very few species have subsequently evolved by adapting to an omnivorous diet (composed of different percentages of plant substances and animal substances), such as badgers, bears, and the almost vegetarian, giant and lesser pandas (bamboo eaters), and the aardwolf (Proteles cristata), an Ienidae, and bat-eared fox (Otocyon megalotis), a Canidae, which have an entomophagous or insectivorous diet (termites). Some Canidae, such as the foxes (Vulpes sp.) and the maned wolf (Chrysocyon brachyurus), being opportunistic species, feed abundantly on fruit in the season that they ripen. Felids belong instead to a group described as hypercarnivorous as their diet is almost exclusively meat (more than 70%).

The red or lesser panda (Ailurus fulgens) is the only species of the Ailuridae family.

Like the giant panda (family Ursidae), it has an almost vegetarian diet based on bamboo. Both these Asian species, have a sort of sixth finger in the front legs, which is the sesamoid bone of the hand, and has the shape and function of an opposable thumb, it is used to grasp bamboo twigs and take them to the mouth.

To this order belong minute species weighing as little as 25 g like the least weasel (Mustela nivalis) up to the giant polar bear and Kodiak brown bear, which can exceed 800 kg, although the greatest of all is the enormous southern elephant seal (Mirounga leonina) with a weight of up to 3000 kg.

The gait can be plantigrade or digitigrade and the paws are always supplied with strong claw nails. The muscles of the hind legs are often massive, equipped with strong tendons that allow in many cases long leaps and rapid snappy movements (sprinters). In the Pinnipeds (Pinnipedia), earless seals, sea lions (eared seals or otary), and walruses, the limbs are transformed into fins.

They have very developed senses, especially sight and hearing, but in some an acute sense of smell prevails as revealed by the development of the nasal mucosa and the olfactory region of the brain, especially in the some Caniformia and Felidae; the latter also have very acute sight, with eyes suitable for night vision.

The following six anatomical features largely define carnivores.

1. All carnivores have elongated and pointed canines with a conical shape called tusks which are used to kill prey; the premolar and molar teeth have a pointed shape with one or more sharp cusps; the so-called carnassial or sectorial teeth are composed of the fourth upper premolar (P⁴) and the first lower molar (M1), they are larger and very sharp, essential for being able to cut (with a scissor-like action) the muscles and tendons of prey, and to obtain pieces of meat of a size suitable for being ingested.

2. The third upper molar (M³) is missing in adult Carnivora dentition.

3. Within the skull it is notable that the braincase (the cerebral portion of the skull) is well rounded and well developed, so that the frontoparietal suture of the cranial bones is placed anteriorly.

4. An important characteristic to determine the taxonomic rank in Carnivores is the particular conformation of the bones of the base of the skull; it is here that there is also the tympanic bulla (auditory bulla), an ovoid hollow bone formation which contains the three hearing bones (the hammer, anvil, and stirrup). Until a few years ago it was believed that the bulla was formed from the tympanic bone only, however, more recent studies have shown instead that its morphogenesis evolved in different ways in the various families of carnivores, and that it may also derive from the fusion of several bones (ectotympanic, rostral, and caudal entotympanics). These different anatomical shapes influence the path of the carotid artery and this set of the features are one of the most important diagnostic factors for the determination of some taxa, including fossil taxa.

5. In the skeleton, the clavicle is missing or very small (vestigial).

6. On the front legs there are always with five digits, often with the first short and vestigial (the dew-claw), which is not used in walking, however, especially in the Felids, it is provided with a hooked claw that is used for predation. There are three small carpal bones (scaphoid, semilunar, and central) that merge into the single bone (scapholunar) of the wrist. The hind legs can have four or five digits depending on the gait adopted by the species. Plantigrade species have five digits and digitigrade species four. Felids (cats) and Canidae (dogs and wolves), which include adept cursorial and sprinter species, are digitigrade; meanwhile Ursidi (bears) and Procyonidae (raccoons) which are not cursorials have legs suitable to their plantigrade gait.

(A) In the order of Carnivora the carpal bones (of the front legs), scaphoid and lunar (semilunar), are fused into a single bone, the scapholunar. (B) Within the Carnivora there are various types of gait, plantigrade, semiplantigrade (some Viverridae like Paradoxurus), semidigitigrade (some Mustelidae like the badger), and digitigrade. The digitigrades put only the end phalanges of their digits on the ground, a gait that allows greater speed.

Chapter 1.2 Features of Caniformia and Feliformia

The order of Carnivora is composed of 16 families divided into two suborders.

