Table Of ContentFELIDS AND HYENAS
OF THE WORLD
Wildcats, Panthers, Lynx, Pumas,
Ocelots, Caracals, and Relatives
José R. Castelló
Foreword by
Alexander Sliwa and Andrew Kitchener
PRINCETON UNIVERSITY PRESS
PRINCETON AND OXFORD
Copyright © 2020 by José R. Castelló
Copyright of the photographs remains with the individual photographers
Requests for permission to reproduce material from this work should be sent to
Permissions, Princeton University Press
Published by Princeton University Press
41 William Street, Princeton, New Jersey 08540
6 Oxford Street, Woodstock, Oxfordshire OX20 1TR
press.princeton.edu
All rights reserved.
ISBN 978-0-691-20845-9
ISBN (pbk.) 978-0-691-20597-7
ISBN (e-book) 978-0-691-21186-2
Version 1.0
British Library Cataloging-in-Publication Data is available
Production and design by José R. Castelló, Madrid, Spain
CONTENTS
FOREWORD 5
ACKNOWLEDGMENTS 7
INTRODUCTION 8
PANTHERA LINEAGE
Tigers, Lions, Jaguars, and Leopards 30
PUMA LINEAGE
Puma, Jaguarundi, and Cheetah 80
OCELOT LINEAGE
Ocelot, Margay, Guiña, Colocolo, and Related Species 96
LEOPARD CAT LINEAGE
Leopard Cat, Pallas’s Cat, and Related Species 134
CARACAL LINEAGE
Caracal, Serval, and African Golden Cat 156
WILDCAT LINEAGE
Domestic Cat, Wildcats, and Related Species 174
LYNX LINEAGE
Lynx and Bobcat 202
BAY CAT LINEAGE
Asiatic Golden Cat, Marbled Cat, and Borneo Bay Cat 226
HYENAS
Hyenas and Aardwolf 238
SKULLS 258
GLOSSARY 263
REFERENCES 267
INDEX 277
FOREWORD
It is with great pleasure and admiration that I write the foreword to Felids and
Hyenas of the World. José Castelló has set the gold standard with his two
major preceding books Bovids of the World (2016, Princeton University
Press) and Canids of the World (2018, Princeton University Press) for
practical, affordable, and visually stunning reference guide books on these
mammalian families. Not only is he a gifted photographer himself, he is
unrelenting in searching for the best available images on the World Wide
Web to show the characteristics of the depicted species and subspecies. There
are billions of images of one of man’s favorite companions and majestic
sources of fear and admiration, the small and large cats in their diverse family
that roam all continents, apart from Antarctica, and that prowl through all
terrestrial habitats. What book can satisfy our urge to revel in the beauty,
elegance, and diversity better than this current one, which systematically
strives to depict the full glory of the Felidae? This tome reflects painstaking
dedication to deliver a reference without comparison.
All wild living cat species and the domestic cat are depicted with a full page,
most often showing currently recognized subspecies on their own spread with
several pictures of adults and kittens. For each species information about its
morphology, distribution, habitat, ecology, biology, and diet, as well as its
main threats and current conservation efforts is provided, taken from the most
current literature available, and here again José Castelló has delved deeply
into both reasonably available and cryptic (gray) literature. This is no small
feat as there is substantial references on some members of the cat family,
particularly the European and North American species and the classic
Panthera cats, while information on particularly the smaller species is
comparatively scarce and hard to obtain. Synthesizing all of this into
digestible form requires great stamina and systematic rigor, which is
presented here.
I am certain that this book will find high acclaim and wide readership and
will foster even more admiration and a drive toward deeper research into the
lesser known felid species, leading to concerted efforts to protect individuals,
populations, and species by providing an affordable and portable current
reference.
Alexander Sliwa
PhD, Curator, Cologne Zoo, Germany
Felid Taxon Advisory Group (TAG) Chair of the European Association of
Zoos and Aquaria (EAZA)
Invited Member of IUCN SSC Cat Specialist Group
Black-Footed Cat Working Group Project Leader; Sand Cat Sahara Team
Co-Leader
I have been waiting a very long time for a book like this one. My interest in
wild cats dates back to when I was in my mid-teens when a new charity, the
Cat Survival Trust, was established locally where I lived. Fascinated by
opportunities to get close to many of the world’s smaller cat species, I
became ever eager to find out more, but good books about cats were lacking.
However, my first significant encounter with a wild cat was when I was
seven years old, with a young female Puma on Sandown Beach in the Isle of
Wight, where the famous lion tamer Nyoka was taking it for a walk on a lead.
When it chased after me, dragging Nyoka with it, I quickly became terrified
and was too upset to meet it once it was under its owner’s control. When I
started work at National Museums Scotland, I met Dr. Nigel Easterbee, who
was carrying out a status survey on European Wildcats in Scotland and
research on how to distinguish wildcats from domestic cats and their hybrids.
When Nigel died tragically in a car accident, I took up this important area of
research and added hundreds of specimens of road-killed wild-living cats to
the museum’s collection, which recently helped to show the imminent
extinction of the European Wildcat in Scotland today through hybridization.
At the same time that I met Nigel, I was finishing off my first book, Wild
Cats of the World (1991, Cristopher Helm), which was a synthesis of what
was known of the behavior and ecology of the world’s cats, but it wasn’t an
identification guide to all taxa, which seemed an ever-distant dream. Soon
after, I was invited to join the IUCN Cat Specialist Group by the late Peter
Jackson. My feline career continued with research on geographical variation
in Tigers in the 1990s and Clouded Leopards in the 2000s, resulting in the
recognition of the first new big cat species, the Sunda Clouded Leopard, in
180 years. In 2010 I was asked by Urs and Christine Breitenmoser to lead the
IUCN Cat Specialist Group’s Cat Classification Task Force, which published
its final report in 2017. This report reduced the number of cat subspecies
dramatically, with only a modest increase in the number of species. But it
was beyond the scope of this report to show what they looked like. So, I am
delighted that the first comprehensive book that illustrates all the world’s
species and subspecies of cat has at last been published. José Castelló’s
Felids and Hyenas of the World is a magnificent guide to these two important
families of carnivorous mammals.
Having been very impressed by José’s two earlier books, Bovids of the
World, published in 2016, and Canids of the World, published in 2018, I
wondered whether he had turned his attention to the felids. So, I contacted
him out of the blue, offering him any assistance that I could. By return email
José confirmed that he was indeed working on the felids, and possibly the
hyaenids, for his next volume. José’s books follow a well-established pattern
with an introductory chapter that provides a brief but interesting, summary of
different aspects of our knowledge of felid and hyaenid morphology, both
inside and out, classification and taxonomy, behavior, including social
organization, feeding and diet, reproduction and communication, with
subsequent sections looking at distribution and habitat, evolution and the
fossil record, domestication and conservation status, and ending with a guide
on how to use the book. What enriches this book and brings it to life is the
stunning photographs of a wide variety of cats and hyenas showing different
aspects of their appearance, anatomy, and behaviors.
The meat of this book is the series of chapters that correspond to the main
felid phylogenetic lineages identified by molecular studies led by Steve
O’Brien and his many colleagues over the last few decades, with the lesser
known hyenas and aardwolves in the final chapter. After an introductory
section to each chapter, each species and subspecies gets a two-page spread
with a description, other names in multiple languages, taxonomy, similar
species, reproduction, behavior, distribution, habitat, and conservation status.
What makes this book remarkable and very useful to a cat specialist like
myself is the array of photographs showing lateral, frontal, posterior, and
other views of each taxon, including differences between the sexes and also
juveniles and younger animals where their appearance differs significantly.
José has established an incredible network of people around the world who
have sent him photos of all species and subspecies from these two
mammalian families. Having worked for seven years in revising their
taxonomy, I find it fascinating to be able to look at the differences between
all the subspecies that are recognized in the IUCN’s A revised taxonomy of
the Felidae.
Cats, of course, have been animals of endless fascination for people over
many millennia. Recorded in cave paintings dating to tens of thousands of
years ago, to images on pottery, standing stones, mosaics, and sculptures
from across a wide array of ancient civilizations from the Incas to the Ancient
Egyptians, cats have continually captured our imaginations with their grace
and beauty as well as their power and strength. Sadly, our fascination with
cats has not prevented us from killing them as troublesome consumers of
livestock and people, or exploiting them for their fur or bones for our vanity
or medicine. Hyenas, in contrast, have been much neglected and vilified as
mere scavengers or cruel killers that are much persecuted wherever they live.
Only the small and dainty Aardwolf, which feeds almost exclusively on
termites, is excluded from these stereotypes, but as a consequence we know
little about it. However, it is important that we are aware that the few hyaenid
species that survive today are a vestige of a much more widespread and
diverse family that populated the Old World, and that despite their canid-like
appearance, they are closely related to the felids and are equally fascinating.
Felids and Hyenas of the World offers a new opportunity to get to know the
world’s hyaenids better. I hope you will take the opportunity to use this
stunning guide, whether in the wild or in captivity, to brush up your
knowledge of and ability to identify the diverse, beautiful, and captivating
felids and hyaenids of the world. I certainly will!
Andrew Kitchener
PhD, Principal Curator of Vertebrates, National Museums Scotland, UK
Member of the IUCN Cat Specialist Group
Chair of the IUCN Cat Specialist Group’s Cat Classification Task Force
Honorary Fellow of the Royal Zoological Society of Scotland
ACKNOWLEDGMENTS
This book is dedicated to my wife, Beatriz, and my children, Alejandro and
Beatriz, for their love and support during the long hours that I spent working
on this book. Also, I need to thank my parents, who taught me the value of
hard work and education. Compiling a book like this one would be
impossible without the help of dozens of experts and wildlife photographers.
I am deeply grateful to all the colleagues and friends who have been part of
this long endeavor.