1. The suborder Caniformia, which means dog-like includes bears, wolves, raccoons, weasels, etc., and the Clade Pinnipedia. It has evolved with diversification centers that have been found in North America and northern Eurasia; the most ancient fossils found in these continents have anatomical structures adapted to an arboreal life. Today’s representatives of the suborder have largely plantigrade gait, except Canidae which are digitigrade; they tend to have an omnivorous diet and often have a large number of teeth—from 34 (only in the weasel) to a maximum of 48–50. A diagnostic characteristic is the tympanic bulla, containing a part of the organs of hearing, which is formed by ectotympanic bones and has no internal median septum which divides it into two chambers. In some families including the Canidae and Ursidae, the alisphenoid canal is present and the internal carotid artery is not obstructed by the temporal bone structure and so is not reduced.

2. The suborder Feliformia, which means cat-like, includes cats, hyenas, genet, mongooses, Malagasy mongooses, etc. It originated with diversification centers located in Africa and southern Asia, with species with an anatomical structure suitable for an arboreal life. The current species almost all have a digitigrade gait (at least in the front legs) and many have nails that are partially retractable or completely retractable, such as the claws of the Felidae. They have a much more carnivorous diet than caniforms and some groups are hypercarnivorous. To improve the possibility of predation the most advanced, like some Felidi, has gradually decreased the number of teeth to up to 28–30; in the other Feliformia there can be up to a maximum of 40 teeth. Their tympanic bulla is formed by entotympanic and ectotympanic bones and it is divided into two chambers by a median septum. With the alisphenoid canal absent, the internal carotid artery is reduced because it is obstructed by the bone structure.

Skull of a carnivore showing the basicranial region, where the two auditory or tympanic bullae are seen at the sides of the occipital hole (or foramen magnum).

Carnivora denture or dentition

Pink, canine teeth; blue, carnassial teeth (M1) (PM⁴)

Chapter 1.3 Evolution, phylogeny, and classification

The assumptions made in order to exactly define the period in which the first true Carnivora appeared are sometimes discordant. It had always been asserted that they first appeared in the late Paleocene, but according to the latest fossil discoveries and based on calculations on the evolution of their anatomical characteristics and molecular phylogeny, today it is generally thought that they probably appeared more than 66 million years ago (Mya), at the beginning of Paleocene. It has always been assumed that the fossil forms belonging to the superfamily of Miacoidea were the first true ancestors of modern carnivores. Miacoidea remains are found in Western Europe, Asia, and North America, with some forms that survived for almost the entire Eocene. These evolved into two families:

1. Miacidae with many genera, including Vulpavus, Paroodectes, Miacis, etc.

2. Viverravidae with fewer genera, including Viverravus, Mustelodon, etc.

Similar to a cross between small genets and martens, they all presented characteristics typical of arboreal animals, such as frontal eyes, an elongated skull and body, short legs, long tail, and pointed and retractable claws. Their dentition, typical of carnivores, with long zanniform canines (fang-shaped canines=tusk shaped canines) and molars with pointed cusps, was generally composed of 42/44 teeth.

However, according to important research in the field of comparative anatomy and paleontology, the true nature of Miacoidea as direct ancestors of the current Carnivora is questioned, with some considering them as only belonging to the Clade of the Carnivoramorpha, which includes the true carnivores, the Miacidae, the Viverravidae, and some forms not well defined from a systematic point of view, including some genera such as Tapocyon and Quercygale (which some authors continue to ascribe to the Miacoidea). Especially Tapocyon, for some of the characteristics of the base of the skull, can perhaps be considered very similar or closely related to the group that originated the true Carnivora. It is, however, quite certain that the first true members of the carnivorous order appeared in Central Asia (perhaps in China) at high latitudes, in regions with a cold temperate climate (Rolland et al., 2015) and there they diversified considerably and then spread to tropical regions later.

Shortly after their appearance (after the middle of the Paleocene) the order was immediately differentiated into the two great present suborders: Feliformia and Caniformia. The results of the latest research indicate that the division between Feliformia and Caniformia began between the beginning and the middle of the Paleocene (57 Mya), whereas before it was assumed to be a much later subdivision, in the middle Eocene about 43 Mya.

Later these Taxa gave rise, in addition to the current families of carnivores, also to many important fossil groups (Nimravidae, Amphicyonidae, Percrocutidae, etc.), which are well represented in many fossiliferous sites and were therefore widely diffused in the past, but which became extinct in various successive epochs but allowing some to survive until the beginning of the Pliocene.

Miacidae: Paroodectes feisti, which belongs to the Carnivoramorpha group. These are animals more or less the size of a genet and with arboreal habits. To this group, that dates back to the Pleistocene, there are many genera and species, some of which represent the true ancestors of the modern carnivores.

Blue branches (blue lines), Caniformia, Red branches (red lines), Feliformia Figure from Van Valkenburgh, B., Wayne, R.K., 2010. Carnivores. Curr. Biol. 20(23), 2157. Figure created by G. Slater and J. Pollinger. Van Valkenburgh, B. Pang, B., Bird, D., Curtis, A., Yee, K., Wysocki, C., Craven, B.A., 2014. Respiratory and olfactory turbinals in feliform and caniform carnivorans: the influence of snout length. Anat. Rec. 297, 2065–2079. https://www.researchgate.net/publication/266857277.

https://upload.wikimedia.org/wikipedia/commons/c/c3/Carnivora_phylogeny_%28eng%29.png

After a long evolutionary history, Caniformia and Feliformia have evolved into the current 16 families composed of about 286 species. These are widespread in almost all the natural and anthropized environments of the Earth, from the Arctic tundra to the tropical forests, in deserts, wetlands, and the sea. The carnivores have reached and colonized all the continents except Australia, but even here they are still present, with at least one species of feral dog, the dingo, imported in prehistoric times (3000–4000 years ago) from Indochina and the Sunda islands by man, but that has now assumed a well-defined ecological role in some Australian ecosystems.

Chapter 2

Family Felidae

Abstract

According to recent systematic reviews, the Felidae family is comprised four subfamilies:

• Two, the Proailurinae (origin of all the Felidae) and Machairodontinae, are both extinct, and known only through fossilized skeletal remains, these include the famous saber-toothed cats.

• The other two, Pantherinae and Felinae, include all known living species and the extinct fossil species related to them (cave lion, American lion, European jaguar, etc.).

The Proailurinae (genera Proailurus, Pseudaelurus, Styriofelis) are not considered by all scientists as the true ancestors of the Felidae family and as the components of a distinct subfamily they appeared at the end of the Oligocene.

According to some researchers the family Felidae (one of the five monophyletic groups of cat-like carnivores or Feliformia) instead has two subfamilies, Machairodontinae and Felinae, to which is added a clade (taxa not named) composed of the genera Hyperailurictis and Nimravides (this last genus is not to be confused with the family Nimravidae which is Carnivora but not part of the Felidae) and a sister group of the Felinae composed of the genus Styriofelis. According to this taxonomic proposal, Pseudaelurus quadridentatus would be part of the Machairodontinae and Proailurus lemanensis is considered the basal taxa of the whole family Felidae.

Keywords

Cats; big cats; evolution; saber-toothed cats; Panthera blytheae; retractable claws; carnassial teeth; hypercarnivorous diet

Chapter Outline

Chapter 2.1 Classification 13

The lineages 17

Classification of the Felidae family 20

Chapter 2.2 Evolution 20

Chapter 2.2.1 The origins: the subfamily Proailurinae 23

Evolution in teeth and development of the coronoid process 25

Chapter 2.2.2 Subfamily Machairodontinae: phylogeny and classification 26

Chapter 2.2.2.1 Tribe Metailurini 28

Chapter 2.2.2.2 Tribe Smilodontini 29

Chapter 2.2.2.3 Tribe Homotheriini 30

Chapter 2.2.2.4 Tribe Machairodontini 31

The Rancho La Brea fossil deposit 33

Biology of Machairodontinae 34

Chapter 2.2.3 The current Felides 40

Chapter 2.2.3.1 Subfamily Pantherinae and lineage Panthera 41

Chapter 2.2.3.2 The occurrences of Panthera blytheae and Its importance in chronological dating of Pantherinae 42

Chapter 2.2.4 Subfamily Felinae 46

Felinae Lineages 47

Chapter 2.3 Anatomical features of current Felids 49

Skeleton and muscles 50

The skeleton (skeletal system) 50

The legs and retractable claws 52

The skull 55

Dentition of current Felides 57

The function of the teeth 58

Musculature (muscular system) 61

Cats always land on their feet 62

The fur 64

Furs: patterns, colors, and habitat 68

Fur as a protection from cold 69

Chapter 2.1 Classification

Carl von Linné is known more commonly as Carl Linnaeus (or Linnaeus) from the Latinized form of his name, Carolus Linnaeus (Råshult, May 23, 1707–Uppsala, January 10, 1778). He was a Swedish physician, botanist, and naturalist, considered the inventor of the modern scientific classification of living organisms. In 1735 and then again in 1758 he published his famous Systema naturae, for Regna tria naturae secundum (systematice proposita per) for Classes, Ordines, Genera, Species (…), with which he founded the foundations of modern systematics, inventing the binomial nomenclature (with genus and species), assigning a universally valid scientific name to every living organism then known. This is why he is recognized as the father of modern taxonomy.