A special thanks to Alexander Sliwa for his expert and gracious assistance in
preparing this book, for sharing his collection of photographs, and his vast
knowledge of felids and hyenas. Alexander is a conservation biologist,
Curator at the Cologne Zoo, Felid Taxon Advisory Group Chair of the
European Association of Zoos and Aquaria, member of the IUCN SSC Cat
Specialist Group, and a major expert in the behavioral ecology of many
species of wild cats.
I am also particularly grateful to Andrew Kitchener, Principal Curator of
Vertebrates in the Department of Natural Sciences at the National Museums
Scotland, Chair of the Cat Classification Task Force, member of the IUCN
SSC Cat Specialist Group, and the author of The Natural History of the Wild
Cats (1991, Cornell University Press).
A very special thanks goes also to my friend Sergey Chichagov, a Latvian
biologist and passionate photographer, who has shared his profound
knowledge of wild cats and hyenas and hundreds of photographs with me. I
wish also to acknowledge the help provided by Roland Wirth, founder of the
Zoological Society for the Conservation of Species and Populations; Klaus
Rudloff, Curator Emeritus at Tierpark Berlin; and Pierre de Chabannes, a
photographer specializing in animal conservation. My grateful thanks are also
extended to Alex Kantorovich, curator of Hai Park Kiryat Motzkin; Jonas
Livet, a French biologist and consultant in zoology; Juan S. Villalba-Macías,
an Uruguayan conservationist and Curator at Bioparque M’Bopicuá; Balázs
Buzás, a Hungarian conservationist and wildlife photographer; and Phil
Myers, Professor Emeritus and Curator Emeritus at the Museum of Zoology,
University of Michigan-Ann Arbor.
Finally, many thanks also to Jim Sanderson, Sebastian Kennerknecht, Laila
Bahaa-el-din, Gregory Breton, Guillaume Dupuy, Francisco Javier Erize,
Mike Gordon, Eyal Bartov, Jaime San Román, Daniel López Velasco, Jesús
Rodriguez-Osorio Martín, Igor Antuna, Javier Amores, Francisco Erize,
Valeriy Maleev, Vladimír Cech, Rodrigo Villalobos, Nayer Youakim, Ricky
Reino, Tomasz Doron, Johannes Pfleiderer, Paulo Barreiros, Michal Sloviak,
Alexander Meyer, Alexander Coke Smith, Otgonbayar Baatargal, Arno
Meintjes, Ulrike Joerres, Johan Bordonné, Milan Korínek, Antonio Núñez
Lemos, Kevin Schafer, Santiago M. Carrillo, Ricardo Fernández Chaves,
Arjan Haverkamp, Minor Torres Salazar, Agustín Emoris, Bruna Zafalon da
Silva, Puch Corinne, Cochahuasi Animal Sanctuary, Mendis
Wickramasinghe, Michael Lorentz, Jonas Van de Voorde, Jean-Francois
Potier, Florent Doko, Joachim S. Müller, Peggy and Marc Faucher, Verónica
Araya García, Nikhil B. Vatsal, Christopher Momberg, Nate Hart, Bharath-
Shreyas Photography, Jorge Serpa, Anda Ciurezu, David Howlett, Javier
Gómez Aoiz, Elena Mashkova, Andrea Riveros Díaz, La Senda Verde
Wildlife Sanctuary, Bruna Zafalon da Silva, Demis Bucci, Tim Sagorski,
Gustavo Muniz, Avijit Sarkhel, Angela Meyers, Huilo Huilo Biological
Reserve, Reserva Nacional Mocho Choshuenco, Parque Nacional Laguna
San Rafael, Eduardo Minte Hess, Enrique Couve, Alfredo Boettiger, Luiz
Mosca, Carmelo Pamies Boluda, David Piaggio De Casanova, SERFOR,
Markus Lilje, Lukas Blazek, McKenzie Greenly, Murthy Kantimahanti,
Vishwanath M. K. Mysore, Vickey Chauhan, Yoann Lombard, Ezra Hadad,
Eric François, Erwin & Annemieke, Greg Pierson, Esteban Argerich, Juan
Reppucci, Eliraz Dvir, Yossi Eshbol, Javier Muñoz Gutiérrez, Parinya
Padungtin, Vivek Sharma, Raymond De Smet, Willie van Schalkwyk, Juan
José Part Oliver, Vijaybabu Kaliappan, Shahab Mirzaean, Narong
Suwannarong, Tambako the Jaguar, Jeremy Holden, Kedar Nath Timsina,
Mike & Glen Heramb Wildlife Photography, Radhakrishnan Rajagopalan,
Tin Man Lee, Ron Gallagher, Zoo Praha, Iman Memarian, Debankur Biswas,
Bjorn Olesen Wildlife Photography, Young Photography, Sid Bramhankar,
Stephen Young, Kevin Hatfield, Cede Prudente, Santiago F. Burneo, Thomas
Retterath, Jasmine Curtis, Ronny Graf, Josh More, Jaime E. Jiménez, Aditya
Singh, Dominique Salé, Armand Meding, Bram Demeulemeester, Jaime
López, Marie Reed, Anuroop Krishnan, Warren Metcalf, Alcey Kangit,