He also classified the representatives of the then known family Felidae, such as the lion, tiger, ocelot, leopard, jaguar, lynx, and the domestic cat and, wanting to give them a taxonomic arrangement based on common anatomical features, he gathered together all these species under one genus Felis (from the Latin felis=cat), taking into account some features of the teeth and the retractile nails, and considering these typical peculiarities to be common to the entire genus.

He then assigned some species to the genus Felis, describing for each of them some characteristic distinguishing features that enabled their recognition:

Felis leo: long floccosa tail, which means having a bow (tuft) of terminal hairs, body yellow-red (Luteo rufus), and the male with a shaggy thorax (with a mane);

Felis tigris: streaked body;

Felis pardus: spotted body;

Felis lynx: short tail with a black tip, ears with tufts of hair on the tip, spotted reddish color.

Two original pages, 41 and 42, of "Systema naturae" edited 1735–58 of Carolus Linnaeus, written in Latin, the official language of science until the end of the 18th century. The taxonomic classification of the family Felidae (Mammalia Ferae: Felis) and of some of the then known species are illustrated; on page 41 are the lion, tiger, leopard, on page 42, the jaguar (Onca), ocelot (Pardalis), domestic cat (Catus), and on page 43 the lynx, with the specific anatomical features. https://upload.wikimedia.org/wikipedia/commons/a/a2/Sistema_Naturae_%281758%29.pdf

Following the Systema Naturae, for a few centuries all felids were grouped under the same genus Felis, given the many anatomical similarities that, beyond their body size, make the family seemingly very homogeneous. Even in the 20th century, an important text on African mammals (Kingdon, 1997) still brought together all the small felids (including the serval, caracal, and African golden cat) in the genus Felis. The cheetah, due to its particular anatomical features, was assigned to a new genus, Acinonyx (Brookes, 1828) and separated into a subfamily of its own: Acinonychinae (Pocock, 1917). Only at the start of the 20th century were the larger Felid species (lion, tiger, jaguar, leopard) divided into their new genus: Panthera. Due to its large size, the puma was assigned to both the genera Felis and Panthera, by different authors; the caracal, because of its short tail and tufts on the ears, was also assigned to the same group as the lynx (lynx caracal) and was sometimes called the desert lynx.

The increase in anatomical knowledge resulted in the genus Panthera being merged into a subfamily, that of Panterinae, which gathered all the species that could roar, a possible feature because of the hyoid bone, which in this genus was not completely ossified but maintained a fibrous front and ligaments that allowed some mobility of the larynx. All the other cats (classified as the Felis genus), ascribed to the subfamily Felinae, instead had the ossified bone completely ossified so that the larynx was blocked, preventing the ability to roar but allowed them to purr, with this latter capacity not being present in the Panterinae. However, with the advancements in the anatomical knowledge of the vocal apparatus of the Felidae, even this definition was questioned, because it was seen that not all big cats could roar, resulting in snow leopards being ascribed for a long time to a different genus (Uncia). It was also discovered that the clouded leopards (Neofelis), although true panthers, had a rigid and ossified hyoid bone and did not roar (in Chapter 7: Ethology, we will see that the roar is also linked to the structure and the dimensions of the vocal folds and of the thorax that acts as a sounding board). Also in the subfamily of the Felinae (small cats), the problems of taxonomic classification were numerous, and before the advent of modern techniques of genetic and biomolecular analysis, studying the comparative anatomy, especially the skulls and colors of the fur of the various species, the Felis genus, was subdivided into many genera, which were not always accepted later, although some are still considered valid, including: Leptailurus for serval, Otocolobus for manul, Herpailurus (Puma) for jagouarundi. Especially in the new species that were being discovered and classified, and particularly within the group of small felids of South American and Southeast Asia, there was an explosion of new genera, with a new one for almost every species. Other new genera included the following:

• South America: Lynchailurus for the pampas cat or colocola, Oreailurus for the Andean cat, Oncifelis for the guigna (or kod-kod) and Geoffroy’s cat, Oncilla for the tiger cat, and Leopardus for the ocelot and margay.

• Asia: