The Wild Cat Book: Everything You Ever Wanted to Know about Cats 9780226145761

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The Wild Cat Book

Fiona Sunquist and Mel Sunquist

The Wild Cat Book

With photos by Terry Whittaker and others t h e u ni ve r s i ty o f c h i c ag o p r ess • chicago and lond on

fio na su nq u ist is a science writer, photographer, and for fifteen years was a roving editor for International Wildlife Magazine. mel sunq u ist is professor emeritus in the Department of Wildlife Ecology and Conservation at the University of Florida, Gainesville. t erry wh it taker is a UK-based photographer specializing in wildlife conservation and the environment. The University of Chicago Press, Chicago 60637 The University of Chicago Press, Ltd., London © 2014 by The University of Chicago All rights reserved. Published 2014. Printed in China 23 22 21 20 19 18 17 16 15 14    1 2 3 4 5 ISBN-13: 978-0-226-78026-9 (cloth) ISBN-13: 978-0-226-14576-1 (e-book) DOI: 10.7208/chicago/9780226145761.001.0001 lib rary of con gr e s s catalogi n g -in -p u b li c at i o n data Sunquist, Fiona, author.   The wild cat book / Fiona Sunquist and Mel Sunquist;   with photos by Terry Whittaker and others.    pages cm   Includes bibliographical references and index.   ISBN 978-0-226-78026-9 (cloth) — ISBN 978-0-226-14576-1 (e-book) 1. Felidae. I. Sunquist, Melvin E., author.  II. Whittaker, Terry (Photographer), illustrator.  III. Title.   QL737.C23S862 2014  599.75—dc23 This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper).

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Contents

Introduction 1

Panthera Lineage Lion  5

Lions are the only social cat. They live in prides composed of related females. Pride members often join forces to hunt large prey.

Jaguar  17

South America’s largest cat, the jaguar’s canines deliver the most powerful bite of any big cat.

Tiger  29

Largest of the living felids and the only striped cat, tigers are built to grapple with prey several times their own weight.

Snow Leopard  41

The stocky, short-legged snow leopard hunts blue sheep and ibex in some of the most rugged terrain on earth.

Leopard  49

The leopard combines the power and strength of the big cats with the grace and versatility of the small cats.

Clouded Leopard  61

Named for the large cloud-shaped markings on their coat, clouded leopards are powerfully built cats with dagger-like teeth.

Bay Cat Lineage Bay Cat  69

The rare bay cat is found only on the island of Borneo. Thought to be extinct, it was rediscovered in 1992.

Asiatic Golden Cat  77

The Asiatic golden cat can be almost any color. Its fur varies from black to spotted, to bright red or gray.

Marbled Cat  73

With its long tail, large feet, and a blotched coat, the arboreal marbled cat resembles a miniature clouded leopard.

Caracal Lineage Serval  81

Specialized for sound-hunting, the tall serval has the longest feet of any cat. These remarkably successful hunters use their height and large ears to locate small rodents in the tall grass.

African Golden Cat  93

The powerful-looking African golden cat is built like a small leopard. Recent DNA studies reveal these cats are closely related to the serval and caracal.

Caracal  87

The caracal’s leaping abilities are legendary—these cats were once used for sport hunting by Indian nobility. Like the cheetah and lion, the caracal’s range covers part of India. v

Ocelot Lineage Ocelot  99

Ocelots are one of the most commonly seen cats in South America. Their short, sleek fur is marked with dark spots and rosettes, creating perfect camouflage.

Margay  103

Andean Cat  121

The Andean cat lives at high altitudes in the Andes, where it feeds almost exclusively on mountain viscachas.

Oncilla  125

Renowned for their climbing ability, margays are the most agile and acrobatic of the cats.

The tiny, delicate-looking oncilla could be mistaken for a smaller version of the margay. Like the ocelot and margay, the oncilla give birth to a single kitten.

Geoffroy’s Cat  109

Pampas Cat  129

About the size of a house cat, the Geoffroy’s cat is an adaptable, flexible predator. This cat has the unusual habit of leaving its feces high in the branches of trees.

Common in South America’s dry open grasslands and high deserts, the pampas cat resembles a burly, long-haired house cat.

Guiña  115

The guiña is the smallest cat in the Western Hemisphere, found only in a small part of coastal Chile and Argentina.

Lynx Lineage Eurasian Lynx  133

Twice the size of the other lynx, Eurasian lynx are forest cats. Deer are their main prey.

Iberian Lynx  139

The Iberian lynx is the world’s most endangered cat—only about 250 cats remain in Spain and Portugal.

Canada Lynx  143

Tall and leggy, the Canada lynx is a specialized hare hunter. Large feet with toes that can be spread to form a snowshoe help these cats travel over snow.

Bobcat  147

The short-tailed bobcat is the most abundant and widespread wild cat in North America.

Puma Lineage Cheetah  153

The fastest mammal on Earth, the cheetah can reach a speed of 70 miles per hour in a few seconds.

Puma  165

The adaptable puma is often thought off as a “big cat,” but it has recently been found to be more closely related to the cheetah and jaguarundi.

Jaguarundi  175

Related to the cheetah and puma, the odd-looking jaguarundi communicates with chirps and whistles. Unlike most cats, jaguarundi hunt mostly during the day.

Leopard Cat Lineage Pallas’s Cat  181

Found in Mongolia and China, the squat, short-legged Pallas’s cat has a face that resembles a Pekingese dog.

Fishing Cat  187

The only species named for its habitat of eating fish, the fishing cat is an excellent swimmer.

Leopard Cat  193

Flat-Headed Cat  197

Flat-headed cats resemble fishing cats; they are excellent swimmers and share the fishing cat’s preference for fish and frogs.

Rusty-Spotted Cat  201

The aptly named “hummingbird” of the cat family, the tiny, quick rusty-spotted cat is Asia’s smallest felid.

The small, lightly built, long-legged leopard cat is Asia’s most common small felid.

Domestic Cat Lineage Domestic Cat  207

Wildcats  225

Black-Footed Cat  219

Sand Cat  233

Domestic cats are believed to have moved in with humans some 10,000 years ago in Mesopotamia, being initially attracted to the rats and mice that fed on stored grain.

The tiny black-footed cat of southern Africa gets its name from the black hair on the soles of its feet. Males and females communicate via a very loud “meow,” which has been compared to the roar of the tiger.

The wildcats of Europe, Asia, and Africa are currently subdivided into five subspecies. The domestic cat is sometimes considered a sixth.

One of the few cats that digs burrows, sand cats are found among the rocky outcrops and sand dunes of the Sahara and the Middle East.

Jungle Cat  239

Adaptable enough to live near villages, the jungle cat is Asia’s equivalent of the jackal.

Acknowledgments 243 Bibliography 245 Additional Readings 259 Illustration Credits 261 Index 263

Pallas’s cat (Otocolobus manul).

Introduction God made the cat to give humankind the pleasure of caressing the tiger. —Anon.

When we returned from a long trip to India, our then-four-year-old daughter was greatly relieved to be back home. As we walked into the house, she flung herself on her favorite cat and exclaimed, “I need to touch a purr!” If you have ever lived with a cat, you know exactly what she meant. Soft and caressable, house cats are comforting companions in the purring body of a wild beast. Their lithe, light-footed movements remind us that they are just a whisker away from their wild relatives, and their do-as-they-like attitudes make us want to woo them. We feel honored when they respond, perhaps because we sense they are by nature solitary creatures that have become social with us. In wild cats we see the beauty of our house cats writ large. For thousands of years, our ancestors and wild cats lived in daily contact. Stealthy, powerful, and intelligent, cats shadowed early human hunters, competing with them for prey. In turn, we scavenged their kills but lived in fear of their covert stalk and deadly canine teeth. We made them our totems

and respected them as symbols of strength, courage, and royalty. Today the image of the crouched tiger or leopard is etched in our consciousness as a symbol of power and of wild places. Keen and intensely familiar, we know the voice, recognize the silhouette. In dark places it stirs a dimly remembered feeling and makes us catch our breath in awe. This admiration, respect, and fear are part of our collective ancestral memory. Our lives have become more distant from the wild, but we remain entranced by the perfect beauty of cats. Personally, as we have learned more about these alluring creatures, our interest has intensified and expanded to include their welfare and survival. We hope this book stirs interest in both and reminds people of the pure pleasure of “touching a purr”! While wild cats vary in size from the tiny 4-pound (2 kg) black-footed cat to the 550pound (250 kg) Bengal tiger, they are nevertheless instantly recognizable as cats. All cat species share many physical similarities, 1

2  i n t r o d u c t i o n C AT FA M I LY T R E E

37 SPECIES Lion Leopard Jaguar Tiger Snow leopard Clouded leopard Sunda clouded leopard

Panthera ancestor 6.4 MYA

Bay cat ancestor 5.41 MYA

Felid ancestor 10.8 MYA

Asiatic golden cat Bornean bay cat Marbled cat Caracal African golden cat Serval

Caracal ancestor 5.6 MYA

9.4 MYA

Geoffroy’s cat Guiña Oncilla Andean cat Pampas cat Margay Ocelot

Ocelot ancestor 2.9 MYA 8.5 MYA

Lynx ancestor 3.2 MYA

8.0 MYA

Iberian lynx Eurasian lynx Canada lynx Bobcat Puma Jaguarundi Cheetah

Puma ancestor 4.9 MYA

7.2 MYA

6.7 MYA

Leopard cat Fishing cat Flat-headed cat Rusty-spotted cat Pallas’s cat

Leopard cat ancestor 5.9 MYA

6.2 MYA

10

Domestic cat Wildcat Sand cat Black-footed cat Jungle cat

Domestic ancestor 3.4 MYA

5

4

3

2

1

Today

MILLIONS OF YEARS AGO (MYA)

A recent revolution in the study of DNA has thrown up several surprises in terms of relationships among the cats, and it is now possible to construct a cat family tree. Geneticists have found that the thirty-seven species of cats fall into eight distinct lineages or groups of related species, which evolved during the past 10 million years.

and they all make their living in the same way. But we now know that there are some unexpected differences among the species, particularly in their assemblages. Early eff­ orts to arrange or classify the cats into related groups were based on similarities in appearance, dental characteristics, the shapes of the bones in their ears, and shared geography. However, with the advancement of technologies to study the DNA of species, over the past two decades our knowledge of cat evolution has been revolutionized. We now have the first cat family tree.

Geneticists have found that the thirty-seven species of cats fall into eight distinct lineages or groups of related species, which have evolved during the past 10 million years. The big surprise was the cheetah, which was removed from the list of the big cats and is now firmly established as a close relative of the puma. We begin with the appearance of the oldest lineage, the Panthera or “big cats,” and make our way through time to the emergence of the most recent lineage, the domestic cat group.

Panthera Lineage

Lion

PAN TH E RA L EO

After watching countless TV specials, most of us associate lions with the prey-rich, rolling short-grass plains of the Serengeti, but these conditions exist in only a small part of the lion’s geographic range; most lions live in woodlands, dry forests, scrublands, and even deserts. Lions are unique among cats in that the core of their social system is the pride. Prides are composed of related females—mothers, sisters, daughters, and aunts and all of their cubs—which live together in groups to defend the best territories. The well-known lion biologist Craig Packer has likened lion prides to “street gangs competing for turf.” Successful prides of lionesses and their offspring often occupy the same area for generations. In the Serengeti, a pride of lions usually consists of two to eighteen adult females, their cubs, and one to seven males. Breeding coalitions

King of the beasts and symbol of power, the lion has been immortalized in history and legend. Their images appear on buildings, flags, in art, and as team mascots.

of males compete with other coalitions for females, and their tenure with a pride can be as short as a few months or last for several years. While the pride is the central unit of the lion’s social system, this system is not inflexible. During a prolonged drought in the central Kalahari, prey numbers declined to such a low level that the normal social system collapsed. Pride males and females abandoned their territories and became nomadic, traveling well outside their normal haunts. Females cooperated with unrelated females in hunts, and associations changed frequently. But lions also inhabit forested areas in both Africa and Asia, where they lead much more solitary lives. In these forested habitats, male and female lions rarely associate with one another except for mating. Studies of lions in India show that males tend to travel alone, make their own kills, and maintain their territories by scent marking and roaring. Females live alone or in pairs, with their young. Physically, lions and tigers are built along very similar lines. Well-muscled forequar5

6 p a n t h e r a l i n e ag e

The core of a lion pride consists of related lionesses and their cubs.

lion  7

European cave lions: Large, maneless lion disappeared about 14,000 years ago Panthera leo spelaea Until about 14,000 years ago, cave lions roamed the forests and grasslands of Britain and Europe, all the way north to Siberia and east to the Bering Strait. These lions were clearly very important totems for early European people, as they were often depicted in painstakingly accurate detail in cave paintings. Fossils and cave paintings show that cave lions were about 25 percent larger than modern lions and lived in prides like today’s African lions. However, the males did not have manes. Cave paintings show groups of cave lions hunting together like today’s Serengeti lions, and studies of the bones from cave lion kills showed that these lions preyed on horses, reindeer, deer, bison, and even cave bears. No one has yet confirmed what caused the cave lion to go extinct about 14,000 years ago. Most likely it was a combination of climate change and vegetation change. What is surprising is that about 6,000 years after the cave lion went extinct, a new, genetically distinct lion species, with a mane, invaded southern Europe. These modern maned lions survived in the Mediterranean region until about 2,000 years ago and were described by Aristotle and other classical Greek authors.

Females give birth to their cubs away from the pride but rejoin the pride when young are four to eight weeks old.

ters, designed to grapple with large prey, taper to slender hind legs. Lions rarely do any sustained chasing. When they do run, lions can reach top speeds of 30 to 37 miles per hour (48 to 60 km/hr). Lions have been recorded as feeding on almost every imaginable land mammal and even a few aquatic species; however, their principal prey are wildebeests, Thomson’s gazelles, warthogs, zebras, and buffalo. There are occasional records of lions killing extremely large animals, including giraffes, hippos, and young elephants. In Chobe National Park in Botswana’s Okavango Delta, some lion prides have learned to hunt elephants when other food

is scarce; at times as much as 20 percent of a Chobe pride’s total food intake consists of elephant meat. Most kills are of young elephants and take place at night. Groups of thirty or more lions try to spook and confuse the herd in the darkness by roaring and charging at juveniles and adolescents. When an individual becomes separated and appears vulnerable, it is swarmed by several lions, which bite the hind legs and cling to its haunches. In scenes that must resemble cave lion attacks on mammoths thousands of years ago, lion prides in Chobe also cooperate to bring down fullgrown elephants. Learning to become a competent hunter is a

lion  9

The mane story: Why do male lions have manes?

Why do male lions, alone among all the cats, have manes? Until recently, there were several different theories, but no completely satisfactory answers. It was only in 2002 that Peyton West, a graduate student at the University of Minnesota working in the Serengeti, resolved the mane question once and for all with a set of elegant experiments. West showed convincingly that the lion’s mane is a signal that advertises male condition. She ran a series of experiments in which she used life-size dummy lions with manes of different color and length, and presented them to wild, free-ranging lions. It turns out that lionesses use manes to choose the best mates, and male lions used manes to assess the condition of rival males and to avoid tangling with stronger males. Male lions begin to develop manes as they become sexually mature, and the mane continues to grow until they are about four or five years old. A lion’s mane reflects its physical condition—injured males have shorter, patchy manes and sometimes lose their mane altogether. West combined the results of her experiments with thirty years of data collected on a population of Serengeti lions. She found that males with darker manes generally had higher levels of testosterone, which meant they were more aggressive fighters. Aggressive males are better able to chase off wandering groups of bachelor males who would kill cubs in a pride takeover. Males with dark manes also have a longer life expectancy and are more likely to survive when wounded, and their cubs have higher survival rates. So, it transpires that the crowning glory of the king of beasts is a vital symbol and a signal to all around him. Lions in good condition have a full dark mane, and lionesses prefer males in good condition because a strong male increases the chances their cubs will survive. Males, on the other hand, want to avoid fighting with other males who are stronger than them, so they use the mane to calculate the strength of a rival.

1 0  p a n t h e r a l i n e a g e

While lionesses do 90 percent of the pride’s hunting, male lions usually tackle larger prey, like this 1,000-pound (455 kg) African buffalo.

lengthy process, and by the time they are two years old, young lions are capable of helping other hunting lionesses make a kill. Young females often stay in their birth pride for their entire life, while young males usually leave. Lions once roamed over most of Africa, Europe, the Middle East, and parts of India. They are thought to have reached Sri Lanka, where

they are commonly depicted in the country’s folklore and art, and likely survived until historic times. Today only about 350 free-ranging Asiatic lions survive in the Gir Forest region of western India. As human populations have expanded throughout Africa, lion populations have shrunk, becoming fragmented and isolated,

l i o n   11

Desert lions: A very different life than savannah lions

Several hundred lean, long-legged lions roam the scorching sand dunes and blisteringly hot boulder fields of Africa’s Kalahari and Namib deserts. These desert lions have learned to go without water for weeks at a time and survive in an environment where the heat is intense—in summer during the middle of the day, the surface temperature of the sand can be 158°F (70°C)—easily hot enough to cook an egg. Life for desert lions is very different from life as a lion in the Serengeti. Family ties are strong among desert lions, but relationships are long distance. Unlike the big groups of the Serengeti, members of the desert prides are not always together; they regularly break off into small groups of two or three. During droughts when conditions are especially bleak, individuals roam alone. Desert lions face a constant battle against thirst and high temperatures. They spend the day in whatever shade they can find and only hunt after dark. In the Serengeti, food is plentiful and lions rarely have to walk more than a few miles before they find a meal, but desert lions may walk 30 to 40 miles (48 to 64 km) in a single night, and pickings are slim. They range over areas 100 times larger than their Serengeti counterparts. Fritz Eloff, who studied Kalahari lions with the help of bushmen trackers, once followed a lioness that walked 25 miles (40 km) and tried eight times without success to catch a meal for herself and her cub. On another occasion, he tracked three lionesses as they walked for seven nights and caught nothing. All they found to eat that week was an ostrich egg. The desert lion’s main prey is gemsbok, a horse-size antelope that is a formidable adversary. Gemsbok have been known to kill lions and leopards by skewering them on their three-foot-long saber-like horns—one gemsbok was found walking around with the dead body of a leopard impaled on its horns. Desert lions have learned to survive on prey that other lions don’t eat; in addition to gemsbok, small prey like porcupines, bat-eared foxes, and aardvarks are a regular part of their diet. Serengeti lions hardly ever kill porcupines, but Kalahari lions depend on this prickly 40-pound (18 kg) food item—a third of their kills are porcupines. Ever resourceful, desert lions can go without water for weeks at a time, surviving on the body fluids of their prey. They lick dew from the grass, and lionesses have even been seen licking raindrops off each other’s fur.

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White lions: Separate species?

With their pale, haunting eyes and cream-colored fur, white lions have become the new favorite of the animal world. Some people view them as sacred spiritual symbols, while others see them as valuable commodities to attract visitors to zoos or to shoot as a rare trophy. In the natural world, these beautiful and popular white cats turn up occasionally in South Africa among the wild tawny lions; but they are rare—only a handful have been seen in the last 100 years. In 1975 two white lion cubs were seen in the Timbavati Nature Reserve near Kruger National Park. They quickly became celebrities, and soon, when another white cub was born and disappeared, people began to worry about their survival. The white cubs were captured and taken to a zoo in Pretoria, South Africa. Today about 500 offspring of these and a couple of other white lions are scattered across the world in zoos, circuses, and breeding farms. White lions are not a separate species from tawny-colored lions; they are a color variation of regular lions. Their white fur is the result of a mutation in the fur color gene, and, like white tigers, they are not an endangered species. Like other white animals, white lions are rare in the wild—and this makes them valuable because people like to see them, so zoos, circuses, and breeding farms create more white lions by inbreeding close relatives. The way to breed white lions is to pair two lions that have the same mutation in the fur color gene, because when both the male and the female carry the same white gene, there is a good chance that one of their cubs will be white. But as most of the lions that carry the white mutation are related, breeding white lions means continually inbreeding close relatives, mating fathers with daughters, and brothers with sisters. As with white tigers, this produces lion cubs with all kinds of physical and mental problems. A few well-intentioned people want to put some of the captive-bred white lions back into the wild in South Africa. The Global White Lion Protection Trust has several white lions living in a large enclosure and would like to eventually release them so they can mingle with the local wild tawny lions. Another group of white lions has been released into an enclosure in the Sanbona Wildlife Reserve in South Africa’s Western Cape Province. Because many of the lions used to create these white lions come from a mixed background of zoos and circuses, biologists believe we would be opening a genetic Pandora’s box into the wild population, and, once done, there would be no turning back. Other scientists suggest that a better option would be to focus on protecting all Kruger lions, because that would be protecting the whole wild population that naturally carries the white fur color gene.

l i o n   13

Roaring lions: A loud, long-range call

The roaring call of a male lion is extremely loud and has been measured at 114 decibels—the intensity of a noisy rock concert. Loud roaring is used for long-distance communication, to promote contact or spacing among lions, and the sound carries for miles. A lion’s roar has been described as “one or two moans, then full-throated thundering, until it dies slowly in a series of hoarse grunts.” The roars of males and females sound similar except that the roars of males are somewhat deeper in tone. Roaring by one lion in a group sometimes stimulates other group members to join in a resounding chorus.

1 4  p a n t h e r a l i n e a g e

Distribution of the lion.

and have disappeared from many areas. In a little more than two decades, lion numbers across Africa have declined to about 23,000, down from more than 200,000. This massive decline is attributed largely to direct conflict with humans, who retaliate for livestock losses with rifles, poison, and snares. The majority of remaining wild lions live in East Africa and southern Africa, where numbers have remained fairly stable for the last thirty years. The largest populations in East Africa are in Tanzania, princi-

pally in the Serengeti ecosystem and the Selous Game Reserve. Ecotourism in Africa has major conser­ vation implications for lions because these big cats are a “must-see” for tourists. While ecotourism is often touted as the way to compensate people for their losses, the market is limited and rarely lucrative enough to encourage ranchers to tolerate lions on their lands. The problem is that lions do not spend their time solely within national parks and game reserves, but often live and hunt out-

l i o n   15

side protected areas, where they frequently kill livestock. Lions that habitually kill livestock are not tolerated, even on commercial ranches that benefit economically from ecotourism. On communal lands, where thousands of people and their livestock share the area, lions have been wiped out. As several lion researchers have pointed out, the best way to conserve lions is to keep them away from cows and goats. Most lion attacks occur at night, when livestock are corralled in an enclosure called a boma. The traditional boma is made from densely interwoven thornbushes, with an opening that can be closed once the animals are inside. The opening is, however, the weak spot

in the enclosure; the approach of a lion often causes the livestock to panic, and in the ensuing stampede, they can push through the gap and are easily killed. Surrounding the boma with chain-link fencing can apparently solve this problem; in the few trials where fencing has been added, it provided an effective defense. Livestock losses can also be reduced by guards, using either people or dogs. status: IUCN Red List—Vulnerable weight: 200–600 pounds (90–272 kg) head-body length: 54–98 inches (137–250 cm) tail length: 24–39 inches (60–100 cm) litter size: 1–7 cubs, usually 3

Jaguar PAN TH E RA ON CA

Jaguars and leopards are closely related and quite difficult to tell apart. However, the jaguar is heavier than the leopard and looks like a much more powerful animal. It lacks the leopard’s lithe grace; instead, it is a strong, deep-chested, stocky-looking cat, with an unusually large head and short, sturdy limbs. Large or small, the jaguar gives the impression of unassailable power. Even a 70-pound (31 kg) jaguar looks as if it could overpower an ox. Its killing technique reflects the extraordinary strength of its muscles and teeth—the jaguar’s canine teeth are more robust and deliver a more powerful bite than those of the other big cats. A million years ago, huge European jaguars were common in the river forests of central Germany and France. Almost twice the size of today’s jaguar, these early jaguars hunted With a broad, powerful-looking head and massive canine teeth, jaguars are well equipped to tackle very large prey. Their bite has more force than that of other big cats, and they can bring down a full-grown Brahman bull by biting through the animal’s skull between the ears or horns.

deer and pigs in the warm floodplain forests of Europe, living alongside ancestral versions of the lynx and puma. Jaguars arrived in the Americas comparatively recently. Both fossil and molecular studies suggest that these cats entered North America via the Bering Strait roughly 800,000 years ago. Then, even more recently—some 280,000 to 510,000 years ago—jaguars moved south and colonized their current home in Central and South America. At that time South America was home to about twenty-five species of large herbivores. However, some 12,000 years ago, during the Pleistocene extinctions, all native South American mammals larger than about 145 pounds (65 kg) disappeared. The only relatively large prey species that avoided extinction were tapirs, capybaras, large armadillos, peccaries, and a couple of deer species. Jaguars are clearly built to grapple with the ghosts of the Pleistocene fauna. Their broad head, massive canine teeth, and short, stocky limbs equip them to tackle prey two to three times larger than their own body 17

18 p a n t h e r a l i n e ag e

Since the early 1970s, the jaguar has been on the list of totally protected species in most South American countries. Though the threat from the commercial skin trade has abated, the jaguar is still losing ground to habitat destruction, and many are shot as cattle killers every year.

weight. However, today these powerful cats live in a landscape that lacks the large animals they are equipped to kill. Despite their robust physique and heavy-duty weaponry, modern jaguars manage to survive on a diet of surprisingly small animals such as armadillos, rodents, birds, and marsupials. Natural prey for today’s jaguar is limited to peccaries, brocket deer, pampas deer, caimans, and capybaras, all of which generally weigh less than 110 pounds (50 kg). Really large natural prey like the occasional tapir and marsh deer are few and far between, so small prey is often the only thing on the menu.

In Peru, Belize, and Costa Rica—where jaguars survive on armadillos, agoutis, iguanas, and sloths—they have become much smaller in body size and weigh scarcely more than a large Labrador retriever. But despite the reduction in body size, these mini jaguars have retained the massive head, powerful teeth, and robust forelimbs of a large-prey specialist. About 500 years ago, early colonists brought cattle and horses to the New World and inadvertently reintroduced Pleistocene-size prey to the South American landscape. The cattle multiplied and roamed semi-wild in large herds, and the jaguar took to this new

j a g u a r   19

Jaguars in North America?: A small number of jaguars appear to be residing in Arizona Jaguars once roamed North America from Oregon to Pennsylvania, and though these cats disappeared from most of the United States before Columbus’s time, the species continued to hang on in Arizona, Texas, and New Mexico until recently. Sixty-two jaguars have been killed or captured in the United States since 1900, most of them within 62 miles (100 km) of the border between the United States and Mexico. There were so few sightings or records of jaguars in the United States over the following decades that the cat was believed to have disappeared from the United States. Then, in March 1996, in southeastern Arizona, a mountain lion hunter and his dogs were hot on the trail of what they thought was a puma. When the dogs treed the cat, the hunter was very surprised to discover the cat was a male jaguar. Six months later another male jaguar was photographed in the mountains of south-central Arizona. These observations prompted Arizona biologists to initiate a series of trail-camera surveys, and between 2001 and 2007, they accumulated more than seventy photos of at least four different male jaguars. Though there has yet to be any evidence of females, two of the males have been repeatedly photographed in the same area for several years, indicating they are residents. These borderland jaguars are thought to be a small segment of a larger jaguar population in Sonora, Mexico. In 1997 the U.S. Fish and Wildlife Service formally listed the jaguar as an endangered species in the United States.

food source with alacrity. Already physically equipped to kill large prey, jaguars grew to rec­ ord size on the millions of feral cattle that resembled its ancestral prey. Today Brazil—with more than 2 million head of cattle—is home to the world’s largest commercial cattle herd. There is no such thing as an average-size jaguar; the weight of this cat can vary by more than 100 percent, depending on where it lives. The smallest jaguars are found in Peru, where males weigh about 80 pounds (37 kg) and females, 70 pounds (31 kg). The largest jaguars are found in the floodplains of the Pantanal region of Brazil and the Venezuelan llanos, where males weigh about 225 pounds (102 kg) and females, 160 pounds (72 kg). Jaguars have an unusual method of killing large prey. Lions, tigers, and leopards usually kill large animals by biting them on the throat or the neck. The jaguar also employs a third killing technique that the other

big cats don’t use: biting through an animal’s skull. A cow killed by a jaguar often has two holes punched through the thick bones on the top of the head behind the ears. Jaguars kill capybaras with a bite to the back of the head, piercing the brain case with their canines: skulls have been found showing that the jaguar had inserted a canine neatly into each ear of the capybara. Although turtles may seem like a rather unlikely prey for a big cat, these reptiles form an important part of the jaguar’s diet in many parts of South America. River turtles can weigh as much as 70 pounds (31 kg), and female turtles are quite vulnerable when they come out of the water to lay eggs. Jaguars use their powerful bite to break open the lower edge of the turtle’s shell to get to the body cavity. On exceptionally large turtles, the space between the top and bottom shells is large enough for the jaguar to insert its paw and scoop out the

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Jaguars are often found near streams and watercourses, where they hunt basking crocodilians or wait for turtles to come on land to lay their eggs. The jaguar’s bite is strong enough to penetrate the tough hides of crocodilians or punch through the thick shells of large river turtles.

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Why are people not on the menu for jaguars?

One of the great puzzles about man-eating is why some big cats don’t or only rarely kill people. Though there are frequent records of man-eating tigers, lions, leopards, and occasionally pumas, jaguars almost never attack people. Many people have addressed the question of what makes a big cat kill humans, but the real question seems to be why do jaguars refrain from attacking humans? It is certainly not their congenial personality that stops jaguars from attacking people. Jaguars are considered to be the most difficult of the big cats to tame, and even when hand-raised, they are moody and temperamental. This is why you never see jaguars jumping through hoops or performing in stage shows. So if their nature is not the reason why jaguars rarely kill humans, what could it be? Prey selection in cats seems to be learned and culturally transmitted, and there are many real examples of tigers and lions that have “taught” their cubs the behavior of man-eating, elephant killing, or how to kill difficult prey such as buffalo. In terms of man-eating by big cats, this is underlined by the fact that the problem usually stops when the offending animal or animals are shot. Throughout Africa and Asia, big cats co-evolved with early humans and primates, and have been killing and eating early hominids and apes for hundreds of thousands of years. In South Africa, a 1.5-million-year-old skull fragment of a young hominid shows the characteristic twin punctures of leopard teeth. However, humans and jaguars have only lived in North and South America for a few thousand years. South American has no large ground-dwelling apes like Africa’s gorillas and chimpanzees—most South American primates are small and arboreal. Perhaps the answer is as simple as the fact that the jaguar has just not had the time or opportunity to develop a taste for humans as cat food.

meat. Empty overturned turtle shells with traces of jaguar hair on them are regularly found on turtle nesting beaches. Unlike the social organization of lions, tigers, and leopards, jaguar society remains a bit of a mystery. Even recent detailed studies using Global Positioning System (GPS) radio collars have not cleared up the question of how jaguar society functions. Despite their apparently confusing social system, jaguars communicate with one another like other big cats, using scent marks and visual marks. Their best-known form of communication is roaring. Early naturalists and explorers likened the call to thunder or to the roar of the caiman. The jaguar’s roar has best been described as five or six repetitions of a short, sharp guttural uh-uh-uh sound. Hunters imitate the jaguar’s roar by grunting into gourds or shells.

Both males and females roar, and there have been instances of two individuals engaged in intense bouts of counter-calling that have lasted as long as two hours. Jaguars live in dense tropical forests, arid scrublands, swampy grasslands, and even mangrove swamps. Throughout most of its range, the jaguar is strongly associated with streams and watercourses. These cats are excellent swimmers that have been seen crossing large rivers, and, like tigers, they sometimes escape the day’s heat half submerged in a stream. The number of cattle killed by jaguars varies from place to place and with the individual cat—cattle make up more than half the diet of some jaguars, while others rarely kill livestock. But because cattle ranchers kill jaguars in retaliation for cattle predation,

The sheep-size capybara is the world’s largest rodent. Semi-aquatic and highly social, they live in savannas and forests, in groups of ten to twenty individuals. The capybara is one of the jaguar’s favorite prey.

Jaguars seem to have a particular fondness for peccaries, and in some parts of South America, collared and white-lipped peccaries make up a quarter or more of the jaguar’s diet.

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Black cats: Melanism in wild felids

Black individuals show up among a quarter of the species in the cat family, including leopards, jaguars, bobcats, servals, golden cats, jungle cats, oncillas, Geoffroy’s cats, and guiñas. Just as albino individuals—all white with unpigmented red eyes—crop up occasionally in many species, so, too, do melanistic, or all-black, individuals. The color comes from melanin, the same pigment responsible for suntans. In domestic cats and leopards, a single recessive gene controls dark coat color, so both black and spotted leopard cubs can occur in the same litter, and black cubs can be born to spotted parents. Among jaguars, melanism is controlled by a dominant allele. Black cats are black because they have a mutation in the gene responsible for coat color. But the genes that control coat color, or pigmentation, often play a wider role in the functioning of the body, and the coat-color gene can influence two or three seemingly unrelated characteristics. As far back as 1828, in his famous book On the Origin of the Species, Charles Darwin commented on the association between blue-eyed cats and deafness. Most people know that white cats are white because they lack pigment cells. But surprisingly, the lack of pigment cells is also implicated in the white cat’s deafness. A clue comes from the fact that white cats with one blue eye and one yellow eye are often deaf on the side with the blue eye. Studies of lab mice suggest that pigment cells are involved in maintaining fluid in ear canals. Without the fluid, the canals collapse, causing deafness as the auditory nerves degenerate. Researchers Eduardo Eizirik, Stephen O’Brien, and their colleagues at the National Cancer Institute’s Laboratory of Genomic Diversity in Maryland have mapped, cloned, and sequenced the genes responsible for black coats in cats. One theory is that animals living in dark, humid forests generally have darker fur for camouflage. African leopards spend most of their lives in open habitats with dappled sunlight, where spots are the best disguise. In dark tropical forests, black coats may be better camouflage than spots. “The most likely benefit of melanism is camouflage for hunting,” Eizirik says. However, O’Brien, chief of the Laboratory of Genomic Diversity, suggests that there are many other selective pressures besides being able to hide. “Another explanation is that about 70 percent of selective pressures associated with the biological environment involve microbes and diseases,” O’Brien explains. Recent studies have shown that coat-color genes also affect the immune system. “The types of receptors used for coat colors are also used by viruses to enter cells,” O’Brien says. “It is plausible that some of these color mutations are adaptive relics of historic epidemics.”

Often the only sign of a jaguar’s presence in an area is its tracks. Male tracks are larger and more rounded than the elongated tracks of females.

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Distribution of the jaguar.

the jaguar’s future is inextricably linked to cattle ranching. Direct persecution by people is considered to be the most significant threat to the long-term survival of jaguars throughout their range. Because of this, bi-

ologists and conservation organizations have invested a great deal of time and effort trying to identify ways to reduce the number of cattle killed by jaguars. Some ranches have experimented with cre-

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ating special “maternity pastures,” in which cows and their newborn calves are kept for three months or longer. These pastures are fenced and are most effective when located well away from forested areas. Shortening the breeding season from year-round to a few months makes it logistically easier and less expensive to protect cows and calves. Some ranches have also tried using different breeds of cattle, especially those known to be more aggressive toward predators. Electric fencing around holding pens works well to deter predators, but maintenance is a problem in

tropical climates. In the end, partial solutions will probably be found in a combination of improved herd management, compensation, and tax incentives; but, ultimately, because cattle are large, available, and easy to kill, jaguars will always prey upon them. status: IUCN Red List—Near Threatened weight: 68–266 pounds (31–121 kg) head-body length: 43–67 inches (110–70 cm) tail length: 17–31 inches (44–80 cm) litter size: 1–4 cubs

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Tiger

PAN TH E RA TI G RI S

In a cage a tiger’s conspicuous dark stripes and brilliant gold coat stand out like a beacon; but in the patchy light of the forest, this bold pattern merges with the shadows, and the huge cat simply disappears. Tigers are powerful, burly animals. Their thick neck, broad shoulders, and massive front legs are designed for grappling with large animals, while their broad forepaws and long claws wrestle the prey into position and hold it for the killing bite. Tigers can single-handedly kill animals four or five times their own weight—a 500-pound (225 kg) male tiger can kill a 2,000-pound (910 kg) bull gaur. Tigers are not runners—they rarely chase their prey. Like other cats, they walk on their toes; soft toe pads distribute their weight over the balls of their feet, giving them not only a fluid walking motion, but also a silent one. They rely on stalking to get close, and Largest of the living cats, tigers are built to grapple with prey several times their own weight.

then overtake their target in an explosive final attack. Tigers are mostly nocturnal; they hunt when their prey is active and travel anywhere from 1 to 10 miles (3 to 30 km) per night. Tigers know their ranges well and rarely wander around while hunting; instead, movements are highly directional. They often move from one hunting area to the next as if they have a mental map of the good hunting sites within their territory, as well as the best routes between them. They use roads, trails, and gullies to travel between hunting areas, so they can move quietly and easily, and will lie in wait in dense vegetation near waterholes or close to places where deer and pigs are feeding. In the Russian Far East, tigers avoid hunting in deep snow cover because the unstable snow crust makes walking difficult and noisy. When moving through areas with deep snow, tigers often take advantage of frozen riverbeds and deer trails, or anywhere the snow depth is less and travel is easier. 29

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Tigers are the only striped cat. Whether it is facial markings or lines on the shoulders and flanks, no two tigers have the same pattern of stripes.

An adult male gaur is a formidable prey, and tigers have been killed during attempts to kill these 2,000-pound (910 kg) wild cattle.

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The “other” tigers: There are as many tigers in captivity as in the wild, but they have little conservation value Today there are more tigers living in chain-link cages in the backyards of America than there are roaming wild in the forests of Asia. An estimated 3,000 to 4,000 tigers live in captivity in the United States, while there are only about 3,500 tigers left in the wild. Astoundingly, in America tigers are almost as easy to buy as cattle—and just about as inexpensive. In Texas, for example, anyone can buy a tiger cub for as little as $500, and it is legal to keep a tiger in your backyard. Once the adorable playful cub becomes an adult, formerly enthusiastic owners are so desperate to find homes for them that they are advertised as “free to good home.” It is difficult to give away an adult pet tiger. People buy tiger cubs with the best of intentions; they are often animal lovers who cannot resist the magical lure of owning a tiger. Tiger cubs are heart-meltingly cute and mostly manageable for the first six months, but as they reach adulthood, they start to become dangerous. The cub that used to play happily with the golden retriever begins to bite, swipe, and urine spray, and owners start to rethink the wisdom of having a big cat around the house. There are literally thousands of unwanted adult tigers in North America. Reputable sanctuaries and rescue facilities are overflowing with them and cannot keep up with the demand for placement. But more homes for unwanted tigers are not the solution. The solution is education—so people understand that it is a horrible idea to try to keep a tiger as a pet—backed by well-enforced laws that ban the private breeding of exotic animals. Another serious problem with so many tigers in private ownership is what Philip Nyhus and his coauthors have termed “warped priorities and inaccurate messages.” Nyhus and his colleagues use Siegfried and Roy as an example, but they might just as well have mentioned any of the tiger breeders who tout their exhibits of “Golden Tigers,” “Snow Tigers,” “Royal White Tigers,” and “Magical White Lions.” Siegfried and Roy and the other breeders emotionally describe how they are conserving the last of these rare and special cats. The message resonates with the public, and people want to help, but the problem is that it is a totally misleading statement. These color morphs are not rare creatures in need of saving; they are genetic anomalies created and perpetuated by humans. Their habitat is an enclosure, and they can never be returned to the wild. Sad to say, they are of little conservation value and devalue the real conservation message. This is not helpful to wild tigers.

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Some authorities believe Siberian tigers were larger in the past, but today tigers in the Russian Far East average about 500 pounds (225 kg).

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Tigers are built to kill prey three to four times their own weight. Broad shoulders and massive forearms are ideal for grappling with gaurs, wild boars, and large deer.

A stalking tiger is the epitome of concentration and focus, continually assessing, making slight adjustments in its position before the final attack. To be successful, a tiger cannot be halfhearted—it must commit totally to the final smashing, explosive charge. Heedless of slashing hooves and antlers, it must go for the killing bite, ignoring the danger. In-

jury can be the ultimate penalty. A wounded tiger will starve if it cannot hunt. Tigers rely on two basic techniques to kill prey. They kill small animals with a bite to the back of the neck. With large, dangerous prey, tigers usually go for a throat bite, crushing the windpipe. Tigers usually make a large kill about once a week, and this will feed them for

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Tiger conservation: Conserving tigers is a “wicked” problem as the cat is worth more dead than alive

The tiger is the focus of hundreds of fund-raising campaigns and countless conservation programs. But despite the fact that governments and conservation organizations spend more than $47 million each year protecting the last tigers and their habitat, the tiger is barely holding on. Today there are only about 3,500 tigers left in the wild. You would think saving tigers would be comparatively easy. Tigers are symbols of nature’s power, viewed with awe by people everywhere. In polls for the world’s favorite animal, the tiger always tops the list. In the wild, tigers don’t cause substantial damage to property or livestock, and even when they do kill cattle, it is not difficult to set up compensation schemes. They do require large areas of connected habitat and plentiful prey, but this has been achieved for other species. The main problem with tigers is how they are valued. Protecting the tiger is more complicated than protecting jaguars, pandas, or most other animals simply because a dead tiger is much more valuable than a live tiger. Conserving the tiger is what scientists call “a wicked problem.” Stakeholders in the battle for the future of tigers have radically different views of the cat—many people, especially in the West, want to see them continue to survive and thrive in the wild, while poor farmers who live alongside them are ambivalent. For others, a tiger’s value lies in its body parts. Today the carcass of a wild tiger is worth more than $50,000. Skin, bones, whiskers, teeth, and claws—every piece of a tiger’s body is a commodity, and there is a bottomless pit of demand for pieces of this magnificent animal. To save the tiger as a species, we have to find a way to make a live tiger worth much more than a dead tiger—that’s the “wicked” problem.

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Tigers often hunt around the edges of clearings, taking advantage of every scrap of cover to get close enough to prey for the final rush.

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Black tigers: No record of a black tiger has ever been documented

There are many historical reports of black tigers, several of which have turned out to be bears or black leopards, but no photos or skins exist. Partially black tigers—in which the black stripes have merged on parts of the body, are more likely, and a skin exists with a black head and back. Poachers in the state of Orissa in India recently killed another partially black tiger.

several days. Once the kill is made, the tiger will lie nearby, feeding intermittently, often eating 30 to 60 pounds (14 to 27 kg) of meat in twenty-four hours. One exceptionally large male in Nepal ate 75 pounds (34 kg) of meat in one night. If the kill is shared among several tigers, it does not last long; a tigress and her two large cubs once ate 225 pounds (102 kg) of meat in two days, until there was nothing left but bones. Young male tigers become proficient killers by about 18 months of age; females seem to develop more slowly and stay with their mother for longer. Daughters also show a pattern of settling next to their mothers when they reach maturity. This results in a neighborhood consisting of clusters of closely related females, like a dispersed lion pride. Unlike their sisters, young male tigers leave their mother’s range at 18 to 24 months of age and travel widely in search of a territory of their own. The distances covered by these males can be extensive; one male in southwestern India was known to have traveled at least 125 miles (200 km), while another covered 175 miles (280 km). As these are straightline distances between beginning and ending points, the actual routes taken would likely be more circuitous. The largest of all the cats, an average tiger measures about 10 feet (3 m) from the tip

of the nose to the end of the tail. Males typically weigh 400 to 500 pounds (180 to 225 kg), while females are slightly smaller and lighter, weighing 220 to 350 pounds (100 to 160 kg). Tigers from Sumatra and other Indonesian islands are smaller and darker than tigers from more northern areas. Interestingly, while historical records of Siberian tigers list some huge cats weighing 700 to 850 pounds (315 to 385 kg), wild Amur tigers today are similar in size to Indian or Bengal tigers. We know this because the Siberian tiger project has been capturing, radio-collaring, and tracking tigers in the Amur region of the Russian Far East for fifteen years and has yet to document any wild tigers weighing more than the largest Bengal tiger. It is likely that the historical heavyweights were either exaggerated field estimates or that super-large tigers were extremely rare and no longer occur in Russia. Part of the tiger’s ability to persist in current and changing landscapes is its flexibility. Tigers are tolerant of an unbelievably wide range of habitat conditions. They are at home in the hot, dry thorn forests of Rajasthan and the steamy tidal mangrove swamps of the Sundarbans. They can live in tropical forest or the pine, oak, and birch woodlands of the Russian Far East, surviving at temperatures of -29°F (-34°C). Tiger populations can

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White tigers: Endangered species or . . . ?

White tigers fascinate people; they attract enormous crowds and bring in huge profits for zoos, circuses, and magic shows—a perfect white tiger may sell for $50,000. But are they really endangered species in need of saving, as some claim? White tigers are a color morph of the regular tiger—a variation that comes from a genetic mutation. In the wild, the genetic mutation that produces the white tiger color morph hardly ever happens. “White tigers are not an endangered species or even a separate species; they are mostly about money,” says Dr. Ron Tilson, former Director of Conservation for the Minnesota Zoo. “They are bred by people to make money.” In 1951 a male white tiger named Mohan was captured in central India. People wanted Mohan to father as many white cubs as possible but that presented a problem. White tiger cubs are born only when the mother and father carry the rare white gene. The odds of finding a female white tiger for Mohan to mate with were very slim. So Mohan was bred with his daughter Radha, who was assumed to have the white gene. Mohan and Radha produced four white cubs. When they grew up, those cubs were mated with

their father, Mohan. This father-daughter mating is called inbreeding. Inbreeding is the reason that the existence of white tigers is controversial. To get more white tigers, you have to breed close relatives, like Mohan and his daughter Radha. This sort of inbreeding causes a lot of physical problems. White tigers are often cross-eyed and have cataracts, cleft palates, hip dysplasia, and crooked spines. “People don’t talk about it much, but to produce white tigers, or white lions, you have to continuously breed father to daughter, father to granddaughter, and so on,” explains Tilson, adding, “The part that never surfaces is the number of cubs that die because of the effects of this inbreeding. The ugly side of the story is the number of cubs that are born horribly deformed. They may be pretty to look at, but they are a mess. More than half the cubs die.” There is also the problem of cubs that are not perfectly white. To get one ideal white tiger—with pure white fur, black stripes, and ice-blue eyes—you end up with hundreds of “throwaway cubs,” cubs that don’t look quite right. These are put to sleep or find their way to pet dealers and roadside zoos. Because of these problems, many reputable zoos now refuse to exhibit white tigers, despite their power to attract crowds. The National Zoo in Washington, D.C., the Minnesota Zoo, and many others no longer breed or keep white tigers. These zoos feel that knowing the truth about where white tigers come from may help people decide for themselves whether they want to support breeding more of them.

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Distribution of the tiger.

also grow rapidly. Tigresses often give birth to three to four cubs and may have another litter within two years. This means tiger populations can recover quickly as long as prey and habitat remain. All across Asia, where forest and village meet, tigers spend the day within hearing distance of the sound of human voices. As a general rule, as long as there is plenty of pro-

tective vegetation and wild prey, and tigers are not disturbed or persecuted, they can live alongside people, invisible and unnoticed. status: IUCN Red List—Endangered weight: 165–574 pounds (75–261 kg) head-body length: 57–91 inches (146–230 cm) tail length: 34–43 inches (87–109 cm) litter size: 1–5 cubs, 2–3 common

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Snow Leopard PAN TH E RA UN C I A

Found only in the mountains of Central Asia, the rare snow leopard makes its living in some of the steepest and most rugged terrain on earth. Prowling a moonscape of jagged rocks and scree slopes, snow leopards spend the winter chasing blue sheep across rocky cliffs, and in summer they search out fat marmots in high mountain meadows. These stocky, short-legged cats are the feline equivalent of high-wire artists, skilled at rushing prey in a landscape where misguided leaps can end in a 300-foot (90 m) drop. Oversize paws grip the rocks, and a tail as thick as a man’s arm acts as a balancing rod. An adult snow leopard is roughly the size of a golden retriever, but it can take down a mountain goat three times its own weight. The snow leopard’s most striking feature is its hooded, hauntingly pale eyes, which always seem to be gazing into the distance. The muscular, stocky snow leopard uses its massive front paws for climbing rocky inclines and for “snowshoeing” through deep, soft snow.

Unlike other cats, their iris is light green or gray. These pale-eyed cats have finger-deep, smoke-colored fur, stenciled with blurred charcoal spots that render them almost invisible among the rocks and boulders. They survive the piercing wind and cold by curling up with their long, thick tail wrapped like a scarf around their body. In profile a snow leopard skull has a marked step in front of the eyes, a short muzzle, and a high-domed forehead created by the enlargement of the nasal cavities. The large nasal cavities are thought to allow the animal to breathe more easily at high altitudes, where temperatures are low and oxygen is in short supply. The snow leopard’s massive paws are well suited both for gripping rocky inclines and for “snowshoeing” through soft snow. They use trails along river terraces, ridgelines, and the beds of deep gorges. Unlike many other cats, they do not seem to mind traveling long distances through deep snow. In western China, scientists followed the tracks of one snow leopard that had walked through 3-foot-deep 41

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The snow leopard’s short muzzle and high-domed forehead provide room for its enlarged nasal passages. This allows the snow leopard to breathe more easily at high altitudes, where temperatures are low and oxygen is in short supply.

(1 m) snow for 6 miles (10 km) without stopping to rest. In places the cat’s belly had created a furrow in the snow. Blue sheep and ibex are the snow leopard’s major prey. In areas where large prey is scarce, they survive by killing marmots and other small animals. In some places nearly half of their summer diet is made up of marmots.

Oddly, snow leopards also eat a great deal of plant material, and several authorities mention finding snow leopard droppings composed entirely of willow twigs and other vegetation. Though all cats eat a small amount of vegetation, there are no reports of other cats consuming such large quantities of greenery. Snow leopards do not roar like tigers, li-

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Counting cats with cameras: A non-invasive technique to estimate the number of cats in an area

In 1991 Indian research biologist K. Ullas Karanth had an idea that would revolutionize one of field biology’s most basic challenges. Struggling with the problem of estimating population densities of tigers in the wild, he wondered if he could set out cameras along game trails to “capture” the cats as photographs. Decades later “camera trapping,” as the animal census technique is now called, is one of the most common methods of estimating the abundance of secretive nocturnal animals like tigers, leopards, jaguars, and snow leopards. Biologists set cameras out in the forest on trails and along other natural routes. Any animal walking past the camera triggers the camera to automatically take the animal’s picture. Biologists now deploy dozens of these remotely triggered cameras at predetermined places in the forest for a set period of time; the resulting photos can be analyzed with complex computer models to produce a close estimate of the number of individuals living in the area. The technique works best with animals that have unique coat patterns. Camera trapping can also be narrowly targeted to answer more precise and practical conservation questions. Scientists have used remote cameras to count insects brought to feed nestling birds, monitor water holes, and indicate where highway crossings should be built. Nearly half of the deaths of the endangered Florida panther come from collisions with vehicles, so when a series of underpasses were planned to allow the cats to cross a major freeway in south Florida, biologists needed to know if the cats would actually use tunnels beneath the road. Camera traps revealed that panthers and all sorts of other animals, including alligators, did indeed use the underpasses, and the go-ahead was given to construct more of the million-dollar units. No panthers have died on the stretch of road with the underpasses.

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Snow leopards become exceptionally tame and gentle in captivity, and in the wild they are known to be the least aggressive of the big cats.

ons, jaguars, and leopards, but they do spit, hiss, growl, and cough-roar. Both sexes have a piercing yowl, which is most often used by females in heat and helps the two sexes find one another at mating time. Unique among the big cats, snow leopards have a well-defined breeding season and birth peak; most young are born in May and June, in a cave or rock crevice. Cubs must grow fast, and this is a time of enormous pressure on mothers to find enough food for their rapidly growing young before the weather window closes. Winter is only a few months away, and cubs must be large

enough and capable of following their mother when the cold and snow arrives. Not surprisingly, snow leopards are sometimes seen traveling in twos and threes during the winter, and occasionally five have been seen together; these groups are almost always mothers with large cubs. In captivity, snow leopards become exceptionally tame and gentle, and in the wild they seem to be one of the least aggressive big cats. There are very few reports of snow leopards attacking people, and they readily relinquish kills of domestic livestock, even

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Innovative conservation programs for snow leopards: Involving locals in conservation program pays dividends for snow leopards Across central Asia, people with different religions, languages, and customs compete with the snow leopard for its prey. In inhospitable environments where their own lives are on the edge, humans kill ibex, blue sheep, and almost anything that moves. Even marmots, a staple summer food for snow leopards, are shot for their meat and skins. In places where wild prey has been depleted by hunting and overgrazing, snow leopards subsist partially on domestic stock, and villagers retaliate by snaring and trapping the cats. In effect, impoverished herders and farmers across Asia are subsidizing snow leopard conservation by providing the cats with a ready source of food—their own animals. Rodney Jackson, founder of the Snow Leopard Conservancy, sums up the situation succinctly: “The snow leopards’ survival hinges upon an uneasy coexistence with subsistence farmers eking out their living from the same environment.” Conservation organizations have realized that for the snow leopard to survive, biologists must involve local people in the development of conservation plans that will suit not just the snow leopards, but also the people of the region. In Mongolia the Snow Leopard Conservancy has developed an unusual program to gain community support for protecting snow leopards. In exchange for signing a contract that says they will not kill snow leopards or their prey, local herdsmen and their families receive training and the tools to create woolen goods and handicrafts that are sold through the Snow Leopard Trust website. About $90,000 worth of Mongolian handicrafts were sold in 2007, and 400 herder families increased their household income by nearly 40 percent. Because the additional income is so significant, peer pressure results in communities policing themselves and outsiders to stop poaching. The scheme has been so successful that similar programs are now being attempted in five Kyrgyzstan villages and three Pakistani villages. In yet another approach, conservation organizations provided the local community near Hemis National Park in India with predator-proofing material such as wooden doors and chain-link fences to help improve their livestock corrals. This has virtually stopped snow leopards from killing sheep and goats. Stakeholders in the conservation of snow leopards and other wild cats include local communities, international conservation organizations, and the general public in developed nations. If you live in London or New York, and you want to keep snow leopards on the planet, it may no longer be enough just to say you love snow leopards. You may have to go to a website and buy something made by a Mongolian herding family. Or contribute to an organization that buys chain-link fences for poor farmers in India.

to a child brandishing a stick. They rarely defend themselves and are sometimes stoned or beaten to death by unarmed villagers. Until recent decades, snow leopards were protected by the sheer inaccessibility of their haunts. Today this huge empty land, known by early explorers as “the dead heart of Asia,” is also home to dozens of different ethnic groups that make up an expanding population of people trying to eke out a meager living raising goats, yaks, camels, and other live-

stock. In this harsh environment, where their own lives are on the edge, mountain residents kill ibex, blue sheep—and almost anything that moves. Even the dachshund-size marmot, a staple summer food for snow leopards, is shot for its meat and skin. In places where wild prey has been depleted by hunting and overgrazing, the cats subsist partially on domestic stock, and villagers retaliate by snaring and trapping the snow leopards. Poachers also search out the cats for their

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Snow leopards are found only in the high mountain regions of central Asia. These cats are found in rugged areas where the terrain is broken by cliffs, ridges, and deep ravines.

pelts and bones. Eugene Koshkarev, a former researcher at the Tien Shan High Mountain Physical Geography Station in Kyrgyzstan, says, “In the winter of 1993–94, the value of a snow leopard’s skin was more than sixty times higher than the minimum yearly wage in Kyr-

gyzstan.” In addition to the pelts, the bones are also sold as a substitute for tiger bones in the traditional Chinese medicine trade. Experts estimate as few as 4,500 or as many as 7,000 snow leopards may remain in the wild. Since 1973 the species has been officially

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Distribution of the snow leopard.

designated as Endangered. The Snow Leopard Conservancy uses innovative grassroots measures to lead local people toward effective stewardship of the endangered snow leopard and its habitat. For more information on the project, field updates, or to make a donation, go to http://wildnet.org/support/.

status: IUCN Red List—Endangered weight: 48–121 pounds (22–55 kg) head-body length: 34–49 inches (86–125 cm) tail length: 33–41 inches (85–105 cm) litter size: 1–5 cubs

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Leopard PAN TH E RA PARD US

Leopards are the very essence of cat. At close range, their strikingly beautiful spots are all you can see; but at a hundred paces, the dazzling markings become an invisibility cloak, and the big cat vanishes into the background like a dream. This quintessential cat combines the power and strength of the big cats with the grace and versatility of the smaller cats. A long, muscled body, thick, powerful limbs, and broad paws give the leopard an aura of enormous physical strength, and it is difficult to imagine that a cat this size can, if need be, survive on a diet of dogs and hares. Leopards are astonishingly versatile hunters. True generalists, they will eat whatever they can catch—and for the most part their diet reflects a list of what is available. They prefer duikers, gazelles, deer, and pigs in the 22- to 88-pound (10 to 40 kg) weight range, but will also eat jackals, bat-eared foxes, porcupines, aardvarks, pangolins, hares, bamboo The leopard combines the power and strength of the big cats with the grace and versatility of the small cats.

rats, and occasionally even ostriches. Leopards also manage to catch and kill a surprisingly large number of primates. They eat baboons, langurs, leaf monkeys, and macaques, and they also hunt and kill adult chimps and even gorillas. A remarkable study in South Africa’s Kalahari Desert was able to follow the lives of desert leopards in fine detail. Biologists working with bushmen trackers followed the footprints of individual leopards for hundreds of miles through the sand. Stepby-step, the bushmen were able to reconstruct the leopard’s daily movements; they could recognize each place where a leopard had crouched to watch a springhare or a bat-eared fox, discern where the cat had rested, and tell how often they had encountered other leopards. With a remarkable degree of accuracy, almost as if the events had been recorded on film, the bushmen were able to re-create hunting attempts; they knew the prey from its tracks, they could tell how long the chase was, and they knew 49

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Leopards are adaptable hunters and can survive on an amazingly skimpy rations, but they prefer gazelle, deer, and pigs in the 22- to 88-pound (10 to 40 kg) weight range.

if the hunt was successful. On one occasion they tracked a male leopard as it stalked a gemsbok cow and calf for a mile (2.5 km) before making the kill. The sand-tracking study showed that Kalahari leopards work incredibly hard and

survive on very meager rations. Both males and females regularly hunted for days without making a kill and went for weeks without drinking. Mothers with cubs often had to walk more than 12 miles (20 km) between each kill; and when they did find something to

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Amur leopards: A rare subspecies in the Russian Far East

In the Russian Far East, a rare leopard subspecies known as the Amur leopard is hanging on by a thread. These beautiful big cats are known for their long, luxuriant coat, an adaptation to surviving the region’s bitterly cold snowy winters. Protected in part by inaccessible terrain and the actions of hardworking conservation groups, about 30 to 35 Amur leopards currently roam the forested mountains of Primorsky Krai. Poaching, habitat loss, and now low genetic diversity are the major threats to their survival. Some 300 Amur leopards live in captivity, mainly in northern European and North American zoos.

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Leopards are solitary cats. Other than females with her young or a mating pair, they travel and hunt alone. Young leopards are dependent on their mother for a year or longer.

eat, it was usually small like a bat-eared fox, jackal, or springhare. Considering the fact that females need to find twice as much food if they are to raise a cub to independence, it is remarkable that the leopard population can survive at all under these harsh conditions. The leopard’s flexible behavior and diet also

allow them to live alongside larger predators like lions and tigers, and they are adaptable enough to survive where these larger cats cannot. Leopards living in Sanjay Gandhi National Park near Mumbai, one of India’s most populous cities, survive primarily on a diet of domestic dogs and rats.

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Perhaps just as importantly, leopards everywhere are better than any other big cat at living silently and unobtrusively alongside humans. When game scouts from Nairobi National Park were brought in to search for a leopard that had escaped from a Nairobi fair, they discovered the tracks of not one but

several leopards in the parks and gardens of the city. In Nepal scientists radio-tracked leopards that spent the day sitting quietly in hedgerows, within a few feet of villagers going about their daily lives, and at night cameras and tracks recorded these same leopards

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Leopards can climb extremely well when they want to. In areas where they have to contend with larger predators, they will often retreat to the safe haven of trees.

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Daughters prefer to live near mother: Results in clusters of related females

When the time comes for young pumas to leave home in search of their own space, daughters behave very differently from sons. Daughters try to stay close to their mother’s range and the place where they were raised; sons disperse to distant areas. This results in clusters of related females living close to one another. The home ranges of mothers, daughters, sisters, and aunts often overlap or are next to one another. These matrilines, as they are called, form because there are benefits to sharing resources with relatives. Though puma mothers and aunts do not physically “help” one another with raising cubs as human mothers might, in evolutionary terms there are advantages to sharing resources with relatives. Leopards and tigers also show similar tendencies for daughters to settle near their mothers, and of course lions have taken this behavior to the max—instead of living next door, mothers, daughters, sisters, and aunts get together and form prides that share a territory.

padding silently among village huts in their secret nocturnal search for dogs and goats. Like many of the smaller cats, leopards are superb climbers, agile and at ease in trees. The mark of a truly accomplished climber is the ability to come down a tree headfirst, and the leopard is one of only a handful of cats that are

skillful enough to descend a tree like this. They are also powerful and strong enough to carry large kills into trees, well away from hungry hyenas, wild dogs, and lions. In many parts of Africa, scavenging pressure from other meat eaters often forces leopards to protect their kills by taking them up into the trees. Any kill

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Leopards clearly have the ability to survive near humans. They can feed on almost any type of prey and can live almost anywhere, from the desert to tropical forest. However, their secretive habits make them extremely difficult to survey, so many current estimates of leopard numbers should be received with caution.

made in the open attracts vultures, which in turn attract other carnivores. In South Africa leopards use caves rather than trees as safe places to cache their kills. Fossilized bones found in these caves show that modern leopards and their ancestors have been living near humans and using caves for millions of years. Leopard-like cats stalked

and killed early hominids in Pleistocene Africa—some of the skulls of hominid fossils found in these caves have puncture marks that match the stabbing canine teeth of leopards. There are far fewer records of man-eating leopards than there are of man-eating tigers, but once a leopard has begun to prey on people, its stealth and boldness make it

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Infanticide: A grisly behavior, common in lions, where young are killed by new pride males Infanticide among animals is a grisly behavior that only makes sense when you understand the evolutionary implications. Ultimately, it’s all about ensuring that your genes survive. Most of the information on animal infanticide comes from studies of lions, where male lions commonly kill cubs when they take over a pride of lionesses—among Serengeti lions, infanticide accounts for a quarter of all cub deaths. Male lions have a relatively short tenure as breeding males. When they manage to seize control of a pride of lionesses, they kill all the young cubs sired by the previous group of males. If the cubs were allowed to survive, the females will not mate again until the young were at least eighteen months old, and the new males would “waste” that time investing in another male’s offspring. By killing the cubs, the new males ensure that the lionesses will come into estrus quickly, be ready to mate, and soon have a new set of cubs that carry their genes. Lionesses try to defend their cubs against infanticidal males because their genes are represented in all their cubs. Playback experiments show that lionesses can distinguish between the roars of their own pride males and the roars of unfamiliar males. When they hear the roars of unfamiliar males, lionesses with cubs become quite agitated and herd the cubs away from the approaching males. Groups of lionesses will sometimes gang up on new males to protect their cubs or lead older cubs away from the pride to prevent them from being killed. Lions are not the only carnivores in which males kill cubs after taking over a territory. The phenomenon is seen in tigers, leopards, pumas, and others—even free-ranging domestic cats. While infanticide occurs naturally in these cats, there is concern that sport hunting of the larger cats is increasing the rate of infanticide. Trophy hunters routinely select for adult males, with obvious consequences to the reproductive performance of resident females. In countries and states where sport hunting of large cats is high, the populations of these species show significant declines.

Males matter: When it comes to raising cubs, removing males is socially disruptive Though males of big-cat species don’t help raise cubs, they do provide vital security by protecting females and cubs from other males. Unstable male ranges do not provide a good environment in which females can raise cubs. In cat species in which the males kill young cubs, the evidence is now clear. Male “care” in the form of protection of cubs from other males is a vitally important factor to the survival of young. When an old male dies or is killed, new males routinely kill their predecessor’s cubs to bring the females into heat so they can sire the new male’s cubs. In places where territorial males are frequently killed, cubs rarely survive. In Nepal one huge male tiger monopolized the breeding in the central part of a park. In four years he and “his” seven tigresses produced twenty-seven cubs, almost all of which survived to adults. When he died, three or four new males fought for two years over his territory and breeding rights to the seven females. No cubs survived during those two years. The recent discovery of the importance of males in raising cubs throws a wrinkle into the previously straightforward calculation of the impact of hunting. It used to be thought that because males were not directly involved with day-to-day feeding and care of cubs, that it was okay for hunters to kill more of them. Males were considered “surplus” or superfluous. Now we know that regularly removing territorial males may mean females raise fewer cubs. Hunters usually seek out the largest males—lions with the biggest manes, the largest leopard, and the biggest puma. These large males are usually the territory holders, the ones who father the cubs and protect the females. The frequent and continued removal of these territorial males increases rates of infanticide, because new males regularly move in and kill cubs so they can father their own offspring.

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Distribution of the leopard.

a formidable killer, capable of claiming an enormous number of victims. Man-eating leopards are feared more than man-eating tigers because, unlike tigers, leopards will break into houses and huts to claim their victims. Man-eating leopards clearly develop a taste for human flesh and often selectively go after people even when there are vulnerable livestock around. In one instance, the infamous “Man-Eating Leopard of Rudraprayag” pushed open the door of a goat hut and made his way through a tightly packed flock of goats to get to a young boy who was looking after the animals. Modern leopards originated in Africa, somewhere between 470,000 and 825,000 years ago. Their spread across Asia was more

recent—170,000 to 300,000 years ago—and they are still found throughout most of their historic range. Today leopards must compete with poachers and bush-meat hunters for prey. But despite the competition, they still manage to persist in conditions where tigers and lions could not live. Ultimately, their flexible diet and ability to live unobtrusively in areas close to humans will likely make them the last big cat left standing. status: IUCN Red List—Near Threatened weight: 37–198 pounds (17–90 kg) head-body length: 36–54 inches (92–137 cm) tail length: 20–36 inches (51–91 cm) litter size: 1–4 cubs, 2–3 common

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Clouded Leopard N EOFEL I S N E B ULOS A

Clouded leopards are instantly recognizable by the large, distinctive cloud-shaped markings on their fur. This cat’s extraordinarily beautiful coat has made it highly sought after in the fur trade. Despite being a protected species, it is heavily hunted and trapped across its range. Skins are often seen in markets, and there is a significant trade in bones, meat, and even live animals. Though they are only the size of a small leopard, clouded leopards have the powerful, robust build of a big cat. Their large muscular head looks almost too big for their body, and with their massive jaw muscles, dagger-like canine teeth, and wide gape, they resemble a scaled-down saber-toothed cat. For its size, this cat has the longest canine teeth of any living felid. The upper canines of a clouded leopard can measure 1.5 inches (4 cm) or more—as long as a man’s little finger. As might be preInstantly recognizable by the large, distinctive cloudshaped markings on their fur, clouded leopards are powerfully built, with a large head and dagger-like teeth.

dicted from their long canines, clouded leopards kill fairly large prey, including bearded pigs, small deer, monkeys, orangutans, palm civets, and porcupines. The clouded leopard’s Malaysian name, harimau-dahan, means “branch tiger,” and these cats are well equipped to move easily through the trees; indeed, their climbing abilities rival those of the highly arboreal margay of South America. A long tail enables balance; large, broad paws provide grip; and short, stout legs give it a low center of gravity and the ability to wrestle large prey. The hind feet have flexible ankle joints that allow the foot to rotate, and the cat can climb slowly down a vertical trunk headfirst, move along horizontal branches while suspended beneath them like a sloth, and even hang from branches by its hind feet. Though clouded leopards are totally at home in the trees, they spend a great deal of their waking hours moving and hunting on the ground; apart from orangutans and monkeys, most of their prey is terrestrial. Since 61

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Clouded leopard: Modern saber-toothed cat?

Often referred to as “small big cats,” clouded leopards are the oldest members of the Panthera lineage, having separated from the “big cat” lineage several million years before the tiger, lion, and jaguar. Clouded leopards are famous for their beautiful cloud-like markings, odd body proportions, and long, dagger-like canine teeth. In addition to long canine teeth, clouded leopards also have several other unusual skull modifications that were also present in saber-toothed cats, including the ability to open their mouths to a nearly 90-degree angle. Both cats independently evolved these specializations to go along with their spectacular canine teeth as specializations for killing prey.

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Sunda clouded leopard: New species found on Sumatra and Borneo

For the last fifty years, the consensus was that clouded leopards were a single species, but in 2006 Andrew Kitchener at the National Museums Scotland, in Edinburgh, became intrigued by the difference in the size and alignment of the cloud-like markings on clouded leopard pelts. He went on to discover some fairly major differences in the coat patterns of clouded leopards from the islands of Borneo and Sumatra and those from the neighboring mainland. It turned out that the pelts of clouded leopards from the islands of Borneo and Sumatra have two or three rows of small cloud-like markings, many distinct spots within the cloud markings, and a double dorsal stripe. And their fur is darker than that of the mainland cats—the pelts of mainland animals have large backward-sloping cloud markings with fewer spots inside the clouds and only a partial stripe down the back. Subsequent morphological studies showed that clouded leopards living on Borneo and Sumatra also differ strongly from mainland cats in many skull and dental characteristics. Island clouded leopards have longer upper canines and thicker cutting teeth in the upper jaw—indeed, their skulls and teeth are more like the primitive saber-toothed cats. Recent molecular genetic studies have now confirmed that the clouded leopard is indeed two species, which diverged more than 1.4 million years ago. Though they look fairly similar to the untutored eye, these cats are as genetically different as lions are from jaguars. Since 2008 the Sunda clouded leopard (Neofelis diardi) has been listed as a separate species on the IUCN Red List, where it is classified as Vulnerable. The species is listed in CITES Appendix I, which bans international trade. The Bornean Clouded Leopard Programme aims to advance and enhance the understanding and conservation of all wild cats on Borneo. For more information on the organization, field updates, camera-trap photos, or to make a donation to their work, go to www.wildcru.org.

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Distribution of the Sunda clouded leopard.

scientists began systematic camera-trapping studies for carnivores, there have been many photographs taken of clouded leopards traveling on forest trails. The clouded leopard has a long moaning call that is reported to carry quite some distance. In Thailand clouded leopards are rumored to use elevated areas known as “tiger hills” to make these calls, which may serve to attract mates or keep animals apart by warning off competitors. They also have a typical repertoire of felid vocalizations; in friendly close-contact situations, they make a short, low-intensity, snorting call—a sound that has

been called “prusten.” Tigers, snow leopards, and jaguars are the only other cats that make a prusten call. Clouded leopards are believed to be an early branch of the “big cat” lineage that diverged from the ancestral Panthera line more than 6 million years ago. However, in 2006, in an unexpected taxonomic twist, scientists revealed that the cat we have always known as the clouded leopard is actually two distinct species, which are as genetically different as tigers and jaguars. All the evidence suggests that when sea levels rose during the Pleistocene, the clouded leopards on Suma-

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Distribution of the clouded leopard.

tra and Borneo became isolated from mainland clouded leopards and subsequently split into two species: the clouded leopard (Neofelis nebulosa) and the Sunda clouded leopard (Neofelis diardi).

status: IUCN Red List—Vulnerable weight: 24–51 pounds (11–23 kg) head-body length: 29–43 inches (75–108 cm) tail length: 22–36 inches (55–91 cm) litter size: 1–5, 2 cubs common

Bay Cat Lineage

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Bay Cat PARD OFE L I S B AD I A

The bay cat is found only on the island of Bor­neo. Until 1992 all the world knew about the bay cat came from nine skins and a few skulls that had been collected between 1855 and 1928. Virtually nothing was known about this felid except that it was about the size of a house cat and that its fur seemed to range from red to gray. Then, in November 1992, a cat appeared—the first live bay cat that any living scientist had ever seen. Caught by trappers, an adult female was brought to the Sarawak Museum in Kuching, Malaysia, emaciated and at the point of death. Knowing that the cat was rare, and probably valuable, the trappers had held the animal in captivity for several months while they tried to find an animal dealer who was willing to take it. Genetic material from this cat showed that In 2013 photographer Sebastian Kennerknecht worked for six weeks with biologist Andy Hearn on Borneo to get this first high-resolution photograph of the elusive bay cat. Before this extraordinary photograph was taken, the only images the world had of wild bay cats were a handful of low-resolution camera-trap photos.

bay cats are grouped with the marbled cat and the Asiatic golden cat. These three little-known Southeast Asian cat species form an ancient lineage that branched off early from the main stem of the cat family tree. The live bay cat trapped in 1992 confirmed that the species was still alive in the forests of Borneo. With the growing number of field biologists working on Borneo and the new technique of camera trapping, more bay cats were photographed on remotely triggered cameras—some of them during the daytime. In 1998 trappers captured another bay cat and sold the rights to photograph the animal to a U.S. photographer. The cat was reportedly “released back into the wild” after the photo session. In 2000 trappers captured another two bay cats, which were sold to an animal dealer. Both died before they could be exported. These cats were mounted as museum specimens and are now on display in the National Museum of Scotland. Local trappers and animal dealers in Bor69

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Ear talk: A cat’s mood is indicated by the position of its ears

All cats communicate with their ears. An alert, attentive cat holds its ears up and points them forward. A nervous or fearful cat lowers its ears. The more anxious the cat becomes, the more it flattens its ears, eventually rotating them so the tips point backward. Cats also communicate by twitching or flicking their ears. Watch two cats staring at each other, one will twitch its ear, then the other will ear flick and look away.

neo are clearly well aware that foreign zoos and breeding facilities will pay as much as $10,000 for a live bay cat. Though it is illegal to capture bay cats, the practice continues. Ironically, by creating a demand for live bay cats, Western zoos and captive-breeding facilities may ultimately become responsible for the extinction of this rare species. The Bornean Clouded Leopard Pro­gramme aims to advance and enhance the understanding and conservation of all wild cats on Bor-

neo. For more information on the organization, field updates, camera-trap photos, or to make a donation to their work, go to www .wildcru.org. status: IUCN Red List—Endangered weight: 6–9 pounds (3–4 kg) head-body length: 21–26 inches (53–67 cm) tail length: 12–16 inches (32–40 cm) litter size: Unknown

Believed to be extinct until 1992, the reclusive bay cat lives only on the island of Borneo.

Distribution of the bay cat.

Marbled Cat PARD OFE L I S M A RM ORATA

Though it is the size of a house cat, the marbled cat looks much bigger—a thick coat, extremely long bushy tail, and characteristic arched-back posture all combine to give the appearance of a much larger animal. With its broad feet and blotched coat pattern, the marbled cat resembles a miniature clouded leopard. Marbled cats live in lowland tropical forests of Southeast Asia, where they probably spend much of their time up in the trees. Large footpads help them grip branches and tree trunks, and their extremely long tail acts like a balancing rod. Marbled cats feed on birds, squirrels, rats, and the occasional monkey. In Sabah, Borneo, Azlan Mohamed and Andreas Wilting observed a marbled cat sitting on a thick branch some 82 feet (25 m) high in the canopy for ten minutes. As they watched with a spotlight, the cat began moving through The coat markings of marbled cats resemble those of clouded leopards.

the branches and then moved headfirst down the trunk of a large tree. Only the highly arboreal margay, the clouded leopard, and the leopard have previously been seen to climb headfirst down a tree. The Bornean Clouded Leopard Pro­gramme aims to advance and enhance the understand-

Catnip: Some cats respond to the chemical in the plant with crazy behavior All cat owners recognize the frenzied rolling, rubbing, licking, and generally crazy behavior of their cat playing with a fresh catnip toy. These hallucinogenic-like fits only last for five to ten minutes and are caused by an aromatic chemical in catnip called nepetalactone. Only about half of all cats respond to catnip; whether or not they react is under the control of a dominant gene. Catnip has been tested on cats in zoos, and so far tigers, lions, snow leopards, leopards and jaguars, servals, lynx, and ocelots have responded enthusiastically. Oddly, both captive and wild ocelots, jaguars, and cheetahs also find the scent of Calvin Klein’s perfume Obsession irresistible. Ocelots in particular are so attracted to the scent that scientists use it to attract the cats to camera traps!

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Spots: Most cats are blotched, striped, or spotted

South American Indian tribes believed that the jaguar made its own spots by dabbing mud on its body with its paws—if you look closely at jaguar spots, you can see what appear to be small toe marks. On the sides of the jaguar’s body, the spots form a series of dark rosettes that enclose one or two smaller dark spots. Leopards have rosette-like spots, similar to jaguars, but without the smaller spots inside the rosette. Cheetahs are covered with evenly spaced, solid black dots about the size of a human thumbprint. The black dots stand out against the coarser golden background fur. Interspersed with the black dots are slightly blurry-looking smaller spots. Desmond Varaday counted all the spots on his pet cheetah and came up with a grand total of 1,967. Serval spots are usually a series of single black dots on a tawny background, but they can vary from fine freckle-size dots to spots as large as a quarter. The serval’s spots sometimes merge into dark stripes on the neck and back. Ocelots have solid or open-centered dark spots that sometimes merge to look like links in a chain. The fur in the middle of the open-centered spots is often darker than the background coat color.

Though little is known of its biology, the marbled cat is thought to be highly dependent on tropical forest. For its body size, this arboreal cat has the longest tail of any cat species.

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Distribution of the marbled cat.

ing and conservation of all wild cats on Borneo. For more information on the organization, field updates, camera-trap photos, or to make a donation to their work, go to www .wildcru.org.

status: IUCN Red List—Vulnerable weight: 5–8 pounds (2.4–3.7 kg) head-body length: 18–24 inches (45–62 cm) tail length: 14–22 inches (36–55 cm) litter size: 1–2 kittens

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Asiatic Golden Cat PARD OFE L I S TE M M I N C KII

In China people believe that the golden cat is a kind of leopard and refer to its varying color phases by different names— those with black fur are called “inky” leopards, and those with spotted fur are called “sesame” leopards. Found in many Southeast Asian forests, the golden cat’s coat can be anywhere from golden brown to dark brown, bright red or gray, and unspotted or marked with spots. A hundred years ago, taxonomists used skull and teeth measurements, foot structure, and nose shape to order the cat family into species. When trying to classify the two golden cat species, taxonomists argued back and forth as to whether the two were “sister” species and should be in the same group, or if they should be classified in two different groups, maintaining that they just

The powerful-looking Asiatic golden cat feeds on monkeys, mouse deer, and birds in the lowland forest of Southeast Asia.

happened to look alike because they lived in similar tropical forest habitats. With the advent of molecular DNA technology, previously confusing relationships in the cat family became clearer. The Asiatic golden cat and the African golden cat are not nearly as closely related as scientists had previously assumed based on their similar looks. In fact, they are in separate lineages, having split from a common ancestor some 8.5 to 9.4 million years ago. Asiatic golden cats are usually found in forested habitats but are also occasionally recorded in scrublands, grasslands, and open rocky areas from sea level to 12,100 feet (3,700 m). They hunt at dawn and dusk, and sometimes during the day, and feed on birds, lizards, squirrels, monkeys, snakes, mouse deer, and barking deer. However, they have been recorded killing buffalo calves, sheep, and goats, indicating that these powerful-looking cats can kill prey larger than their own body weight.

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Distribution of the Asiatic golden cat.

Asiatic golden cats remain essentially unstudied in the wild—we know almost nothing about their biology and behavior. They show up surprisingly often in camera-trap studies, so they may be more common than previously thought. The little information we have from the few animals held in zoos suggests that they may breed at any time of

the year and typically have only one kitten in a litter. status: IUCN Red List—Near Threatened weight: 20–35 pounds (9–16 kg) head-body length: 26–41 inches (66–105 cm) tail length: 16–22 inches (40–57 cm) litter size: 1–3 kittens

Caracal Lineage

Serval

LEPTAI LURUS SE RVA L

As a light breeze shuffles the grass in Tanzania’s Serengeti Plain, a serval’s spotted shape dissolves and reappears, fading in and out of focus. The long-legged cat waits, still as a sentinel. Its eyes are almost closed as if in a trance, but its enormous, dish-shaped antenna ears slowly turn, listening intently. Minutes pass, then suddenly the ears focus, pinpointing the exact source of the sound. Two cautious steps and the cat springs, floating above the meter-high grass. Its body swerves and twists in midair, reorienting to each move of the rat. Two forepaws slam into the dirt. The pounce connects. Another meal is secured. Like the cheetah, the serval is one of the more specialized members of the cat family. Stalking through the grasslands on their stiltlike legs, servals look strangely un-catlike in The serval is a tall, lightly built cat with a small, slim face dominated by very large oval-shaped ears. Built for height rather than speed, the serval uses its long legs in combination with its large ears to capture small rodents in long grass.

shape. These tall, lightly built cats have a small, slim face dominated by very large oval-shaped ears. Rather than the sleek, muscular, crouching body plan of most felids, servals seem to have been designed for maximum height. For its size, the serval has the longest legs and feet of any member of the cat family; it stands as tall as a German shepherd dog, but weighs only half as much. Strictly speaking, it is not the length of the serval’s legs that allow it to tower over other cats, but rather the length of its feet. Elongated metatarsal bones in the palms and soles of the serval’s feet add extra inches to this cat’s height. Long legs usually mean speed, but the serval is not a particularly fast runner; rather, its legs are long so it can hear better. For sound-hunting to work most efficiently, ears need an elevated platform. Servals are so tuned into noises that they often stop and sit with their eyes closed for ten minutes or more, just turning their heads and listening. Unless they are extremely hungry, these cats rarely bother to hunt in windy weather—wind noise seems 81

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Serval pounce like a fox, springing high, then slam their paws down on the prey. Nearly half of all pounces results in a kill.

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to interfere with their ability to pinpoint the sound of prey moving through the grass. A serval pounces like a fox, springing high with all four feet off the ground. If the first attempt misses, the cat follows its prey— chin tucked into its chest, tail up, bouncing high into the air in a swift succession of stilllegged jumps. This capriole jump, in which all four feet are off the ground at once, is a characteristic serval hunting technique. Servals also use this high-bouncing pounce to flush animals from cover. They gallop through the grass in a zigzag pattern, leaping high into the air; anything that moves is caught immediately. Servals are prodigious leapers— their jumping abilities are equal to those of the caracal. A single pounce may span up to 12 feet (3.6 m), and they have been seen leaping 6 to 10 feet (2 to 3 m) into the air trying to knock down a bird or an insect in flight. Besides providing a vantage point for the serval to “hear into” tall grass, this cat’s long legs also pack a powerful punch. The serval uses its front feet to deliver a series of formidable blows that can kill or stun prey. When hunting larger animals that might fight back, the serval springs into the air, strikes with all four feet, bites, and then leaps away. Servals can kill quite large snakes by bashing them with a series of swift hammer-like blows. Not many cats are known for their digging abilities, but servals can, and often do, dig for their meals. Huge ears make their hearing so acute that they can locate prey underground, and they use their long mobile toes and strong claws to hook rodents and fledgling birds out of tunnels. One young serval was seen standing on his hind legs, system-

atically poking his front paw into the nest holes of a colony of swallows. Servals have also learned a highly refined “dig and wait” hunting technique that they use specifically to catch African mole rats. Mole rats live exclusively underground, so they are generally not accessible to cats. A serval locates a mole rat tunnel, scratches a hole in it, then sits and waits with one paw upraised—ready to capitalize on the fact that mole rats usually hasten to repair any damage to their tunnel system. At the first sign of any movement, the serval slams its paw down, hooks the mole rat, and flings it away, then follows with a lightning pounce before the stunned victim can recover. Whether it is feeding on frogs, mice, or fledg­ling birds, the serval’s big ears, un-catlike shape, and specialized hunting technique translate into more meals per pounce than the average cat. Most cats are lucky if they get a meal for one in every ten hunting attempts, but nearly one of every two serval pounces results in a kill, which makes it one of the most successful of all feline hunters. A single serval can kill about 4,000 rodents and 250 snakes in a year. Though they sometimes kill storks, flamingoes, and young antelope, servals survive on surprisingly small prey. Most of the serval’s diet is made up of animals weighing under 7 ounces (200 g)—less than 2 percent of an average female’s body weight. Servals hunt in the early morning and late afternoon, but females have to step up their efforts when they have kittens; mothers have to spend a major part of the day hunting. When the kittens are about a month old, the female begins to bring food back to the den.

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Servals are specialized sound-hunters, with the longest legs and feet of any cat. Long feet add extra inches to their height and provide their large ears with a raised platform to hear “into” the tall grass.

Biologist Aadje Geertsema, who studied servals in Tanzania, watched one serval mother carry a mouse for 1,750 feet (1.6 km) to the spot where her three kittens were hidden in some reeds. When she was about 165 feet (50 m) from the den, the female began to call, stopping periodically to listen for an answer. Near the den, the three kittens ran out meet their mother. One picked up the mouse, while the other two settled down to suckle. As the kit-

tens get older, it becomes more and more difficult for the mother to persuade them to stay at the den site while she goes hunting. Geertsema watched one mother spend more than an hour growling and spitting at her kitten before she could get away to hunt. The serval has been able to exploit high rodent populations in open grasslands and woodlands by virtue of its physical and behavioral specializations. These specializa-

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Distribution of the serval.

tions are very similar to those found in the dog family, a group that also preys extensively on small rodents in open habitats. With its large ears, small slim face, and long legs, the serval resembles a fox or a maned wolf rather than a cat. The hunting behavior of the serval is also rather doglike. The high-bouncing pounce, scanning for sounds, and digging are all hunting behaviors shared more with the dog family than with other members of the cat family. Like foxes, servals thrive on the rats and mice

that go hand in hand with agricultural activity, and they are quite common in rural areas in many parts of Africa. As long as they are not persecuted, these graceful, longlegged cats can survive in man-altered habitats and can even be beneficial to humans. status: IUCN Red List—Least Concern weight: 15–40 pounds (7–18 kg) head-body length: 24–36 inches (63–92 cm) tail length: 8–15 inches (20–38 cm) litter size: 1–4 kittens

Caracal CARACA L CARACAL

With its long legs and tufted ears, the caracal looks like an African version of the lynx, and for many years this cat was known as the caracal lynx. However, in 2006 DNA studies found that caracals are much older than lynx—the Caracal lineage broke off from the ancestral felid line more than a million years before the true lynx appeared. Caracals, it turns out, are not lynx, but instead are more closely related to the serval and African golden cat. For several hundred years, caracals were used by Indian nobility for sport hunting. They were trained to hunt small game such as hares and partridge, and each cat was carried out to the hunt on a horse, sitting on a small quilted pad behind the rider. A painting from 1602 of King Solomon’s court shows a tame caracal sitting in front of the throne

The caracal’s leaping abilities are legendary. Tame caracals were once used by Indian nobility for sport hunting, with the cats pitted against one another in pigeon-catching contests.

while two chukar partridges stand nearby; photographs taken as recently as the 1920s show tame caracals and cheetahs with their handlers, ready for the hunt. Caracals were still being sold in the marketplace in Amritsar until the 1940s. Covering parts of both Africa and India, the historic range of the caracal mirrors that of the cheetah, and both coincide with the distribution of several small desert gazelles. In many parts of the caracal’s present range, these 30- to 65-pound (15 to 30 kg) gazelles have either disappeared or are under intense pressure. Though caracals seem to be able to survive on hares and hyrax-size prey, they are clearly “gazelle cats,” as the nomads of North Africa have always called them. Cats are often said to have stern, aloof expressions or cruel, fierce eyes, and this description fits the caracal better than most felids. In strong light the caracal’s upper eyelid seems to cover the top half of its eye, giving it a narrow-eyed appearance that could be anthropomorphically described as cruel. 87

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The caracal resembles the

lynx, but DNA studies show that these cats are actually related to the serval and African golden cat. In Africa, caracals occasionally kill large antelope that can be two to three times their own size.

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Like the cheetah, caracals are found in parts of Africa and India, and the range of these cats coincides with the distribution of several small desert gazelles. Where these gazelles no longer occur, caracals survive on birds, reptiles, and mammals. In Ranthambhore National Park, India, this female caracal with two large cubs was feeding on a monitor lizard.

However, the lowered upper eyelid is probably a protective adaptation against the sun’s glare for this often diurnally active cat, rather than a reflection of the caracal’s fierce nature. When standing, a caracal appears taller at

the rump than at the shoulders because its hind legs are somewhat longer than its forelegs. The powerfully built hindquarters enable this cat to make spectacular leaps, sprint short distances at high speed, and climb well

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Amazing leaper: Only a few cats use a highflying leap to knock down birds in flight Unless they are chasing butterflies or grasshoppers, cats rarely leap into the air; they prefer to keep their hind legs firmly planted on the ground. This gives them a stable footing if the prey proves difficult to handle. The caracal is an exception. These cats are well known for the ability to jump—a tame caracal was lounging on the floor and when startled leaped 11 feet (3.4 m) into the air. This cat’s acrobatic abilities were recognized by Indian princes and potentates, who kept tame caracals for hunting. Tame caracals were pitted against one another in pigeon-catching contests, in which the cats were released into a flock of feeding pigeons while bets were made as to which one would bring down the largest number of birds. A skillful caracal could knock down nearly a dozen birds before the remainder of the flock escaped. This contest was almost certainly where the expression “to put the cat among the pigeons” originated.

Distribution of the caracal.

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when it needs to. It is an adaptable predator that can survive on a surprising variety of birds, mammals, and reptiles. Caracals kill birds ranging in size from quail to ostrich and will feed on just about any mammal from mice to hares and antelope. Most of their prey weighs less than 11 pounds (5 kg), but caracals are unusual among the small to medium-size cats in that they can and do kill prey that is two to three times their own size. They have been known to kill large antelope, sheep, and goats, and in some areas their appetite for domestic stock has resulted in them being regarded as a problem animal. In western India, where caracals are reported to be surviving on small rodents, these cats are clearly living on the edge. Despite its adaptable feeding behavior, a cat this size is unlikely to be able to sustain itself on rats and mice unless they are superabundant. When ro-

dents are not plentiful enough and gazelle-size natural prey have been eliminated, this large, powerful cat often turns to killing any available gazelle-size ungulate—sheep, goats, and other domestic stock—as they have in parts of the United Arab Emirates. Caracals are in the strange position of being endangered in the Asian portion of their range while being hunted as a problem animal in Namibia and South Africa, where they can be killed without restriction. The caracal is quite common in Israel, considered to be rare but holding its own in Pakistan, and on the verge of extinction in India. status: IUCN Red List—Least Concern weight: 14–40 pounds (7–18 kg) head-body length: 24–46 inches (61–106 cm) tail length: 7.7–13 inches (19–34 cm) litter size: 1–6 kittens, usually 2

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The powerful-looking African golden cat is built like a small leopard. Recent DNA studies reveal that these cats are closely related to the serval and caracal.

African Golden Cat CARACA L AURATA

The African golden cat is a powerful-looking cat, built like a small leopard. In parts of West Africa, these cats are sometimes called the “boy of the leopard” or the “leopard’s brother,” probably because of the likeness. African golden cats come in two distinct color phases—red and gray—but their fur varies so much through these two color morphs that they can look like a completely different cat in different regions of Africa. Their coat can be almost any color from plain orange-red to dark gray or even black, and anywhere along the continuum from plain and unspotted to heavily spotted. To add to the color confusion, the coat of an African golden cat in the London Zoo changed from red to gray over a period of four months. Golden cats are one of Africa’s least-known cats. They are found only in the dense tropical forests of Equatorial Africa, often along rivers and watercourses. They also seem to be able to survive in recently logged forest as long as plenty of heavy undergrowth remains.

They were once thought to be mainly nocturnal, but a recent camera-trap study, in which the cats tripped motion-sensitive cameras as they moved through the forest, revealed that they also hunt during the daytime. Analysis of their scats shows that they feed on a variety of rats, forest antelope, and birds. One study in the Central African Republic found that these cats sometimes hunt monkeys and duikers weighing 6 to 33 pounds (3 to 15 kg)—surprisingly large prey for a 22-pound (10 kg) cat. Because African and Asiatic golden cats look so similar, they were once thought to be close relatives, but new studies of felid DNA show that the African golden cat is more closely related to the caracal and serval. status: IUCN Red List—Near Threatened weight: 11–31 pounds (5–14 kg) head-body length: 24–39 inches (61–99 cm) tail length: 6–15 inches (15–38 cm) litter size: 1–2 kittens

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African golden cats are found only in the dense forests of Equatorial Africa.

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Distribution of the African golden cat.

Ocelot Lineage

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Ocelot

LEOPARD US PARDAL I S

Perfectly patterned to blend in with a world of dappled sunlight and shadows, ocelot fur is short, sleek, and marked with solid and open-centered spots that merge into lines. The subtle patterns of spots and chains of elongated dots on the ocelot’s coat vary so much that no two animals look alike. Ocelots live in a broad range of tropical and subtropical habitats from southern Texas and Arizona through Central America and as far south as Argentina. They are one of South America’s most frequently seen cats. Ocelots do most of their hunting on the ground at night, but they are occasionally active during daylight hours, especially when it is cloudy or rainy. They are strong swimmers, a not-unexpected ability for a cat that lives in places that sometimes flood for several months of the year. In Brazil scientists discovered that two radio-collared ocelots had swum across the swift Iguaçú The ocelot’s spotted coat blends perfectly into a world of dappled sunlight and shadows.

River four times. At night ocelots sometimes hunt in the open, but these cats basically depend on dense cover to stalk their prey. An ocelot needs about 1.5 pounds (680 g) of food a day to sustain itself, and most of this energy is obtained by preying on small abundant animals such as opossums, mice, rats, and rabbits. These cats are, however, also known to kill larger creatures such as agoutis, pacas, capybaras, sloths, armadillos, and even the occasional deer. During a night’s hunting, an ocelot crosses and recrosses its range, using the area intensively. They have two basic hunting strategies, a move-and-sit method and a slow hunting walk. With the move-and-sit method, the cat selects a spot where it can see the surrounding area, then crouches and waits for a rabbit or an opossum to come by. After half an hour or so, it walks quickly to another area, where it repeats the sit-and-wait. A cat using the slow hunting walk moves continuously, watching and listening for prey as it travels slowly and quietly along forest trails. 99

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Expensive taste: Some cats respond to expensive perfumes Knowing that trappers often use commercial scents to lure wild cats to traps, zoos have begun using “novel odors” to give captive cats something new to investigate in their enclosure. They found that some of the most expensive perfumes work just as well as the nastiersmelling traditional scents. Calvin Klein’s Obsession and Chanel No. 5 cause most cats to approach the scent, sniff, and twirl and head rub for two to three minutes. Scientists are now carrying the perfume into the jungle and sprinkling it around camera traps to encourage their subjects to have a picture taken.

The ocelot is found from Texas and Arizona in the north, to Argentina in the south. In some parts of its range, this cat is known as manigordo, meaning “fat hands,” because of its large front feet.

One radio-tracking study in Peru showed that bright moonlight hinders hunting. On fullmoon nights, ocelots avoid walking on trails and hunt only in dense brushy areas. Despite its perfectly camouflaged coat, it is difficult for an ocelot to creep up on a rat or a rabbit in bright moonlight unless there is plenty of vegetation to hide behind. Like margays, ocelots are unusual in that they have only one kitten per litter, and in the wild females have a litter only every other year. Kittens begin following their mother when they are about three months old and

remain dependent on her hunting skills for several more months. Young ocelots acquire their permanent teeth when they are about eight months old; but even after they have learned to hunt on their own, they sometimes continue to use their mother’s range until they are two years old. Compared with other cats of similar size, ocelots have longer pregnancies and smaller litters. Furthermore, ocelot kittens have the slowest growth rates of all the small cats— they do not open their eyes until they are fifteen to eighteen days old. If an ocelot female gives birth to her first litter at two and a half years old and raises one kitten every other year after that until she is twelve years old, she probably gives birth to only five to seven young in a lifetime. This is less than half the number of young a female bobcat might produce. The fact that ocelots have so few young in a lifetime makes them particularly vulnerable to hunting, deforestation, and other catastrophic events as populations are slow to recover.

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Distribution of the ocelot.

status: IUCN Red List—Least Concern

tail length: 10–17 inches (25.5–43 cm)

weight: 14.5–35 pounds (6.6–16 kg)

litter size: 1–4 kittens, usually 1

head-body length: 27–39 inches (69–100 cm)

Margay LEOPARD US WI E D I I

Margays are highly specialized arboreal cats. They live only in dense forests, where they do most of their hunting high above the ground, stalking small birds, squirrels, and rats among the branches. Agile and acrobatic, margays are the most talented climbers in the cat family. Broad velvety paws with flexible toes grab and hold even the narrowest branch, and a long tail helps them keep their balance as they move about in the treetops. Margays can run along a vine the diameter of your little finger or hang from their hind legs like a monkey. Very few people have been able to observe these cats in the wild, but one scientist studied the behavior of two captive four-month-old margay kittens. These young margays were able to jump 8 feet (2.4 m) straight up into the air and 12 feet (3.7 m) horizontally. Their reactions were so quick

The margay’s large eyes suggest they are strictly nocturnal, and most of the studies to date have found them to be active only at night. They have excellent eyesight and can spot a fly at 30 feet (9 m).

that if they fell while climbing, they could grab a branch with one paw and scramble up again. One of them could even hang by his hind legs while manipulating objects with his front paws. He also had excellent eyesight and could detect a fly at 30 feet (9.1 m)—leaping into the air, he would catch the fly with both paws, then draw it into his mouth before his hind feet touched the ground. Though in many places it lives alongside the larger ocelot, the margay is more specific in its habitat requirements than the ocelot. Margays live almost exclusively in humid evergreen forests and usually spend the day resting in a tangle of liana vines or the bole of a palm tree, usually at least 20 feet (6 m) off the ground. These cats hunt in the trees at night, but they also move on the ground when traveling between hunting areas. They kill whatever small prey they encounter while traveling between hunting areas or moving around in the trees. They eat cane rats, spiny pocket mice, squirrels, rabbits, mouse opossums, small birds, insects, and even fruit. Surpris103

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Known for their climbing ability, margays are the most agile and acrobatic of the cats. They are strongly tied to tropical forests and can exploit the upper levels of the canopy, where they hunt birds, small rodents, and insects. They sometimes eat fruit.

ingly, in captivity, margays readily eat figs and are also known to eat lettuce. The general rule for mammals is that the usual litter size is half the number of nipples, and this holds true for the margay—unlike other cat species, female margays have only two nipples and usually give birth to a single kitten in a litter. The kitten grows quickly and starts to eat solid food when it is about two months old. By the time the kitten is eight to ten months old, it is fully grown. Margays are members of the ocelot lineage, which separated from the ancestral felid line about 8 million years ago. The oce-

lot and margay are closely related, and genetic studies indicate that they split from a common ancestor that migrated into South America via the Isthmus of Panama about 3 to 5 million years ago. The margay looks like a scaled-down, bigeyed version of the ocelot, and both species have very similar spotted markings. Margays are smaller and more slimly built than ocelots and have a longer tail, but the pelts of the two species are easily confused. Fur traders have been known to take advantage of the similarity, and margay skins with the end of the tail cut off are sometimes traded as oce-

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Climbing cats: Accomplished climbers can descend headfirst

Really accomplished tree climbers like margays, clouded leopards, leopards, and marbled cats are able to climb down a tree headfirst like a squirrel. They can do this because they have the ability to rotate their ankles sideways, so they can grip the tree trunk equally well with their hind legs and forelegs. Instead of the narrow, firm feet of a domestic cat, these cats have extra large, soft feet with very mobile toes. These broad feet are a good platform for precision balancing and jumping, as well as an effective gripping surface for climbing and hanging on to branches.

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The margay looks like a scaled-down version of the ocelot. Both cats have similarly spotted coats, but margays have a longer tail and much larger eyes.

lot. One of the most striking differences between ocelots and margays is their fur texture. Ocelots have sleek, shorthaired coats, while margay fur is thick, soft, and longer than that of many of the other spotted cats. No one really knows why margays have such long fur, but they have an unusually low metabolic rate compared with other cats, so they may need the thicker fur coat to keep warm. The margay’s more arboreal lifestyle makes it much more dependent on forested habi-

tats, and thus more sensitive to deforestation. These characteristics, combined with the fact that females usually give birth to only one young at a time, make this cat a potentially vulnerable species. status: IUCN Red List—Near Threatened weight: 5–11 pounds (2.3–4.9 kg) head-body length: 20–31 inches (51–79 cm) tail length: 13–20 inches (33–51 cm) litter size: Usually 1 kitten

Distribution of the margay.

Geoffroy’s Cat LEOPARD US G EOFFROY I

Geoffroy’s cats have an unusual habit of sitting upright on their haunches or standing on their hind feet and using their tail like a tripod as a balancing aid. Even two-monthold kittens have been seen sitting on their haunches in this “prairie dog” stance. These versatile little cats are found in a wide variety of habitats in southern South America, including grasslands, dry steppe, open woodlands, and marshlands. They can climb but spend most of their time on the ground. Geoffroy’s cats are about the size of a domestic cat, but they have a shorter tail and a slightly flattened head. The background color of their fur varies from smoky gray to lion-colored and is dotted with small black spots, which in some places merge to form dark bands. Black individuals are fairly common. This beautiful coat has given the Geof-

In 1979 and 1980, more than a quarter of a million Geoffroy’s cat skins were traded in the international market. The species is now protected, and international trade is prohibited.

froy’s cat the unfortunate distinction of being the world’s second most frequently traded fur. Records show that in a two-year period (1979–80), more than a quarter of a million Geoffroy’s cat skins were traded in the international marketplace. Since 1994 Geoffroy’s cats have been protected under Convention on International Trade in Endangered Species (CITES) laws. International trade is now prohibited, and most of the hunting and trapping for the fur trade has ceased. While the international market is no longer a threat, rural people continue to kill Geoffroy’s cats for their skin, meat, or to reduce predation on poultry. In central Argentina, two-thirds of all Geoffroy’s cat deaths are caused by poaching and by free-ranging ranch dogs. Geoffroy’s cats are adaptable, flexible predators, and their diet varies from place to place. There are records of them feeding on mice, rats, hares, tree porcupines, guinea pigs, armadillos, viscachas, reptiles, and birds. In the desert-like dry rocky hills and creosote 109

About the size of a house cat, Geoffroy’s cats are adaptable, flexible predators. Their attractive fur and tractable nature have made them a popular cat among hybrid breeders—crossed with domestic cats, they produce hybrids known as Safari cats.

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Designer cats: The search for a wild-looking house cat

A famous beauty once approached playwright George Bernard Shaw and suggested they have a child together. “Just imagine,” she said, “a child with your brain and my looks.” Shaw replied, “Yes, but what if it has my looks and your brains?” Even the most knowledgeable and educated cat lover would admit to having a “wouldn’t it be fantastic” moment of longing, imagining how amazing it would be to see a tame, gentle mini-leopard strolling into your living room or sleeping on the couch. Not surprisingly, breeders continue to try to create a pet that has the exotic beauty of a wild cat combined with the gentle nature of a domestic cat. Domestic cats are mated with servals, leopard cats, fishing cats, and jungle cats to produce exotic-looking hybrids known as Bengals, Jambis, Safari cats, and Savannah cats, which sell for anywhere from $2,000 to $16,000 per kitten. Most hybrid kittens start out playful and friendly, and a few stay that way; but the vast majority of hybrids become more and more difficult to handle as they become adults. Neutered or not, by the time they are a year and half old, most hybrids spray urine and scent mark all over the house, just as they would in the wild. They bite, scratch, chew the furniture, fight with other pets, and are generally destructive. At this point, overwhelmed owners usually give up and call the pound or a cat rescue organization. So, every time you fantasize about having a pet wild cat, think of Shaw, and the reality of dominant genetic traits.

Man-Made Hybrid Cats • Savannah cat is a cross between a domestic cat and a serval. • Safari cat is a cross between a domestic cat and a female Geoffroy’s cat. • Bengal cat is a cross between a domestic cat and a female leopard cat.

• Caracat is a cross between a domestic cat and a caracal. • Jambi is a cross between a domestic cat and a fishing cat. • Chausie is a cross between a domestic cat and a female jungle cat. This hybrid is also called a Stone Cougar.

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Distribution of the Geoffroy’s cat.

flats of Argentina, two-thirds of their diet consists of vesper mice and grass mice. In a coastal lagoon in Argentina, they hunt migratory waterfowl; when the birds move on, the cats switch to feeding on mice and wild guinea pigs.

Surprisingly, Geoffroy’s cats can also kill relatively large animals. In the grasslands of southern Chile, more than two-thirds of their diet consists of 7-pound (3 kg) Cape hares, and on two occasions these cats were seen struggling to carry hare carcasses into

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trees. In Argentina a scientist watched a female Geoffroy’s cat wrestle with the body of a seriema—a large, long-legged coursing bird that resembles a secretary bird. The cat made several attempts to carry the bird up to a high branch but fell each time. An hour later the cat had managed to drag the chicken-size bird into a nearby burrow and could be seen sleeping behind its prey. Geoffroy’s cats have an unusual habit of leaving their scats high in the branches of trees, often 10 to 15 feet (3 to 4.5 m) above the ground. The fact that they go out of their way to do this suggests that the droppings serve as a scent or visual signal to other cats.

Geoffroy’s cats breed well in captivity, but they are not often seen in zoos. A recent boom in the popularity of hybrid cats has resulted in a heavy demand for captive animals. This cat’s beautiful pelt and its tractable nature have created a demand by private breeders and the general public for Geoffroy’s cat–domestic cat hybrids, known as “Safari cats.” status: IUCN Red List—Near Threatened weight: 7–15 pounds (3–7 kg) head-body length: 17–35 inches (43–88 cm) tail length: 9–16 inches (23–40 cm) litter size: 1–3 kittens

Guiña

LEOPARD US G UI G N A

The tiny South American guiña is not only one of the smallest cats in the world, but it also has one of the most restricted ranges. It lives only in a small part of coastal Chile and Argentina within an area roughly the size of the state of Texas. Until 2002 almost nothing was known about this miniature cat, but two recent studies have given us a glimpse into their lives. The guiña is the smallest cat in the Western Hemisphere; at 3 to 6 pounds (1.4 to 2.7 kg), it is about half the size of a domestic cat. Black guiñas are quite common; on mainland Chile, two-thirds of the cats captured during a radio-tracking study were black. Guiñas live in the cool rain forests of Chile and Argentina, and in the temperate beech forests of southern Chile, but they also use eucalyptus plantations and patches of trees between cultivated areas. They avoid open arThe diminutive guiña and oncilla share the distinction of being the smallest cats in the Western Hemisphere. They hunt small mice and birds.

eas and prefer habitats with thickets of dense brush and lots of stalking cover. Small mice and birds are the staple prey of this tiny predator. In the deep ravines and almost impenetrable bamboo forests of Chiloé Island off the coast of Chile, Jim Sanderson managed to radio-collar and follow several guiñas to see what they were feeding on. He discovered they hunted in the densest undergrowth, where they caught a variety of small birds, including thrushes, lapwings, and huet-huets. They also ate small mammals and lizards, and occasionally killed domestic chickens. Rachel Freer radio-tracked guiñas in the more open temperate beech forests of mainland Chile. She found that guiñas here also ate birds, but nearly three-quarters of their diet consisted of tiny South American field mice, Chilean tree mice, and 1-ounce (30 g) opossums. The cats were most active at dusk and spent about twelve of every twenty-four hours searching for prey—though they are good climbers, guiñas usually hunt on the 115

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Landing on their feet: A surprising number of cats survive falling from great heights Folklore has long maintained that cats always land on their feet, and a study by the Animal Medicine Center in New York has now confirmed the truth behind this legend. Doctors in the New York veterinary hospital studied 132 cats that had fallen from tall buildings— anywhere from the second floor to the thirty-second floor. Astonishingly, 90 percent of cats that fell survived, and two-thirds of them required no medical treatment. Cats survive these incredible falls because they turn their legs downward and extend their limbs outward, essentially assuming a gliding position like a flying squirrel. This prevents them from tumbling head over heels through the air while falling and saves them from hitting the ground headfirst. The legs-out position also changes the aerodynamic drag on the cat’s body and slows it down so it hits the ground with the least possible force. One of the record falls involved a cat named Sabrina, who fell from the thirty-second floor of an apartment building and landed on a concrete sidewalk. Sabrina walked away from the incident with a chipped tooth and a minor chest injury— an achievement not many other animals could match.

ground. They walk slowly, intensely searching a small area, then move rapidly to a new location and repeat the slow search pattern. Because this cat is so small, its skins have not been of much interest to the fur trade. However, it does kill chickens and is said to attack young goats, so it is hunted and killed wherever its presence is suspected. A questionnaire sent to local people in southern Chile revealed that most people held negative attitudes toward the guiña. On Chiloé Island, local farmers killed two of the seven radio-collared guiñas in Sanderson’s study

Guiñas are forest cats, active both day and night. These 4-pound (1.8 kg) mini-cats are hunted and killed whenever they are found near humans, since they are known to attack young goats and chickens .

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How cats “see” with their whiskers: Touch-sensitive whiskers work as “feelers” in the dark

Researchers have found that a blindfolded cat can negotiate a path across a room crowded with toys and other cats without running into any of them or even touching them. Cats are among a wide array of mammals with specialized touch-sensitive hairs—called vibrissae, or

because the cats were killing their chickens. In addition to conflicts with people, habitat destruction and logging in the ancient southern beech forests are probably major contributors to the guiña’s decline. The species is currently on CITES Appendix II and is fully protected in Argentina and Chile.

whiskers—that work as “feelers” for locating objects in darkness or low visibility. Whiskers grow on either side of the muzzle, around the eyes, and below the chin. Cats also have sensitive vibrissae on their ankles. Stouter than other body hairs and embedded more deeply in the skin, whiskers are extremely sensitive to movement. They rest in tiny sacs of fluid, pivoting like a straw in a soda bottle. When anything brushes the hair shaft, the information zips down to a rich supply of nerve endings that line the sac. Whiskers provide a sort of vision by touch, a tactile third eye. Cat whiskers are so sensitive that they can detect minor changes in air currents moving around objects, permitting blindfolded cats to avoid obstacles without even touching them. This sensitivity is critical to cats’ ability to hunt at night. By way of high-speed photography, researchers have found that a blindfolded cat can catch and kill mice as long as its whiskers are intact. A cat on the prowl holds its whiskers out on either side of its face like a fan. Just before pouncing on its prey, the cat shifts its whiskers forward, extending them like a net in front of its mouth. When the cat makes contact with its prey, the whiskers convey exactly which way the mouse is dodging in the final split second. A cat carrying a freshly caught mouse wraps its whiskers around the prey, sensitive to any twitch warning that the mouthful would squirm free.

status: IUCN Red List—Vulnerable weight: 3–6 pounds (1.3–2.5 kg) head-body length: 15–20 inches (37–51 cm) tail length: 8–10 inches (20–25 cm) litter size: 1–4 kittens

Distribution of the guiña.

Andean Cat LEOPARD US JACOB I TA

Until the year 2000, virtually the only proof that the Andean cat still existed came from the occasional appearance of fresh skins in the Buenos Aires fur markets and half a dozen chance sightings of this cat in the wild. These few early sightings all noted that the cat ignored the presence of humans and allowed observers to approach within a few yards. The Andean cat’s characteristically nonchalant behavior around people seems to have remained unchanged. In 1998 Jim Sanderson was following up on an earlier sighting of the Andean cat in Chile when he came face-to-face with one right outside his camp. Sanderson was able to follow and photograph the cat for more than five hours—even when he approached within 6 feet (2 m), the cat ignored him; it stretched, Andean cats are one of only four cat species listed as Endangered. Their long thick fur and fluffy tail are adaptations for surviving in extremely cold temperatures. The Andean cat lives at high altitudes in the Andes Mountain chain from Peru to Argentina.

yawned, urine-sprayed a rock, slept for a while, then resumed its search for mountain viscachas as he watched. In 2004 a park ranger in the Caverna de las Brujas Provincial Reserve in Argentina came across two Andean cats in a small cave at the foot of a cliff. He was also able to watch and photograph the cats—an adult and a kitten— for more than thirty minutes. The Andean cat’s unusually fearless nature has provided some amazing photo opportunities for researchers, but it has also had fatal consequences. The cat’s unafraid behavior makes them highly vulnerable to persecution, and there are many accounts of villagers and reserve guards killing these cats by stoning them. The Andean cat is about the size of a large house cat. The cat’s long, thick fur is pale silvery gray, spotted, and striped with dark markings. The tail is long and fluffy, like the tail of a snow leopard, and banded with about seven dark rings. Andean cats live high in the Andes Mountains of Argentina, Bolivia, Chile, and Peru, on open, tree121

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The new scoop on poop: New survey technique relies on identification of individuals from DNA in scats At one time field biologists picked through cat scats looking for feathers, hair, and bones so they could figure out what the cat had been eating. Recently, cat scats have become even more informative, because a new technique using the unique DNA signature on the scat is allowing scientists to identify individual cats. When food passes through the gut of an animal, it collects a thin layer of cells from the intestine. The DNA in these cells can now be collected and analyzed, and each scat identified and matched to an individual cat. This gives biologists the ability to discover what each cat has been eating, whether there are differences in the diets of males and females, and, even more importantly, reliably count the number of individual tigers, leopards, or jaguars in an area.

The Andean cat’s life is tied to the presence of viscachas—rabbit-size rodents who live in colonies and make their homes in rock piles.

less, rocky slopes, usually between 9,800 and 16,500 feet (3,000 and 5,000 m). At these high altitudes, daytime temperatures are often around 32° to 39°F (0° to 4°C), freezes are frequent, and water is scarce. There

are, however, recent records of the cat from much lower elevations in Argentina. The life of the Andean cat is closely linked to the presence of mountain viscachas—a rabbit-size animal that lives in large colo-

a n d e a n c a t   1 23

Distribution of the Andean cat.

nies. Andean cats typically hunt in a viscacha colony for a few days, then move on to another colony several miles away. To successfully raise kittens, a female needs an area that encompasses at least one or two large, productive mountain viscacha colonies. The only Andean cat that has been radio-collared to date was a female with a home range of 18 square miles (47 km2). Males likely roam over even larger ranges that encompass the smaller ranges of several females. Like the closely related Geoffroy’s cat, Andean cats leave their feces in latrines. This

convenient habit and the new science of “molecular scatology” have allowed researchers to discover some of the finer details of their diet. While mountain viscachas make up more than 90 percent of the Andean cat’s diet, the cats also eat leaf-eared mice, chinchilla rats, and quail-size birds called tinamous. In less than a decade, the Andean cat has moved from being an almost completely unknown species to a model for cat conservation. In 2002 the Andean cat was moved from Vulnerable to become one of only four cat species listed as Endangered. Despite the fact that the IUCN classifies the Andean cat as an Endangered species, the long-term conservation prospects for this species are encouraging. In late 1999 a highly motivated, multinational group of biologists from Argentina, Bolivia, Chile, Peru, and the United States founded the Andean Cat Alliance, or Alianza Gato Andino. The group rapidly developed a Conservation Action Plan to provide information on the Andean cat’s distribution and basic biology, to deal with specific threats, and most importantly to involve local people in conservation actions. For more information on the project, field updates, and new camera-trap photos, go to www.gatoandino .org; or donate to the research via http:// wildlifeconservationnetwork.org/support/. status: IUCN Red List—Endangered weight: 7–15 pounds (3–7 kg) head-body length: 22–26 inches (57–65 cm) tail length: 16–18 inches (41–48 cm) litter size: 1–2 kittens

Oncilla

LEOPARD US TI G RI N US

Oncillas look a lot like margays, but they are smaller and more delicately built, with larger ears and a narrower muzzle. Until a few years ago, the oncilla was thought to be strictly a forest cat, confined to lowland and cloud forests; but recent information from camera traps shows that they also thrive in wet and dry savannas and thorn scrub. As might be expected, this 6-pound (2.7 kg) cat hunts very small prey. Oncillas eat birds, small mice and rats, and lizards, as well as centipedes, grasshoppers, beetles, and small opossums—most prey weigh less than 3.5 ounces (100 g), or roughly the size of a mediumsize tomato. In forested habitats, oncillas hunt at night, but in savannas and scrub forests, they hunt during the day. Like the margay and ocelot, oncillas commonly give birth to a single kitten after a long gestation period. This makes them more vul-

Cats are “sugar blind”: Cats lack the gene to taste sugar All cats, from tigers to tabbies, lack one of the receptors that allow animals to detect sweet tastes. Unlike dogs and most other mammals, cats are “blind” to the taste of sugar. Sweet compounds are recognized by a special taste-bud receptor controlled by two genes, one of which is nonfunctional in cats. This “sugar blindness” probably played an important role in the evolution of the cat’s strictly carnivorous diet.

nerable to hunting and trapping, because populations take a long time to recover from losses. A recent genetic analysis suggests that the oncilla may be two species. status: IUCN Red List—Vulnerable weight: 4–8 pounds (1.8–3.5 kg) head-body length: 15–23 inches (38–59 cm) tail length: 8–17 inches (20–42 cm) litter size: 1–4 kittens, usually 1

The tiny, delicate-looking oncilla resembles a smaller version of the margay. Like ocelots and margays, oncillas have only a single kitten in each litter. 125

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o n c i l l a   1 27 Opposite, top. In 2013 Brazilian scientists studying the genetics of the South American Leopardus cat group proposed that oncilla from northern and southern Brazil should be two different species. The northern and southern populations are genetically distinct and have been separated for at least 100,000 years. Common names have yet to be decided for these new species. Opposite, bottom. Oncilla from both areas are the same size and look almost identical, but the northern oncilla has slightly lighter fur marked with smaller rosettes. Scientists anticipate that as more genetic information becomes available from other parts of the cat’s distribution, we may see a multitude of oncilla species.

Distribution of the oncilla.

Pampas Cat LEOPARD US COLOCOLO

Pampas cats look like a burly domestic cat, and their long hair makes them look bigger than they really are. On some individuals the long hairs on the back form a sort of mane, which can be erected when the cat is frightened or nervous, giving it a larger and more formidable appearance. These wild cats are fairly common in South America in dry open grasslands and high deserts from Ecuador and the Brazilian highlands in the north to Patagonia in the south. They feed mainly on small mammals but are known to attack domestic poultry and have even been seen raiding the nests of Magellanic penguins in Patagonia. Pampas cats are flexible predators, varying when and where they hunt, what species they hunt, and changing these patterns as circumstances warrant. In Argentina’s high-altitude

The pampas cat looks like a heavyset house cat, and their long hair makes them appear larger than they really are. Pampas cats are common in South America’s dry open grasslands and high deserts.

deserts, pampas cats hunt at night, feeding on leaf-eared mice, birds, and a pocket gopher– like rodent called the tuco-tuco. In this same area, they seem to have worked out a timeshare arrangement with the slightly larger Andean cat, which hunts during the daytime for rabbit-size rodents known as mountain viscachas. In northern Chile’s bleak altiplano, the same two cats share an arid treeless area of exposed rocky terrain dotted with high-altitude lagoons. Both species depend on mountain viscachas for basic survival, but the pampas cat spends more time hunting migratory waterfowl and flamingoes around the wetlands, while Andean cats focus their efforts on the steep rocky areas, where they search for mountain viscachas. Molecular analysis by scientists in Steve O’Brien’s Laboratory of Genomic Diversity in Maryland shows that the pampas cat and Andean cat are “sister species” and diverged from the other small South American cats some 2 million years ago. Further, it has been suggested, based on differences in skull and 129

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Distribution of the pampas cat.

skeleton measurements of cats from different regions, that geographic isolation has since divided the pampas cat into three separate species. Given the cat’s distribution and the fact that populations are separated by high mountain ranges and large river systems, the potential for speciation certainly exists. However, genetic analysis indicates that the pampas cat is still a single species.

status: IUCN Red List—Near Threatened weight: 6–9 pounds (3–4 kg) head-body length: 16–31 inches (42–79 cm) tail length: 9–13 inches (22–33 cm) litter size: 1–3 kittens

Lynx Lineage

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Eurasian Lynx LYN X LY N X

About 2 million years ago, a puma-size cat known as the Issoire lynx roamed Europe and northern Asia. This ancestral lynx had a larger head and shorter, sturdier legs than modern lynx, and its teeth suggest it fed on roe deer–size prey. The Issoire lynx is believed to be the forefather of the Eurasian, Canada, and Iberian lynx and the bobcat. When lynx from Eurasia reached North America, there were no roe deer–size prey available. The most abundant prey was the much-smaller snowshoe hare. Consequently, the animal that we now know as the Canada lynx became a specialized hare hunter, and all aspects of its lifestyle are now closely tied to the snowshoe hare. The Iberian lynx—another descendant of the Issoire lynx—remained in Europe but retreated to Spain and Portugal ahead of the glaciations. Over time the Iberian lynx also became smaller, and it, too, specialized on hunting rabbits. Eurasian lynx are twice the size of other lynx; a large male can weigh 65 pounds (30 kg).

Ear tufts: Signaling devices?

Why some animals have ear tufts has always been a bit of a mystery. Dozens of different mammals from squirrels, marmosets, African bush pigs, and lynx have ear tufts, as do rockhopper penguins and owls. Just about everything from camouflage to hearing aids has been proposed as likely functions of ear tufts, but no one has yet to come up with a plausible unifying explanation. Among the cats, ear tufts are likely used as signaling devices to enhance communication. Caracal, bobcat, and Eurasian, Canada, and Iberian lynx all have pronounced ear tufts, and, coincidentally, these are all short-tailed cats. Cats use their tails to convey information, and it may be that in the absence of a long tail, ear tufts have assumed a more important role in communication among these species.

The Eurasian lynx is the only one of the four modern lynx that has retained the size and many of the characteristics of the ancient Issoire lynx. It is twice the size of other lynx—a large male can weigh 65 pounds (30 kg)—and looks like a bobcat, except that it is much larger, longer-legged, and has bigger feet. Eurasian lynx have a distinctive tall “leggy” 133

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e u r a s i a n l y n x   1 35

Unlike the other lynx species, which feed on rabbits and hares, Eurasian lynx feed primarily on roe deer.

look and a rather odd, tipped-forward stance; when they move, their hind legs are noticeably longer than the front legs. In the Lynx family, long hind legs and short front legs are a compromise between speed and strength. The long hind limbs help them reach sprinting speed more quickly and allow them to run fast when chasing prey. But a cat’s forelimbs are used mainly for close-quarters wrestling with a panicked prey; for the Eurasian lynx, which regularly kill deer, shorter, stronger front legs help them catch and hold larger prey. Like the Canada lynx, Eurasian lynx have Lynx spend the day hidden in dense cover and emerge to hunt in late afternoon. They are most active at dawn and dusk.

wide paws with well-developed webbing between the toes, and in the winter the undersides of the paws are covered with long, dense, shaggy hair—all adaptations for moving easily through deep, soft snow. These lynx are forest cats, and their vast geographic range essentially mirrors the range of the roe deer. Across much of Europe and Asia, the single most important prey in the Eurasian lynx’s diet is the 45-pound (20 kg) roe deer, but these cats also eat reindeer calves, wild pigs, young red deer, hares, and other small creatures. Lynx spend the day hidden in dense cover and emerge in late afternoon. Most active at dawn and dusk, they hunt intermittently throughout the night. Lynx have excellent vision and hearing. One scientist was able

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Chamois are another favorite prey of Eurasian lynx. Unfortunately, chamois are also popular game animals, which brings the lynx into conflicts with hunters.

to test the abilities of two hand-reared lynx and found that they could see a mouse at 245 feet (75 m), a hare at 985 feet (300 m), and a roe deer at 1,640 feet (500 m). While hunting, lynx follow well-used trails made by deer or hares, frequently stopping to survey their surroundings from the tops of boulders and downed trees. These cats generally avoid water and cross streams by walking along fallen trees or leaping from rock to rock. Eurasian lynx are capable of living close to humans, but they are cautious, timid cats, and history shows that they have not fared well in their relationship with people. They

were once found across Europe, but by the end of the nineteenth century, this cat had been eradicated over much of central and southern Europe due to a combination of deforestation, persecution, and loss of prey. Some lynx populations survived in Scandinavia, Russia, the Balkans, and the Carpathian Mountains. Efforts to reintroduce lynx to Europe began in the 1970s, and over the last forty years lynx have been released at more than twenty sites within their former range. The results of these efforts have been mixed. Reintroductions in Switzerland, Slovenia, and the Czech Republic have been quite successful,

e u r a s i a n l y n x   1 37

Distribution of the Eurasian lynx.

with lynx expanding into adjoining forested areas, but most populations have remained small and isolated by habitat fragmentation. While lynx numbers have increased and they are now found in more areas, they have recolonized less than 20 percent of the Alps. As the populations of reintroduced lynx have expanded, so have the concerns of farmers and hunters. Lynx predation on domestic sheep, roe deer, and chamois has caused conflicts with livestock breeders and hunters. Financial compensation for livestock losses has worked fairly well to reduce conflicts with

sheepherders, but the other major conflict is with hunters, who contend that lynx diminish roe deer and chamois numbers. This antagonism leads to illegal killing of lynx. Resolving this conflict will go a long way toward the recovery of Eurasian lynx populations. status: IUCN Red List—Least Concern weight: 37–66 lb (17–30 kg) head-body length: 35–47 inches (90–120 cm) tail length: 7–9 inches (18–23 cm) litter size: 1–4 kittens

Iberian Lynx LYN X PARD I N US

The Iberian lynx is the world’s most endangered felid and the only wild cat to be classified as Critically Endangered. About 250 individuals remain in the wild in two small, isolated populations in southern Spain. Over the past twenty-five years, lynx numbers have been in free fall, and despite current Herculean efforts to rescue the species from extinction, the Iberian lynx is likely to be the first cat to go extinct since the European cave lion. Some 2 million years ago, a lynx-like cat known as the Issoire lynx roamed Europe and northern Asia. Recognized as the common ancestor of all of today’s lynx, the Issoire lynx had shorter legs than modern lynx and, judging by its jaw morphology and teeth, was more of a generalist predator, living on roe deer–size animals. The Critically Endangered Iberian lynx is a rabbit specialist, and the cat’s numbers have plummeted as disease and habitat change have decimated rabbit numbers throughout the Iberian Peninsula. Only 250 Iberian lynx remain in the wild.

About 1 million years ago, when the quaternary ice ages swept south through Eurasia, wildlife retreated in advance of the cold, moving south to warmer parts of Europe. The Iberian Peninsula—Spain and Portugal—became a refuge for a variety of species, including the lynx. At about the same time, the European rabbit appeared in the fossil record of the Iberian Peninsula. All indications are that this now-widespread rabbit originated in the Iberian Peninsula and, indeed, until 10,000 years ago was found only in Spain and Portugal and southern France. Isolated in the Iberian Peninsula with this new abundant source of food, the lynx began to focus on rabbits, which at that time were larger than their present-day relatives. Over time the cat became a rabbit specialist, and its distribution mirrored the natural range of the European rabbit. Today the Iberian lynx is a unique species, somewhat smaller in size than the ancestral Issoire lynx and now completely dependent on rabbits. Eighty to 99 percent of 139

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the Iberian lynx’s diet consists of rabbits, and this cat has become a virtual prisoner of its specialized diet. Rabbits are the ideal prey for a medium-size carnivore. Rabbits are relatively large and not too difficult to catch, have several litters a year, and live at high densities. Lynx need to kill about one rabbit per day to sustain themselves and cannot live in places where rabbits are absent or scarce. The Iberian lynx might have been better equipped to survive twenty-first-century life in Europe if it had not become a rabbit specialist but had remained a more generalized carnivore like its ancestor. Today habitat change and disease—first myxomatosis in the 1950s, then rabbit hemorrhagic disease in the 1980s—have combined to reduce the Iberian rabbit population by 95 percent since the 1950s. Despite its legendary ability to procreate, the once-abundant European rabbit is now languishing. The IUCN has officially classified the rabbit as Near Threatened in Spain and Portugal, and sadly the fate of Iberian lynx is tied to its dwindling prey. Iberian lynx are about twice the size of a domestic cat but with longer legs. They stand 15 to 20 inches (40 to 50 cm) at the shoulder and weigh 17 to 35 pounds (8 to 16 kg). Like all lynx, they have a short tail, a short body, tufted ears, and a relatively small head. Both sexes have prominent facial whiskers and ears tipped with long, erect tufts of black hair. Not surprisingly, lynx are closely associated with scrublands, meadows, and open forests—all habitats that are favored by

rabbits. Breeding lynx females need safe places for birth dens, such as hollow trees or rock cavities, and a regular supply of water. While nonbreeding lynx can survive on a mix of prey, breeding females require an abundant and reliable source of rabbits. Litters of two to three kittens are born in March or April. Young are born blind and helpless, unable to regulate their own body temperature, so for the first few weeks the mother must spend a great deal of time in the den, nursing the kittens and keeping them warm. Kittens are born with their ears folded and tiny tufts of black hair on the tops of their ears. Mothers tend to keep their kittens in the birth den for about a month, then begin moving the young to different dens every few days. Kittens become highly competitive when they are about six or seven weeks old; they fight intensely, and siblings sometimes kill one another. With the help of the international conservation community, Spain and Portugal have mounted extensive efforts to save the Iberian lynx. Experts believe that if the lynx is to survive, conservation efforts will have to include habitat restoration, reintroductions, supplementary feeding, and captive breeding. Many of the cats have been killed while crossing roads, so wildlife underpasses have been constructed to reduce road kills. In 2003 a large-scale captive-breeding program was initiated. Though there have been several setbacks, at least forty-six kittens have been born, and in 2009 sixteen young lynx were sent to Portugal to start a separate breeding program. Also in 2009 the first cats were released from the captive-breeding pro-

i b e r i a n l y n x   141 Distribution of the Iberian lynx.

gram into the wild near Córdoba. Provided with supplemental food in the form of rabbits, the lynx survived and bred. Further releases are planned as more lynx become available from the breeding efforts. The current captive population includes about eighty-four breeding females.

status: IUCN Red List—Critically Endangered weight: 17–35 pounds (8–16 kg) head-body length: 25–36 inches (65–92 cm) tail length: 4–6 inches (11–16 cm) litter size: 1–4 kittens

Canada Lynx LYN X CA N AD E N SI S

Canada lynx and bobcats look so much alike that at one time it was believed that the bobcat was just a “southern” version of the Canada lynx. Now we know that these two cats are the product of two separate invasions of North America; the bobcat has been in the United States for more than 2 million years, while the ancestors of the Canada lynx moved from central Asia across the Bering Strait and colonized Alaska and Canada only 200,000 years ago. The tall, leggy Canada lynx is a highly specialized hare hunter and one of the few cats that do not have the usual felid body shape. Though they are only about twice the weight of a domestic cat, Canada lynx have disproportionately long legs, and their much longer hind legs give them a tipped-forward appearance. These long hind limbs are an adaptation The lynx is managed and trapped for its fur across much of Canada, and the harvest is regulated through trapping concessions, closed seasons, and quotas. Included in these regulations is the establishment of refuge areas where lynx are not trapped.

for running and bounding through snow just like their snowshoe hare prey. Canada lynx also have specialized paws to help them move more easily on top of powdery snow. Their footprints can be as large as the palm of a man’s hand. Canada lynx get their big paws from the elongated bones in their feet. Their phalanges—similar to our finger bones—are long and can be spread to create a larger foot surface. Combined with dense, coarse hair on the bottom of the feet, these long toes create a snowshoe-like foot, twice as large as the foot of the similar-size bobcat. These big feet support the lynx’s weight on top of the snow just as a snowshoe supports a person. The bobcat’s smaller, narrower feet sink into the snow and hamper their movements; they simply cannot match the lynx’s specializations to hunt in deep snow. The lynx’s ability to move in deep snow allows them to live and hunt in places where bobcats cannot. However, in the few places where the lynx and bobcat overlap, the lynx does not fare well in competition with the 143

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The tall, leggy Canada lynx is a specialized hare hunter. Elongated bones in their feet allow them to spread their toes to form a larger surface, creating snowshoes for moving through deep, soft snow.

The distribution of the Canada lynx mirrors that of the snowshoe hare, and at times it eats little else. The availability of hares influences every aspect of the lynx’s life.

more aggressive bobcat. Any climate change that reduces snow depth and allows bobcats to move farther north will not bode well for lynx. The geographical distribution of the Canada lynx mirrors the distribution of the snowshoe hare, and hares make up most of this

cat’s diet—at times they eat little else. During the summer, lynx add birds, voles, meadow mice, and squirrels to their diet, but most of the meat they consume still comes from hares. Trapping records of the Hudson’s Bay Company stretch back to the early 1800s and show that lynx populations typically cycle through highs and lows every ten years. The pattern is roughly synchronous from east to west across northern North America and tracks the dramatic changes in hare abundance. At the peak of the cycle, hare densities can reach 2,300 animals per square kilometer, and then in the low, a few years later, they plummet to 12 hares in the same size area. Lynx numbers rise and fall with hare numbers. When hares are at their most abundant, it is not unusual to see lynx females with as many as six kittens, most of which survive to adulthood. When hare numbers crash, few young lynx

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Distribution of the Canada lynx.

survive, and adults abandon their ranges and move long distances—as far as 310 to 620 miles (500 to 1,000 km). Snowshoe hares feed at night, and this is when lynx do most of their hunting. They need to find and kill one hare every day or two to survive. A hunting lynx zigzags and circles back and forth in a small area, probing patches of cover in an attempt to flush a hare. At other times they sit or lie in wait, sometimes long enough for the snow to melt and form an ice-encrusted impression of the cat’s body. Ambush hunting from these “snow beds” may be a way for the lynx to conserve energy when hares are scarce. Canada lynx are unusual among cats in that they sometimes cooperate by hunting in groups—in most cases the groups are mothers hunting with their almost adult-size young,

but sometimes two or three adult lynx will hunt together, especially when hare numbers are low. Biologists have pieced together the information from snow tracking, and there are even YouTube videos showing groups of two to four lynx walking single file across a forest opening, then fanning out and moving through the area on a common front, flushing hares to other group members. Lynx are trapped for their fur across much of Canada, and the harvest is regulated through trapping concessions, closed seasons, and quotas. status: IUCN Red List—Least Concern weight: 11–38 pounds (5–17.3 kg) head-body length: 29–42 inches (74–107 cm) tail length: 2–5 inches (5–12.7 cm) litter size: 1–8 kittens

Bobcat LYN X RUFUS

Bobcats were once thought to be a southern form of the Canada lynx; however, fossils and DNA studies show that the bobcat’s ancestors entered North America long before the ancestors of the Canada lynx. Even though they look almost identical, the bobcat and Canada lynx evolved separately. And while bobcats have been present in North America for nearly 2 million years, Canada lynx are a relative newcomer to the continent, having been in North America for only 200,000 years. Bobcats and Canada lynx are very similar in appearance and quite difficult to tell apart. Both have tufted ears, relatively small heads, and long legs. However, the bobcat has smaller feet, and its short tail is black on the upper surface and pale colored underneath. Unlike lynx, bobcats are not well adapted to living and hunting in deep soft snow, and they also have a hard time regulating their body temperature in cold weather. A combiHunting and trapping for the fur trade almost extirpated bobcats from the midwestern United States in the 1970s, but populations have recovered due to legal protection.

nation of deep snow and cold temperatures most likely limits the bobcat’s northern distribution. Bobcats are generalists, both in their habitat and diet. They are good climbers and will rapidly climb a tree if pursued by dogs. They can also swim if they have to but don’t often venture into water. Bobcats prefer areas with dense cover or uneven, broken terrain, and they hunt just about anything, including rabbits, hares, opossums, rats, and mice. Bobcats are also serious predators of fawns, and surprisingly deer are a major part of their diet, especially in the northeastern United States. Though they mostly eat smaller animals, these cats can single-handedly kill an adult deer more than ten times their own size. Bobcats also kill chickens, turkeys, sheep, and goats, but they are not known to be serious predators of livestock. On a few occasions, they have been known to cause havoc in a henhouse or lambing pen by killing large numbers of chickens or lambs in a single night. These cats usually hunt while walking 147

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Easily recognized by its short tail, the bobcat is the most abundant and widespread wild cat in North America; it looks very similar to the Canada lynx but has much smaller feet.

slowly along trails, stopping to listen intently to any sound. The mere flicker of a feather or the slight turn of a tuft of fur caught on a branch will attract their attention. Fur trappers capitalize on this by using tufts of fur,

feathers, or a dangling piece of silver foil to lure bobcats to a trap site. Bobcats are prolific breeders. They have large litters and can give birth to twenty-five to thirty kittens over a lifetime. The similar-size

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Why cats can’t be vegetarians: Cats are obligate meat eaters A cat’s body is, in essence, a reflection of its diet. Stabbing and shearing teeth, sharp claws, and a short intestine adapted to process meat—this is the body plan of a true hyper-carnivore. Many of the vitamins and amino acids cats require can only be obtained from animal sources. Unlike humans and dogs, cats cannot manufacture taurine in their bodies, and they cannot obtain other key nutrients in sufficient quantity from plant-based food. Cats also require a higher proportion of protein in their diets than almost any other mammal. Domestic cats need a diet that contains at least 12 percent protein—kittens need 18 percent protein. This is why it is very difficult to create a vegetarian diet that will keep a domestic cat healthy and happy. Dogs, on the other hand, can survive and thrive on a vegetarian diet; they do well with only 4 percent protein.

female ocelot would be doing well to produce seven kittens in the same-length lifespan. If food is plentiful, female bobcats can become pregnant when they are nine to twelve months old, but when prey densities are low,

few young females even breed. Pregnancy lasts about sixty-three days, and the mother gives birth to two to three kittens in a wellhidden den, typically a rock pile, cave, or hollow log. There have been reports of females

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Distribution of the bobcat.

using abandoned buildings and even a beaver lodge as a den site. When the kittens are about two months old, the mother starts to bring live and dead animals back to the den for them to eat, but she continues to nurse them for another month. At three months of age, the kittens begin following their mother, and they depend on her to hunt for them until they are about seven months old. Bobcats are hunted and trapped across most of their range in North America. After trade in tropical cat skins was halted in the early 1970s, the skin trade turned to bobcats, and they became the number-one spotted cat in the international skin trade. By 1977 concerns over decreasing bobcat numbers led to them being legally protected—skins were no longer allowed to be traded internationally.

By the year 2000, populations rebounded and recently there have been several petitions to remove the protective legislation. However the bobcat remains on Appendix II of CITES and is the only spotted cat in the world that can be legally trapped and hunted. Today the bobcat is recolonizing many parts of its historic range. A combination of good conservation laws and the cat’s own prolific breeding ability has made it one of the most successful small cats. status: IUCN Red List—Least Concern weight: 13–44 pounds (6–20 kg) head-body length: 20–47 inches (50–120 cm) tail length: 3–10 inches (9–25 cm) litter size: 1–8 kittens

Puma Lineage

Cheetah ACI N ON YX J UB ATUS

The next time you’re driving along the freeway at 70 miles an hour (113 km/hr), look out the window and imagine a cheetah racing alongside you, keeping pace with the car— that’s how fast these graceful cats can sprint. Cheetahs are the greyhounds of the cat family—built to run, they have longer legs than other cats, a long, supple spine, and a long tail that they use as a balancing rod. Though they are about the same size as a leopard, cheetahs lack the heavily muscled, wrestler-like body of the larger cats; rather they have the slim, lanky, deep-chested build of a sprinter. Most cats can retract their claws when they are not using them, but a cheetah’s claws are shorter and straighter than the claws of other cats; and there are no claw sheaths for the claws to pull back into, so even when they are retracted, the claws are easy to see. This Cheetahs use old termite mounds and other raised areas to scan for prey as well as keep an eye out for other large, dangerous carnivores. Even when resting, cheetahs are always alert to danger, and sentry duty is shared among family members.

accounts for the common misconception that cheetahs have non-retractile claws like a dog. Cheetahs use these strong blunt claws like cleats on a running shoe, to help them push off and build speed. The large center pad on the cheetah’s foot is another running aid; the pad is covered with ridges that act like treads on a car tire, providing traction and grip during fast turns. A running cheetah covers about 23 feet (7 m) with each stride, the same distance as a galloping horse. As the cat accelerates, it increases the number of strides per second, until at top speed it is traveling about 82 feet (25 m) per second, or nearly three times faster than the best human sprinter. There is some dispute over the cheetah’s top speed, but the most reliable estimate over a short distance is 68 miles per hour. These cats do not chase prey for long distances. Instead they are designed for a short, explosive sprint. Aided by enlarged lungs, heart, and nasal passages, cheetahs accelerate rapidly, reaching a speed of 46 miles per hour (75 km/hr) in just two seconds. During 153

c h e e t a h   1 55

these sprints their breathing rate increases from 60 to 150 breaths per minute, and heat production increases fiftyfold. Cheetahs store about 90 percent of the body heat they produce while sprinting, and it is likely that the length of a sprint is determined by the amount of heat the cat’s body can store before its body temperature reaches a dangerous level. For a cat that hunts mainly during the day, this heat buildup during a chase can affect the success or failure of a hunt. In an experimental trial, cheetahs refused to run when their body temperatures reached 104.4°F (40.5ºC). The cheetah is an animal of open country, and throughout most of its range in Africa, this cat is associated with grassy plains, open woodlands, and semi-desert. A few cheetahs still survive among the sandy plains and rocky hills of the Sahara, where they spend the day among the rocks and under bushes avoiding the intense heat of the sun. In the Sahara, cheetahs do most of their hunting at night and in the relatively cool hours just after daybreak. In the Serengeti, cheetahs hunt later in the morning and earlier in the afternoon to avoid competition with lions and hyenas. If they have the opportunity, lions, leopards, and hyenas will kill adult cheetahs and cubs and steal cheetah kills, so the timid, retiring cheetah goes out of its way to avoid these larger, more aggressive carnivores. Wherever cheetahs are found, they usually hunt gazelle or gazelle-like antelope that The slim, lanky cheetah has the deep-chested body of a sprinter. Cheetahs also have comparatively small heads, and their canine teeth are reduced in size—meaning they lack the bite power of the other big cats. Cheetahs kill by strangling their prey with a throat bite.

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Asiatic cheetah: Only remaining Asian cheetahs are found in Iran

Once widespread in Asia, the last representatives of the Asiatic cheetah now exist only in the cold, dry, hilly mountainous regions of central and eastern Iran, where there are about 70 to 100 individuals. Recent studies show that Asiatic cheetahs are quite distinct genetically from their African cousins and suggest that their populations have been separated for 30,000 to 70,000 years, making the Asiatic cheetah much older than previously thought.

weigh less than 88 pounds (40 kg). If more than one cheetah is involved in the hunt, larger prey such as wildebeest may be taken. Cheetahs make full use of cover to stalk as close as possible to prey. Head lowered to shoulder level, body in a semi-crouch, they alternately creep or run forward, then freeze, maneuvering until they are close enough to launch a short, fast chase. They focus on one animal before the chase begins and rarely switch targets during the chase. At the end of the chase, while running slightly behind or beside the gazelle, the chee-

tah reaches out and, using its large hooked dewclaw, snags the prey on the leg or hindquarters, which causes the gazelle to stumble and fall. Once the animal is down, the cheetah grabs it by the throat and lies beside it, maintaining a stranglehold for five minutes or more. Cheetahs often drag or carry their meal to the nearest shade and rest beside the carcass for some time, panting heavily. If more than one cheetah is present, those not directly involved in the kill begin feeding immediately, while the cat that made the kill cools down. Cheetahs eat fast, but they

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King cheetah: A separate species or abnormal markings?

Though it was originally described as a separate species, the king cheetah is in fact just an ordinary cheetah with abnormal markings. The king coat pattern is controlled by a single recessive gene; if both parents have the gene, about a quarter of their offspring will have the king coat. The king cheetah was unknown to science until 1926, when a farmer in southern Rhodesia purchased a cheetah skin patterned with stripes and blotches instead of the normal spots. Geneticists now believe that the king cheetah coat pattern results from a mutation in the “tabby” gene. In the wild, king cheetahs are found exclusively in Zimbabwe, Botswana, and the Transvaal, but they were virtually unknown in captivity until 1980. Today there are several dozen king cheetahs in zoos and captive-breeding centers around the world.

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Dogs working for cat conservation: Guard dogs protect livestock from predators

Recently in Africa, livestock-guarding dogs have become an integral part of Namibia’s highly effective cheetah conservation program. The idea of livestock-guarding dogs is based on encouraging a strong social bond between the dog and livestock. By rearing the puppies with the sheep and goats, with very little human contact, the dog comes to think of the sheep as pack members, and it will defend the sheep against all predators. In 1994 the Cheetah Conservation Fund in Namibia began breeding Anatolian shepherds—a special breed of livestock-guarding dogs that has been used in Turkey for 6,000 years to protect sheep from wolves. Puppies are placed with the livestock when they are about six to eight weeks old. The dogs bond with the sheep and goats, and move with them as they graze. When a leopard, cheetah, or other predator approaches the herd, the dogs bark loudly and stand between the cat and the flock. Farmers have reported seeing their dogs fighting with predators, and the dogs have killed jackals, baboons, and leopards that threatened “their” herd. Namibian farmers are extremely pleased with how well the dogs work to protect their flocks—almost three-quarters of farmers report a large decline in the levels of stock loss since getting the dogs.

Aided by an enlarged heart, large nasal passages, and a flexible spine, the cheetah is the fastest land mammal on earth.

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Cheetahs have many adaptations for speed and maneuverability. They have large thigh muscles, long limbs, a long tail, and a long, flexible back. Their hard footpads are covered with ridge-like threads like those on a car tire, and their stiff, unsheathed claws provide traction and grip during sharp turns.

are careful, tidy eaters, and groups feed together quite peacefully, with only occasional growling and snapping. Ranchers say that a skeleton with the bones still connected, not torn apart, is considered to be the characteristic sign of a cheetah kill. These slender cats

are quite capable of eating large quantities of food, and individuals have been known to eat more than 20 pounds (9 kg) of meat in less than two hours. Each cheetah has a unique pattern of spots on its face and chest; scientists have used these

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Cheetahs have more cubs than most other cats—litters of eight cubs are known, and as many as six cubs have been seen traveling with their mother.

spot patterns to identify and recognize individuals. The arrangement of black bands at the ends of a cheetah’s tail is also unique to individuals, although the tails of littermates look very similar. A conspicuous black tear line runs from the inner corner of the eye

to the mouth, which may function like the black-line face paint that that football players wear, to help the cheetah cope with glare from the sun. Cheetahs make a variety of unusual calls. They have a yelp—a brief high-pitched

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Distribution of the cheetah.

“yow”—which is used as a contact call, most commonly by mothers separated from their cubs or young that have lost their mother or siblings. The yelp can be heard for a mile (2 km). Another call is the “churr,” which is used by mothers to encourage their cubs or by males to find their partners. After a meal or when resting, cheetahs also purr, which sounds like a domestic cat only much louder. Cheetah have larger litters than other cats— often five or six cubs in a litter—and the young become independent at a relatively

early age and breed early. In places where they are the only large predator, cheetah populations are expanding. However, where cheetahs live alongside lions and hyenas, these predators kill an astoundingly high percentage of cheetah cubs. In the Serengeti, only 5 percent of cheetah cubs survive to adulthood. Cheetahs present an extraordinary challenge to conservationists. As long as they are not hunted, cheetahs may actually be better off outside national parks than inside these

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protected areas where there are high densities of lions and hyenas. The Cheetah Conservation Fund is dedicated to saving the cheetah in the wild. For more information on the project, field updates, or to make a donation to their work, go to http://www.cheetah.org/.

status: IUCN Red List—Vulnerable weight: 46–143 pounds (21–65 kg) head-body length: 44–55 inches (113–140 cm) tail length: 24–33 inches (60–84 cm) litter size: 1–8 cubs , 3–4 common

Puma

PUMA CON COLOR

Pumas are usually classified with the “big cats,” but there has always been something a little different about them. Biologists sometimes call them “small cats in a big cat body” because though they weigh almost as much as leopards, pumas just don’t have the robust muscular head and heavily muscled forelimbs of the other big cats; rather they look halfway like a cheetah. In addition, pumas don’t roar like the big cats but have a shrill, whistle-like call. And pumas purr, something that none of the big cats do. A few years ago these oddities came together as molecular geneticists confirmed that the puma is genetically a very different cat; its closest living relative is the cheetah, with the two species diverging more than 3.5 million years ago. With this established connection, it is easy to see that pumas do bear a closer resemblance to cheetahs than to the Recent genetic studies show that the puma is not one of the big cats but is actually more closely related to the cheetah and jaguarundi.

other big cats. Both pumas and cheetahs have small, rounded heads; slim, lanky bodies; and moderately long limbs. Like cheetahs, pumas are timid, retiring cats, more eager to flee than to fight, and both species rarely confront humans. Pumas also have unusually long hind legs and a relatively long spine, which provides increased flexion while running. The cheetah shares this characteristic, but the puma relies more on ambushing prey than running it down. Twelve thousand years ago in North America, a catastrophic extinction event abruptly wiped out 80 percent of the continent’s large animals. Ground sloths, camels, capybaras, mammoths, and many other herbivores disappeared, along with the lions, cheetahs, and saber-toothed cats that preyed upon them. Pumas were probably eliminated from North America at the same time but survived in South America, and a few of these survivors quickly moved north to repopulate North America after the great extinction. DNA evidence shows that all 165

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Continent-crossing puma: Young male makes record journey

In June 2011 a puma or mountain lion was hit and killed on a highway in Milford, Connecticut. The 140-pound young male made the headlines as the first wild mountain lion recorded in the state in a hundred years, but then a few weeks later, officials announced even more incredible news. This cat came with a barely believable backstory. Incredibly, scientists had matched this cat’s DNA to hair and feces left by a mountain lion that had wandered through Minnesota and Wisconsin in 2009 and 2010. Further tests showed that the Connecticut cat had been born in a population of mountain lions somewhere in South Dakota. During the previous two to three years, he had walked across more than half the continental United States in what was one of the longest-ever recorded journeys for a land mammal. Considering the highways and hazards he must have encountered in this epic trip, his journey makes the Pleistocene migrations look easy.

In Canada, moose, elk, and deer make up

p u m a   1 67 more than half of the puma’s diet, but in the

tropics these cats survive on armadillos and other small prey.

Homeward bound: Tendency for translocated cats to find their way home Having a domestic cat in your house can teach you a great deal about the behavior of wild cats. Take, for instance, the stories of homing cats. Like most scientists, we were dubious—until we relocated. When we moved to a new house, we kept the cats indoors for six weeks. The first day we let them out into the yard, they vanished. Two weeks later they were found: they had traveled the 6 miles back to our old house. Relocation has always seemed like an easy option for problem wild cats. “Can’t you just move them?” is always the first question when a puma or jaguar causes a problem. But this is rarely a solution because wild cats are just as good at homing as domestic cats. In a classic study of translocated pumas in New Mexico, two male pumas were moved nearly 310 miles (500 km) then radio-tracked as they returned to their original ranges. It took one male 469 days to travel the 288 miles (465 km) back to his original range. The other male traveled 305 miles (490 km) in 166 days, regained his territory, and in the following years fathered three litters of kittens.

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In the late 1800s and early 1900s, pumas were classified as vermin and systematically eliminated from two-thirds of their former range, including most of the eastern United States. In the 1970s, wildlife agencies began to manage pumas as a game animal; puma numbers have rebounded across the United States, and the cats have begun to recolonize many parts of their historic range.

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Black pumas: Many people claim to have seen a black puma, but do they exist? In North America there are literally thousands of reported sightings of black pumas, also known as black panthers, but none of these sightings have ever been authenticated. Despite the fact that people regularly report seeing black pumas and sometimes even claim to have shot them, there are no skins in collections or specimens in any museum. Even though panthers are shot by hunters and hit on the road by cars, no one has ever produced or photographed a black specimen. Despite this, sightings of black pumas continue to crop up year after year in newspaper reports. In Florida, where only 160 or so of these cats exist, black panthers are the most commonly reported cats. Biologists insist that there are no black pumas in Florida and maintain that they should know, because most of Florida’s pumas have been radio-collared, photographed, and followed since birth. Throughout the puma’s range, officials of state wildlife agencies remain skeptical about the existence of black panthers, ascribing the sightings to large house cats, excited observers, or poor lighting conditions. However, most of the public continues to believe in black panthers, arguing that so many observers cannot be wrong.

North American pumas are descended from just a small number of animals. The adaptable puma managed to thrive on the smaller prey that remained, and this adaptability continues to be the key to their success. Pumas are now found from the tip of South America and north to southeastern Alaska. They live in deserts, mountains, and tropical forests, where they survive on prey that ranges in size from mice to moose. In the snows of Alberta and British Columbia, they regularly bring down moose and elk, prey three to ten times their own body weight, and in Montana in summer, they often spend the day chasing after 1-pound ground squirrels. But across most of North America, deer make up more than half the puma’s diet. In South America these adaptable cats survive on hares, armadillos, peccaries, capybaras, and guanacos. Pumas can also be quite individualistic or idiosyncratic when it comes to the prey they kill. In Nevada some have learned how

to kill wild horses, and in Mexico a female puma killed 72 horses, mules, and colts over a nine-month period. Pumas will, like other cats, indulge in surplus killing sprees, particularly with domestic sheep. In one incident, 192 sheep were killed in a single night. These incidents usually occur when a cat ends up trapped inside a pen with a lot of frightened animals, and many sheep are killed in the melee. Pumas will kill almost any animal that puts itself in a vulnerable position, but when hunting large prey such as deer, elk, or horses, the attack can be a hazardous moment for the cat. Pumas have been found dead—impaled on branches, with broken necks or smashed skulls. All of these injuries happened when they were trying to make a kill. One study suggested that as many as a quarter of natural deaths are caused by injuries that happen while the cat is trying to capture prey. One way or another, an adult puma needs to kill the equivalent of a deer about every

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Florida panther: Collisions with vehicles threaten remaining cats

Splashing through belly-deep water in the Everglades and living their secret lives in Florida’s inaccessible swamps, a small population of puma eke out a living on deer, hogs, and raccoons. Known as Florida panthers, this southern subspecies of puma was thought to be extinct until a hunter accidentally treed one in 1973. Follow-up investigations discovered 35 or so of the cats hidden in the swamps and wetlands at the southernmost tip of the Florida peninsula. Today approximately 100 to 160 Florida panthers hunt in the swamps, hardwood hammocks, and pinelands of south Florida. Male panthers need an area of about 200 square miles (518 sq km); females require about 75 square miles (194 sq km). While moving about their home ranges, Florida panthers also suffer fatal consequences of living in such a heavily populated state; many of the cats are killed while attempting to cross the state’s busy highways. An astounding 40 percent of all Florida panther deaths are animals killed in collisions with vehicles. Partly in response to this massive road kill, the state of Florida pioneered the development and use of wildlife underpasses. When Alligator Alley, the major east–west highway across southern Florida, was upgraded to a four-lane highway, a 40-mile (64 km) portion of the road was completely fenced and equipped with thirty-six underpasses. Wildlife attempting to cross the road is funneled into the underpasses by a 9-foot-high chain-link fence installed along both sides of the highway. The underpasses have been extremely effective at reducing panther road kill. Though the toll on cats continues on other roads, no panthers have been killed on the sections of road that have been equipped with underpasses.

eight or nine days; and with more mouths to feed, females with large cubs have to hunt more and kill twice as often. In zoos pumas are fed 4 to 11 pounds (2 to 5 kg) of fresh meat a day, depending on their size. Pumas that intend to return to a kill sometimes cover the remains with leaves, stones, grass, or snow. This habit is probably to protect the remains from scavengers, but the behavior has provided fertile ground for some wonderful tall tales and apocryphal hunting stories. Many woodsmen have written of spending the night out in the forest wrapped in a blanket, only to awaken the next morning covered with a pile of leaves. Most of the stories conclude with the hunter awakening just as the female puma returns with her cubs, and almost all of the hunters manage to shoot the puma just before it attacks. Unlike the big cats, pumas don’t roar but

maintain contact with one another with chirps and shrill, whistle-like calls. Females also have a long-range call that has been described as sounding “like the scream of a vampire woman” and “unearthly.” This scream, or caterwaul as it is sometimes referred to, is like a louder version of the sound made by mating domestic cats. The call carries for 1,300 to 1,650 feet (400 to 500 m) and is probably intended to let the male know that the female is ready to mate. Except when they are courting or mating, pumas live solitary lives, avoiding each other whenever possible. This is not surprising, because these cats fight frequently and with fatal consequences. Male pumas kill other males, females, and cubs; in fact, fights with other pumas are the major cause of death among pumas of all ages, and infanticide and cannibalism combined are the main cause of cub

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Pumas and people: Attacks on people increasing; how to avoid becoming a victim Recent high-profile stories about pumas standing on decks, looking through sliding-glass doors, and killing deer in people’s backyards have ramped up public apprehension about living near pumas. Between 1890—the date when records were first available—and 2011, there have been roughly 158 verified cougar or puma attacks on humans in the United States and Canada, and 22 deaths. Most attacks on humans occur in California, British Columbia, and Colorado, and there have been 12 deaths in the past twenty years compared with 10 in the previous hundred years. Most pumas avoid direct encounters with people, but as more people move into puma habitat, the likelihood of cat-human encounters increases. The push for space is coming from both sides. On one hand, increasing puma numbers are repopulating western wilderness areas; and, on the other hand, people are building more houses in the wilderness and choosing to live in habitats that have plenty of hunting cover, as well as deer and elk—perfect puma country. People are also expanding their recreational use of parks and private land; there are more walkers, joggers, and people hiking the backcountry than ever before and more opportunities for encounters between people and cats. The potential for being killed or injured by a puma is low, but you need to follow a few good rules for hiking in puma country. The most important of these is never hike alone, and if you are walking with children, never let them roam ahead or lag behind. Children are at a much greater risk of being attacked, as are solitary hikers. If you encounter a puma • Don’t run—running stimulates the instinct to chase. • Avoid crouching or bending over—pick up children quickly.

• Do everything to appear larger—raise your arms, open your jacket, throw sticks, speak in a loud voice. • Fight back if attacked—use rocks, sticks, backpack, anything to hand.

death. Some males are more aggressive than others, and since they can be as much as 70 percent larger than females, serious injuries and fatalities are common. Why pumas fight so much is a bit of a mystery. Most likely, males are fighting for territory and mating rights, but why they would kill breeding females and potential mates remains a puzzle. Given the opportunity, young female pumas prefer to settle next to their mother, and in areas where they are not hunted, pumas develop clusters of related females, in which the ranges of mothers, daughters, and sisters overlap or abut one another. Within these clumps of related females—or matrilines, as they are called—cub survival is higher.

Young male pumas are frequent targets of adult male aggression, as they are seen as potential competitors for mating opportunities. To avoid these confrontations, young males typically leave the areas where they were born and travel extensively, searching for areas without resident males. They can move long distances surprisingly fast—one male traveled nearly 30 miles (48 km) in two days. Finding an area that they can claim as their own is a long process; it sometimes takes months and involves traveling over hundreds of miles. Several males have been found more than 100 miles (160 km) from their birth areas, and these long-distance movements are obviously an important component of the ex-

Distribution of the puma.

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Signs warning hikers and joggers of the presence of mountain lions are common in many California parks and along hiking trails.

pansion of pumas back into areas from which they were eliminated many years ago. The fact that pumas are quietly exploring landscapes now dominated by highways and humans is a reminder of their historic reinvasion of North America 10,000 years ago and testament to their remarkable adaptability.

status: IUCN Red List—Least Concern weight: 66–167 pounds (30–80 kg) head-body length: 39–59 inches (100–150 cm) tail length: 24–35 inches (60–90 cm) litter size: 1–5 cubs

Jaguarundi PUMA YAG OUA ROUN DI

The jaguarundi is an enigma. It does not look like any of the other small South America cats and shows some behavioral similarities with the puma. For a long time, it was placed in its own genus, but recent genetics studies have cleared up the confusion. It turns out that the jaguarundi is more closely related to the cheetah and puma than to the other South American cats. Molecular DNA place the jaguarundi, puma, and cheetah together in the Puma lineage, and fossils show that all three species originated in North America. With their small heads, long, slim shape, and long tails, the puma, cheetah, and jaguarundi resemble one another physically, and all three communicate with un-catlike whistles and chirps. The confusion over the jaguarundi’s position in the cat family has also been fueled by its appearance—it hardly looks like

The jaguarundi is very different in appearance and behavior from the other small South American cats. Recent DNA studies reveal that this unusual-looking cat is more closely related to the puma and cheetah.

a cat at all. Its elongated, low-slung body is more reminiscent of a marten, although some people have compared it to a weasel or otter. Like pumas, jaguarundis have a plain-colored coat. There are two main color phases—an iron-gray morph and a red-brown morph—both color phases can be born in the same litter. Kittens are born spotted, but the markings disappear as they grow. Jaguarundis hunt in more open areas than other cats, and they are most often seen in places where dense cover is mixed with openings and edges. They hunt during the day, on the ground, but they are also agile climbers, adept at moving along branches. They may spring 6 feet (2 m) off the ground to swat at a bird or to investigate a noise in the bushes while standing in a tripod positions, balanced on their haunches and tail like a kangaroo. They hunt mainly rodents, birds, and reptiles, but sometimes manage to catch larger animals such as rabbits, opossums, and armadillos. 175

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Alone among the small cats, the jaguarundi has a plain unmarked coat. There are two main color phases: the iron-gray morph and the red-brown morph. These phases were once thought to represent two distinct species, but both colors can be born in the same litter.

Although the jaguarundi is the most commonly seen cat in Central and South America, its biology and behavior are still largely a mystery. There are several reports of the cat in the United States, including Florida, but the last confirmed jaguarundi was a road kill in Texas in 1986.

status: IUCN Red List—Least Concern weight: 7–15 pounds (3–7 kg) head-body length: 21–30 inches (53–76 cm) tail length: 12–20 inches (31–52 cm) litter size: 1–4 kittens, usually 2

Distribution of the jaguarundi.

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Purring: Cats purr in a variety of different contexts

Except for lions, leopards, jaguar, tigers, snow leopards, and clouded leopards, all cats purr. Though most people would agree that cats purr when they are happy and content, there is no such clear agreement among scientists as to the function of purring. Cats purr when they are relaxed and happy, but surprisingly they also purr when they are injured or sick. Veterinarians report that cats sometimes purr when they are in pain or in highly stressful situations. Purring is a continuous murmuring sound produced both on the inhaled and exhaled breath. It is also a closerange sound—mother cats purr while nursing kittens, and cats purr when snuggling up to people—so physical vibration of a purr and body contact between the purrer and the receiver is undoubtedly important.

Ligers and tigons: Products of mating between lions and tigers

A liger is a cross between a male lion and a female tiger. They share the physical characteristics of both parents, and their fur usually has spots and stripes on a lion-colored background. Ligers usually grow larger than lions or tigers and may weigh 800 to 1,000 pounds (360 to 450 kg). Male ligers are often sterile, but females can be fertile. There are several ligers in captivity, said to be the result of “accidental” pairings. Most reputable zoos do not encourage these cross-species matings, but as ligers can be great moneymakers for their exhibitors, some less scrupulous wildlife parks keep them on exhibit and charge for viewings and photography sessions. Tigons are a cross between a male tiger and a female lion and can be striped or spotted. Unlike ligers, tigons do not grow larger than their parents; they are less common than ligers and not often seen in captivity.

Leopard Cat Lineage

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Pallas’s Cat OTOCOLOB US M A N UL

The Pallas’s cat is a squat, short-legged little feline with a long, shaggy coat and a thick bushy tail. A broad head with a low forehead and a flattened face makes it look like a pug or Pekingese dog. Thick fur and a stocky shape help reduce heat loss while the cat hunts in the high-altitude grasslands and rocky outcrops of the Mongolian steppe. Winters in Tibet and Mongolia are severe; temperatures can be very cold: -58°F (-50°C)—colder than the temperature of a household freezer. There is not much in the way of cover to hide behind on the bleak, treeless Mongolian steppe, and Pallas’s cats are not fast runners, so they have become masters of hiding in plain sight. Their small round ears are set so far down on the sides of the head that the tips of the ears are level with the outer corner of the cat’s eyes. This helps them keep a low profile. When threatened, they crouch and lie flat on the ground, head down and motionless—pale, The squat, short-legged Pallas’s cat resembles a pug or Pekingese dog.

shaggy fur helps them blend into the short grass and rocks, and they become almost invisible. Pallas’s cats inhabit the high-altitude steppe grasslands and semi-desert areas of central Asia. In the Tien Shan, Govi-Altai, and the Pamirs, the cat is found at elevations of 9,850 to 13,120 feet (3,000 to 4,000 m), but most records come from lower elevations. In Ladakh the cat is found in barren, stony valleys and hilly terrain above the tree line, from 11,800 to 15,750 feet (3,600 to 4,800 m). Little rainfall, low humidity, and a wide range of temperatures characterize typical habitat for the Pallas’s cat. Summer high temperatures may reach 100°F (38°C); in central Asia the winter air temperature is commonly -58°F (-50°C), and snowfall is light and uneven. Pallas’s cats cannot navigate deep, loose snow, and they are rarely found in areas where the snow is more than 4 inches (10 cm) deep. These cats hunt by stalking or slowly creeping up on prey; they are not good or particularly fast runners, so they need to get close before pouncing. They hunt mostly by sight, 181

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Pallas’s cats live in central Asia, in semidesert areas with rocky outcrops. They spend the day in caves, rock crevices, and marmot burrows, emerging in late afternoon to begin hunting.

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Pallas’s cats are poor runners and rely on camouflage to creep close to prey. The cat’s low-set ears are adaptations for hunting in open terrain.

using the sparse vegetation as stalking cover. Their low-set ears and flat forehead are adaptations for hunting in open terrain, allowing the cat to peer over rocks or low bushes without exposing too much of its head. Pallas’s cats feed on a broad range of small rodents, insects, birds, and carrion. In areas where pikas are common, they become the cat’s favorite prey, making up more than half their diet. Pallas’s cats prefer to hunt these 4.5-ounce (125 g) rodents because they are larger than gerbils and jerboas. As might be expected of a cat that lives in

such a harsh climate, the Pallas’s cat is a seasonal breeder: most litters are born in April and May. For many cat species, two or three kittens in a litter are the rule, but Pallas’s cats have large litters; two to four kittens are common, and as many as five or six is not unusual. There is even a record of eight kittens in a litter. The ability to have such large litters is not uncommon among felids living in highly seasonal environments where prey abundance fluctuates widely. Canada lynx and European wildcats also give birth to as many as many as eight kittens in a litter when food is very abundant.

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Distribution of the Pallas’s cat.

Related to rabbits, the rat-size pika is a plant-eater. Pikas can produce two or more litters of as many as ten young over the course of a single year and are considered a keystone species over much of their range in the highaltitude grasslands of Asia.

Pallas’s cats do not dig their own dens, but instead shelter in caves, rock crevices, or abandoned marmot burrows. If they are caught out in the open, these cats are very vulnerable to predators such as eagles, wolves, foxes, and herders’ dogs. Having to constantly fend off larger predators may explain the cat’s reputation for meanness in captivity. Pallas’s cats

in zoos are notorious for their fierce attitude. Bill Swanson, of the Cincinnati Zoo, remembers a litter of newborn Pallas’s cats that zookeepers thought were having trouble breathing. “When they listened closely, they realized that the noise they were hearing was the kittens growling and hissing at each other—before they had even opened their eyes!” Pallas’s cats live at extremely low population densities and are very vulnerable to overexploitation. They are legally hunted for fur and traditional medicines—among other things, herders use the fat to relieve frostbite. State-sponsored pika-control programs and overgrazing by livestock also threaten their survival. status: IUCN Red List—Near Threatened weight: 5–10 pounds (2.5–4.5 kg) head-body length: 18–26 inches (46–65 cm) tail length: 8–12 inches (20–31 cm) litter size: 1–6 kittens, 3–4 common

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Fishing Cat PRI ON A I LURUS VI VE RR IN U S

Several cat species will catch and eat fish when the opportunity arises, but the fishing cat is the only species to be named for its habits and diet. Fishing cats and leopard cats are closely related and share several physical characteristics, including similarly spotted coats; long, narrow skulls; and small, rounded, black-backed ears with prominent white patches. But here the resemblance ends. The powerful, stocky fishing cat and the slim, agile leopard cat look as different as a weight lifter and a ballet dancer. Rather than the lithe, light-footed grace that one normally associates with a small cat, the fishing cat presents an aura of power and strength. Its deep-chested body and comparatively short legs give it the look of a much bigger cat. The fishing cat is about twice the size of a large house cat. The toes on the front feet are partially webbed, and the tips of its claws While fishing cats are found in wetland habitats, their distribution across Southeast Asia appears to be very localized and is still not well known.

protrude from their sheaths, even when fully retracted. The short tail is about one-third of head and body length and unusually thick and muscular near the body. The fishing cat lives in marshes, swamps, oxbow lakes, reed beds, mangroves, and tidal creeks. It is very much at home in the water and can swim long distances, even underwater. With its powerful build and strong swimming ability, it is not surprising that the fishing cat takes a wide range of prey, most of it in and around water. Fishing cats can be seen crouching on sandbars and rocks along rivers, using their paw to scoop out fish, and swimming underwater to catch ducks and coots. One was watched by S. Mukherjee, an Indian graduate student, as the cat hunted along the edge of a small canal. It pounced on a frog, ate it, then moved to another spot, where it sat, focusing intently on the scene. A few moments later, the cat leaped into the water and, putting its head under the surface, grabbed something, then ran onto the bank with its prize. In zoos fishing cat kittens take to water at 187

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The fishing cat’s partially webbed feet, a thick, muscular tail, and a long, narrow skull are all adaptations for swimming and capturing fish.

f i s h i n g c at   189

Fishing cat mother with her three-month-old kitten.

an early age. They begin to play in their water bowl at four weeks, and by three months they are spending a lot of time in the water, play fighting, wrestling, and pouncing on imaginary fish. They hook fish out of their pool with a paw, and in deeper water the cats wade in and submerge their head to seize the fish with their teeth. Fishing cats are not common in captivity, but the few institutions that do keep them have found that when provided with a pond and live fish to catch, they make striking exhibits. Fishing cats seem to be unusually tolerant of each other, so several adults can be kept in the same enclosure. Fishing cats belong to the leopard cat lin-

eage, and recent DNA studies show that the fishing cat, leopard cat, and flat-headed cat— another fish-eating specialist—form a tightknit group of species that appeared in Asia in the late Pleistocene, when sea levels were lower and the islands of Southeast Asia were connected. Interestingly, the flat-headed cat is found only on Borneo, Peninsular Malaysia, and Sumatra, where the fishing cat’s presence is disputed. It may be that the “fishing-cat niche” is not wide enough to support two such specialized cats. The fishing cat is found in scattered pockets of suitable habitat across much of Southeast Asia. There are large gaps in its distribu-

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Fishing cats are unusual in that they will plunge their entire head underwater in search of prey. They have been seen diving headfirst into water, catching fish with their mouth.

tion, and the species seems to have been in serious decline for the last twenty years. The new survey technique of camera trapping has revealed just how rare fishing cats have become. In many parts of Southeast Asia, remotely triggered cameras routinely record the leopard cat, the Asiatic golden cat, and even the elusive marbled cat, but rarely photograph fishing cats. Wetland destruction, elimination of mangroves, and development of wetlands for human settlement and shrimp farming seriously threaten this species. Fishing cats are still trapped for their fur and have a comparatively high market value. The cat’s large size, a sought-after spotted pelt, and a distribution tied to easily convertible wetlands and grass-

lands almost certainly mean that this species will suffer accelerating declines in the future. With this in mind, the cat specialist group recommended that fishing cats be uplisted to Endangered, and in 2008 they became one of the few felid species to be moved from Vulnerable to Endangered on the IUCN Red List. International trade is now controlled, and the species is listed on Appendix 2 of CITES, which limits trade only to within countries with fishing cats. status: IUCN Red List—Endangered weight: 11–35 pounds (5–16 kg) head-body: 26–33 inches (65–85 cm) tail length: 9–12 inches (24–30 cm) litter size: 2–3 kittens

Fishing cats are very much at home in the water and can swim long distances even underwater.

Distribution of the fishing cat.

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Leopard Cat PRI ON A I LURUS B E N G A L E N S IS

The small, lightly built, long-legged leopard cat is Asia’s most common small felid. It is also the only small cat with a geographic range that extends from the tropical forests of the equator, north to the below-zero winter temperatures of the Amur Peninsula. Across this vast north–south distribution, leopard cats show a fourteen-fold increase in body size; they commonly weigh as little as 1.2 pounds (0.5 kg) in Malaysia, while in the Russian Far East they can weigh 15 pounds (6.8 kg). Leopard cats are closely related to fishing In the tropics, rats and mice make up the bulk of the leopard cat’s diet. In northern areas, the larger cats occasionally kill roe deer fawns. Leopard cats are the only small cat with a geographic range that extends from the tropical forests of the equator, north to the cold and snow of the Russian Far East. Cats from the southern and northern ends of this range look like different species. On the equator, leopard cats have dark coats and weigh about 2 pounds (0.9 kg); in the Russian Far East, they are pale silver gray and can weigh as much as 15 pounds (6.8 kg).

cats and flat-headed cats, and, interestingly, these three are the only small-cat species that show a strong affinity for water. All three have well-defined webbing between their toes and tend to hold on to struggling prey, rather than dropping it and reattacking. This behavior is thought to be characteristic of cats that prey on fish because, unlike a rodent, once released a fish is not easy to recapture. Leopard cats are excellent swimmers, completely at ease in the water, which may help to explain why the cat occurs on more islands than any other felid. Indeed, the first leopard cat specimen to be described by scientists was reportedly captured as it swam to a ship anchored in the Bay of Bengal. In Sumatra we watched one young cat playing belly-deep at the edge of a small stream; it eventually swam confidently into the swift-flowing deep water and crossed to the other side. Captive leopard cat kittens spend a lot of time playing in water, and in Borneo two young males kept as pets began playing in a small pond when they were only two months old. According to the 193

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How cats drink: Cats don’t lap like dogs but pull water into mouth with tip of tongue

High-speed photography reveals that cats don’t drink like dogs, which lap up water with their cuplike tongues; instead, cats draw water into the mouth using just the tip of their tongue. With the tip turned downward to form a J-shape, the upper surface of the tip lightly touches the surface of the water. The tongue is then rapidly pulled upward, pulling with it a column of water. A domestic cat laps at four times per second, which appears as a blur to the human eye.

owners, “They tried to catch water weeds or played with each other, falling over and getting thoroughly wet. Frequently they swam right across the pond.” Leopard cats are not tied solely to wetland habitats. Today in Southeast Asia, leopard cats are found in tidal forests, tropical evergreen rain forests, dry coniferous forests, rubber and oil palm plantations, and around villages. In the Russian Far East, leopard cats usually live near rivers, in river valleys, forested ravines, and coastal areas. These cats have small, narrow feet, which make it difficult to travel through deep snow; and in Rus-

sia they are found only in areas that don’t get much snow. A small population of leopard cats on Iriomote Island, about 125 miles (200 km) east of Taiwan, were at one time considered to be a distinct species, but molecular studies show these cats to be an island form of the leopard cat. The DNA evidence suggests that the Iriomote cat population was established from leopard cats from northern China or the Korean Peninsula. Leopard cats are also found on Japan’s Tsushima Islands nearest Korea. In keeping with their wide distribution, leopard cats have a broad catholic diet, with rats and mice making up the bulk of their food; they also kill tree shrews, ground squirrels, bats, and even an occasional animal as large as a hare or a mouse deer. The much larger leopard cats in the Russian Far East also feed on waterbirds, fish, pheasants, and roe deer fawns. Leopard cats are agile climbers and can be quite arboreal. In oil palm plantations, leopard cats are quite at home 9 to 13 feet (3 to 4 m) off the ground as they move lightly and easily among the palm fronds hunting rats. In Sabah, Malaysia, we were able to watch a leopard cat hunting. From its perch near the base of a low-hanging palm frond, the cat spotted a rat moving in some ground creeper. With a long bound, it sprang from the tree and pounced on the rat, which had by then disappeared under the mat of vegetation; after a few seconds of rummaging around under the creeper, the cat raised its head, holding the rat in its mouth, and, without pausing, began to chew. In twenty seconds the event was over, and the cat moved on, pushing its

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Distribution of the leopard cat.

head under the creeping vines that carpeted the ground. In the 1970s the leopard cat was used to create the Bengal cat, a hybrid cross that would appeal to domestic cat lovers. The idea was to create a gentle tame house cat with the beautiful spotted coat of the leopard cat. Bengal cats have become a popular breed, with over 60,000 cats registered with the International Cat Association. But unfortunately, like other hybrids, Bengal hybrids also have significant problems. Once they reach adulthood, these cats tend to bite, scratch, fight, destroy furniture, and urine mark throughout the house. Cat rescue organizations get hundreds of calls and e-mails from people who have succumbed

to the appeal of having a cute, wild-looking kitten, and then discovered that they can no longer handle the adult cat. The Bornean Clouded Leopard Programme aims to advance and enhance the understanding and conservation of all wild cats on Borneo. For more information on the organization, field updates, camera-trap photos, or to make a donation to their work, go to www .wildcru.org. status: IUCN Red List—Least Concern weight: 1.2–15 pounds (0.55–7 kg) head-body length: 15–29 inches (39–75 cm) tail length: 7–12 inches (17–31 cm) litter size: 2–3 kittens

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The flat-headed cat resembles a fishing cat, and both share a preference for rivers, streams, and ponds, where they hunt fish, frogs, and small mammals.

Flat-Headed Cat PRI ON A I LURUS P L A N I CE P S

The odd little flat-headed cat has short, stum­py legs; a short tail; and a long, flat head with tiny, low-set ears. Smaller and shorter than a house cat, it looks like a mini fishing cat and shares the fishing cat’s preference for fish, frogs, and small mammals. The few flatheaded cats that have been seen in the wild have all been along mud banks, near rivers, or at the edge of small streams and ponds. These little cats seem to be designed for finding and catching food in the water. Their teeth are pointed and specialized for seizing and gripping slippery prey; and their large eyes are set farther forward and closer together than those of other cats, giving them better binocular vision. The flat-headed cat also has the uncommon trait of having socalled non-retractile claws. The claws actually retract, but the covering sheaths are reduced, such that about two-thirds of the claws are exposed. Only the flat-headed cat, the fishing cat, and the cheetah share this trait. Flat-headed cats live only in the tropical forests of southern Thailand, Malaysia, Su-

matra, and Borneo. Flat-headed cats are rare and elusive; even in areas with many camera traps, they are rarely photographed. Most of what we know about this cat’s behavior has come from observations of captive animals. In zoos flat-headed cats show a great affinity for water. They search for food on the bottom of pools with their paws spread wide, like a raccoon feeling for prey in the water. Kittens play in water for hours, submerging their entire head to seize pieces of fish. The Bornean Clouded Leopard Programme aims to advance and enhance the understanding and conservation of all wild cats on Borneo. For more information on the organization, field updates, camera-trap photos, or to make a donation to their work, go to www .wildcru.org. status: IUCN Red List—Endangered weight: 3–5 pounds (1.5–2.2 kg) head-body length: 18–20 inches (45–52 cm) tail length: 5–7 inches (13–17 cm) litter size: 1–2 kittens 197

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Small cat swimmers keep it in the family: Three closely related cats are excellent swimmers

Everyone knows cats hate getting their feet wet. Well, it’s true—mostly. The majority of cats avoid water and rarely swim unless pressed. However, there are three closely related small cats from Asia that are excellent swimmers. Three members of the leopard cat lineage—the leopard cat, fishing cat, and flat-headed cat—are all completely at ease in the water. In zoos the kittens of all three species spend a great deal of time playing in the water. Flat-headed cats live on mud banks along rivers and streams and in flooded areas in Borneo, Sumatra, and Malaysia. They are very rarely seen in the wild, but in captivity they play in water for hours, plunging their head underwater to seize frogs and pieces of fish. They also use their forefeet like raccoons to grope around in the mud on the bottom of pools. Leopard cats are also excellent swimmers, completely at ease in the water. In Sumatra we watched one playing belly-deep at the edge of a small river; it eventually swam confidently into the swift-flowing deep water and crossed to the other side. The few zoos that keep fishing cats have found that when provided with a pond and live fish, these cats make striking exhibits. Fishing cats are very much at home in the water and can swim long distances, even underwater. One fishing cat chased by dogs was seen to swim “a considerable distance down a narrow channel without exposing any part of its body. It appeared to have its eyes open the whole time and propelled itself with powerful strokes of its hind feet.” Among the small South American cats, the only one known for its swimming ability is the ocelot. In some areas, ocelots live in seasonally flooded savannahs and during the wet season routinely swim between points of high ground while hunting. Among the larger cats, only the tiger and jaguar spend a lot of time in the water. During the hot dry season, tigers will spend the day submerged up to their neck in a lake or water hole.

About the size of a domestic cat, the flat-headed cat swims and has webbed feet and partially retractile claws.

Distribution of the flat-headed cat.

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Rusty-Spotted Cat PRI ON A I LURUS RUB I G I N O S U S

The tiny rusty-spotted cat is closely related to the leopard cat. It has been called the “hummingbird” of the cat family—a very apt description, because not only is this miniature feline about half the size of a house cat, but it is also extremely quick and agile. Speed and the ability to climb like a squirrel help this cat avoid larger predators. Rusty-spotted cats are rarely seen in zoos, and the Frankfurt and Colombo (Sri Lanka) zoos hold the majority of the world’s captive population. Captive rusty-spotted cats are mainly nocturnal but have bursts of activity during the day. They pace rapidly and their quick, darting movements make them look like speeded-up versions of the other small cats in nearby exhibits. Zookeepers report that these speedy little cats also have tremendous appetites: a cat weighing 5 pounds (2 kg) will eat 4 to 6 ounces (113 to 170 g) of a meat mixThe tiny rusty-spotted cat of Sri Lanka and India is one of the world’s most mysterious felids. Almost nothing is known of their behavior in the wild.

ture every day, topped off with a chick. In the dry forests and scrublands of India and Sri Lanka, rusty-spotted cats feed on birds, rodents, insects, and lizards. Rusty-spotted cats are said to be extremely fierce, but this is hardly surprising as they are a potential meal for just about any other predator, including foxes, jackals, and other larger cats. In all likelihood, these cats hunt mostly on the ground but use their superb climbing ability to quickly escape into the trees when danger threatens. Very little is known about the behavior of rusty-spotted cats in the wild or even their distribution—although based on a recent increase in sightings and camera-trap images of this tiny cat, they may be more widespread than previously thought. status: IUCN Red List—Vulnerable weight: 2–5 pounds (1–2 kg) head-body length: 14–19 inches (35–48 cm) tail length: 8–10 inches (20–25 cm) litter size: 1–3 kittens

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The rusty-spotted cat of Sri Lanka and India is one of the world’s smallest cats.

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Distribution of the rusty-spotted cat.

Seeing in the dark: Cats have a reflective layer in their eye, which helps them to see in the dark If you shine a flashlight on a cat at night, its eyes glow with an almost metallic yellow-green luster. This eye shine has been the ingredient of legend—witch hunters in the Middle Ages thought it sprang from the fires of hell—but in reality it is simply a result of the cat’s eye structure and an adaptation for hunting in low light. When light enters a human eye, the retina absorbs and uses some of it, but the rest is lost. Not so in the cat’s eye. In darkness, its pupils expand until they seem to occupy the whole eye. The wide-open pupils let faint light pass through the iris and strike the tapetum lucidum, a special reflective layer beneath the retina. The tapetum lucidum acts like a mirror, reflecting light back to the retina, in a sort of Ping-Pong effect. Thus, the cat gets a second chance to use light that would have escaped and been lost as it is in the eye of a human. It is this reflecting light that we humans see as the cat’s yellow-green eye shine. For cats, the tapetum lucidum helps improve vision in the dark. Cats’ vision at night is about six times better than that of humans. At night cats can see in what to humans is pitch-dark, thus improving their chances of finding prey.

Domestic Cat Lineage

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Domestic Cat FE LI S SI LVE STRI S CATU S

Cats are an enigma. Proud, dignified, and independent, they are also graceful, soft, and endearing. They charm us with their calming purrs and make us laugh with their obsession for cardboard boxes and paper bags. They are unpredictable and ungovernable, so we are honored when they demand to be petted or decide to sleep in our lap. Most other domesticated animals such as cattle, sheep, goats, and horses naturally lived in groups—they were already used to following one dominant individual as leader, and they soon adapted and transferred their allegiance to humans as herd leaders. Even dogs originally lived in packs and are programmed to obey the pack leader. Not so cats. They are carnivorous, The coat color of the domestic cat has its origins in the striped tabby coat of the African wildcat. With narrow stripes and bars on its legs and tail, and the classic “M” marking on the forehead, the “mackerel” tabby is one of the most common tabby patterns, often seen in feral cat populations. The majority of the world’s cats are still tabbies.

solitary, and fiercely defend their own turf. They truly do “walk by themselves.” Genetic evidence points to the Fertile Crescent—the area that is now Turkey, Lebanon, Iraq, Syria, and Israel—as the site of the early domestication of the cat. Neolithic hunter-gatherers first settled in this area and began to cultivate crops about 12,000 years ago. The houses, fields, and grain stores of these early settlements created new niches for wildlife. House mice moved into grain stores and silos—and wildcats (Felis silvestris) followed the newly abundant food source. Whereas people domesticated most animals by capturing them, then taming and training them to perform useful tasks, wildcats are thought to have domesticated themselves, most likely by hanging around settlements. In early times, cats that tolerated human presence probably had an advantage: people accepted them because they caught rodents, and, charmed by their cute characteristics, a few humans might even have fed them milk and 207

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For domestic cats, a box or a paper bag is a convenient and an attractive den. In zoos even big cats like leopards and tigers have been known to squeeze into cardboard boxes.

scraps. No one knows how long the transformation took, but we have evidence that around 10,000 years ago cats were starting to become important in the lives of Neolithic people. A recent discovery on the Mediterranean island of Cyprus provides an insight into the early relationship between man and cats. Sometime around 9,500 years ago, people from the northern Levant (now Lebanon and

Syria) colonized the island of Cyprus, bringing with them their tools and early domesticated animals. They made the 40-mile crossing by boat with their sheep, goats, pigs, and cattle. They also must have brought some of the earliest relatives of the house cat. In 2004 archaeologists reported that while excavating a late Stone Age village in Cyprus, they had uncovered an elaborate Neolithic human

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Unlike dogs, cats were not bred for specific tasks, and even today most of them remain almost indistinguishable in shape, size, and color from their wild ancestors.

burial site. The grave contained the remains of a 30-year-old person as well as pieces of polished stone, axes, flint tools, two dozen seashells—and the body of a cat. There was no way to know if the cat was a pet, but the careful burial indicates that 9,500 years ago cats had some sort of special significance to the people who lived in that village. Cats are not native to Cyprus, so humans must have carried them to the island along with their other domesticated animals. Beginning about 4,000 years ago, the domestic cat’s history becomes easier to follow as Egyptian artists began to depict the cat in mosaics and paintings. Statues, amulets, and paintings show cats sitting under chairs, riding in boats, and being worshipped. One tomb

Allergies: Kids with pets have fewer allergies Contrary to many parents’ fears, having cats and dogs around the house does not increase a child’s risk of becoming allergic, but actually seems to protect children. Children who grew up with two or more dogs or cats were found to be half as likely to develop common allergies. Researchers think that the effect is the result of early exposure to bacteria carried by dogs and cats. Exposing babies and young children to these bacteria when their immune system is developing during the first year probably helps “exercise” the immune system and might force the immune system to develop in a different way.

painting shows a cat killing a mouse under its owner’s chair; another depicts a cat tied to the leg of a chair with a red ribbon, strain-

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Can you stop a cat killing birds? Do bibs/bells work to reduce cat predation on wildlife?

Many people think that if they give their cat plenty of food, they won’t need to hunt, but the domestic cat’s drive to hunt is independent of hunger—this means that cats will hunt even when they are well fed. Over the years, there have been many attempts to find ways to curb the wildlife-killing tendencies of pet cats. Scientists have attached bells, bibs, and ultrasonic devices to cats in the hopes of interfering with their ability to catch prey. The results are not clear-cut. One study found that cats with bells on their collars brought in fewer prey, while another found that cats with bells actually caught more prey. The most successful device so far is the CatBib, a specially designed piece of synthetic fabric that attaches to the cat’s collar with Velcro and hooks. In the study, cats spent three weeks wearing the bib and three weeks without it. The bib stopped 81 percent of cats from catching birds and 45 percent of the cats from catching small mammals. Though it seems unlikely, cats apparently adapted well to wearing the bib.

ing to get at a bowl of food. But the illustrations also show us that the tamed cats of the pharaohs were not that far removed from the wild. Unlike modern domestic cats, the cats illustrated in the Egyptian images have many physical features common to today’s wild cats.

They are larger, with long legs, a long tail, an elongated body, and a narrow chest. Their shoulder blades stick up above their back, and they walk with a gait like a cheetah—all characteristics of the wildcat rather than today’s domesticated tabby.

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Bastet represents the great cat goddess who was in charge of all growing things; she was a symbol of fertility for both crops and women, and eventually became known as the goddess of love and joy.

About 3,000 years ago, the daughter of Isis and Osiris the sun god was more and more frequently depicted with a cat’s head and came to represent the benevolent aspects of the sun. Bastet, as she was called, eventually became the great cat goddess who was in charge of all growing things; she was a symbol of fertility for both crops and women, and eventually came to be the goddess of joy and love. Some

2,000 years ago, Bastet emerged as a major deity and the focus of the famous cat cult, whose center was the great temple of Bastet in the city of Bubastis, east of the Nile. The shrine contained a giant statue of the goddess Bastet and thousands of cats that were fed and cared for by priests and attendants. As the goddess of joy and love, Bastet was an extremely popular deity, and each year thousands of people celebrated the festival of the cat goddess with a pilgrimage to Bubastis. Paintings from this time showed that cats came in several colors, including ginger, orange-brown, and tabby. At this time, many people owned cats, and the death of a pet would send the entire family into mourning, almost as if a family member had died. The dead animal was embalmed and buried in a special cat cemetery. Food for the afterworld, in the form of small pots of milk and sometimes even mummified mice and shrews, was buried with the cat. The penalty for killing a cat was death, and people would flee if they saw a sick or injured cat on the street for fear of being held responsible. Cats spread from Egypt to the rest of the world soon after Christianity arrived in Egypt. Once in Rome, the cat dispersed through Europe as a camp follower of the constantly traveling Roman armies. By the tenth century, the cat was becoming more common in Europe, even written into legal definitions. In Wales in the tenth century, a hamlet was defined as a place that contained “nine buildings, one herdsman, one plough, one kiln, one churn, one bull, one cock and one cat.” Cats most likely spread through Europe and around the world by way of sailing ships and barges, and there are many nautical terms

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Wildlife killers: Cats are serious predators of birds and small mammals

Almost every cat owner has experienced that “Oh nooo” moment, when his or her cat appears at the door with an offering of a dead mouse or fatally injured bird. Sadly, this moment is happening in a million U.S. households every day. It is a hard truth for cat owners to accept, but outdoor domestic cats really do kill a lot of wildlife. There are over 86 million pet cats in the United States, and at least half of them spend time outdoors. It is almost impossible to calculate exactly how many small animals and birds each cat kills, but there have been several studies in which home owners reported the number of “kills” their cats brought home. In these studies of hundreds of home owners and their cats, the cats averaged two to three kills per month. This number seems small, but it is actually huge when multiplied by millions of cats. Based on these and other studies, scientists estimate that in the United States, domestic cats kill hundreds of millions of birds and more than a billion small animals every year. When our cat brings us a gift of a dead mouse, most of us reassure ourselves that it is only once in a while, and that there are plenty of birds, lizards, and mice. The reality is that our pets are a serious hazard to wildlife, and we each need to find a way to deter our cats from hunting. Whether we keep them indoors, confine them to a fenced area in the yard, or use collars, bells, and bibs, it is up to us as responsible pet owners to address this problem. The price of continuing to delude ourselves is too high.

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Modern cat breeds evolved from two main types: the European, a cold-weather type, which has a large head, stocky body, and thick coat and resembles the European wildcat; and the foreign, a warm-climate type with a slender body, long limbs, and a short coat that resembles the African wildcat.

and weather descriptions that make reference to cats. A light breeze that ripples the surface of the water is known as a cat’s paw, and a cat scratching the leg of a table or chair was thought to foretell a storm. Many other words with nautical associations began with the word cat, such as cat-o’-nine-tails, catboat, catwalk, and cat rig. Carried across oceans or walking from village to village, cats gradually spread across the globe, and by the tenth century they had reached Japan by way of China. Recent genetic studies found that oriental cats have been separated from European cats for at least 700 years, and once they arrived in the East they lived in separate populations, which each be-

gan evolving along their own path. Small isolated groups of cats gradually developed distinctively different coat colors, bobbed tails, kinked tails, and other unique characteristics, which led to today’s breeds such as the Korat, the Siamese, and the Birman, all of which are described by Thai Buddhist monks in a book of cat poems that dates to 1350. Like the early cats of Egypt, many other breeds have a written, pictorial, or oral history as well as a genetic fingerprint. The Japanese bobtail is one of these—its long history is recorded in paintings and supported by historical accounts. We can follow the history of the Japanese bobtail in art because this cat is easy to recognize because of its short

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Modern domestic cats have a much greater variety of coat colors, textures, and patterns than they did two hundred years ago, but the majority of the world’s cats still have some variation of the original tabby coat pattern.

tail. Two Japanese bobtails are featured in a fifteenth-century painting in the Freer Gallery of Art in the Smithsonian Institution in Washington, D.C., and the walls of Tokyo’s Gotokuji Temple, built in 1697, are adorned with paintings of bobtail cats. According to historical accounts, the cat was brought from China to Japan sometime between the sixth and ninth century. These

Cats rule! The most popular pet in the United States In 2011 cats were the most popular pet in the United States. According to the Humane Society of the United States, there are some 86.4 million cats living in U.S. homes, and over half of them live in multi-cat households. The really good news is that 88 percent of these cats are spayed or neutered.

white cats lived in the Imperial Palace of Kyoto and were treated as royalty. Even today there is a long-held Japanese superstition that all white cats bring good luck, and the Japanese “welcoming cat” (Maneki Neko) with one raised paw is believed to bring good luck to the owner. A raised right paw supposedly attracts money, while a raised left paw attracts customers. Cats were not always so admired. During the years that humans and cats have lived around each other, the cat has been regarded as both a god and an agent of the devil. But if you watch a cat for a while, you can begin to understand how these opposing descriptions apply to them. When a cat sits with feet and tail neatly arranged, gazing calmly at the world, you can see why it might have been considered

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What to do about the problem of feral cats? Humane treatment of cats and impacts on native wildlife are major concerns

Millions of feral and free-roaming domestic cats live in cities, parks, and wild spaces in the United States, Europe, and Australia. Many people feed them, but the cats still hunt birds, small mammals, and lizards, killing millions of small animals and birds each day. For cat lovers and conservationists, the two separate but central questions of the issue have become: (1) What do we do about this overabundance of cats? (2) How do we safeguard native wildlife species? To solve the overabundance problem, humane groups are increasingly turning to trap-neuter-return (TNR) programs, which involve the capture, sterilization, and return of cats to the site where they were caught. Volunteer caregivers feed and monitor the feral cat colony on site, which may be around an inner-city Dumpster, on an industrial site, or in a city park. Because neutering cats is always a good thing, TNR is an intuitively satisfying idea to most people, but there is very little evidence to show that releasing neutered cats back into neighborhoods and parks solves the problem. One would think that neutering would result in fewer kittens and thus fewer cats, but oddly this is not always true. At two TNR-operated feral cat colonies in Miami-Dade County, cat numbers initially declined but then increased as illegal dumping of cats and the attraction of stray cats to the provisioned food more than offset any losses. Surprisingly, it turns out that it is not enough to neuter most of the cats in a TNR colony—studies show that of the hundreds of thousands of TNR cat colonies in the United States, only a few have documented a decrease in cats. The second question raised in association with trap-neuter-release programs for feral cats concerns the impact of millions of free-roaming cats on wildlife. TNR programs may help reduce the rate of growth of feral cat populations, but they do little to stop cats from hunting wildlife—as cat owners know, even well-fed sterilized cats continue to hunt. Feral cats hunting wildlife may not be a problem if the cat colony is located at a Dumpster or an inner-city apartment block and the cats are killing rats and pigeons, but it can be a disaster when a cat colony is located next to a park, near a nesting area on a beach, or in a wildlife sanctuary. Ultimately, TNR cat colonies may be appropriate in inner cities and urban areas where cats are unlikely to harm wildlife. However, colonies of free-roaming cats near refuge lands and in state and city parks should not be permitted.

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Tails communicate information about a cat’s mood. A cat holding its tail high and straight is friendly and willing to socialize. A cat that wraps its tail around a person’s leg or another cat is conveying friendship.

sacred—being adored seems to come naturally. It does not take a huge leap of imagination to see the other side—the witch’s cat—a frightened creature in full defense mode, hissing and spitting with back arched and fur on end. The Middle Ages marked the beginning of three centuries of persecution of the cat, and by the fourteenth century they were considered to be the familiars of witches and disciples of Satan. Women, especially the old and ugly, became a special target, and many were accused of being witches. Witches were thought to be able to bring all sorts of misfortune upon people, and as the servant of the witch, cats were extremely suspect. An unknown cat appearing at the doorstep often precipitated a crisis of confidence in the house-

hold as people wondered if the creature was a cat or a witch come to cast a spell on someone. Witches were thought to be able to transform themselves into cats to disguise their activities, and some believed that witches rode to their midnight meetings on the backs of giant cats. The first official trial for witchcraft took place in 1566, and as a result Agnes Waterhouse and her daughter Joan were executed. At the trial it was said that the women had a “whyte spotted catte.” Hundreds of years later, cats were still associated with the devil and bad luck, and were often viewed as witches’ familiars or companions who carried out her evil plans. Even the well-known saying that a cat has nine lives has its origins in witchcraft. According to a book

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entitled Beware the Cat, which was written in 1584, “It was permitted to a witch to take on her catte’s body nine times.” Also connected with witchcraft is the old superstition that a black cat crossing your path brings bad luck; this saying originated with the belief that the black cat was marking a path to Satan. It is astonishing that any domestic cats at all managed to survive this period in Europe. People killed cats whenever they encountered them. Cats were burned, beaten, drowned, and generally abused. No feast day was complete without the torture and death of a cat—it was considered a symbolic way to drive out evil. Cats did not return to public favor until the end of the seventeenth century, when French Cardinal Richelieu took to keeping dozens of them at court. Other people followed suit, and the cat soon regained some of its popularity in Europe. In Rome and Greece in the early days of domestication, cats that had different coat colors probably would not have survived, so all cats would have had some variation of the wild tabby coat pattern. The coat color of the domestic cat is believed to have originated by mutation from the original striped tabby coat of the African wildcat. Today’s domestic cats have a much greater variety of coat colors, textures, and patterns than they did two hundred years ago, but the majority of the world’s cats are still tabbies. When cats began living under human protection, individuals with different coat color mutations became sought after. The selective breeding of cats began comparatively recently, in the late nineteenth century. The move to breed new types of cats began in Brit-

ain and coincided with a surge of public interest in the newly introduced theory of evolution and the widespread fascination with the idea of improving and enhancing different breeds of animals. Some of the more ancient breeds are derived from populations of cats that became geographically isolated—like the tailless Manx or the Japanese bobtail, both of which have existed for several centuries. But apart from these few, almost all cat breeds have been created quite recently. A hundred fifty years ago, cat lovers began to create different cat breeds by selecting random genetic mutations for coat color, fur length, face shape, and ear folds. Today the Cat Fanciers’ Association recognizes forty-two breeds; most of them are defined by body conformation, geographic origin, color, and hair type. Many of today’s breeds are varieties that have been given fancy names: the Balinese is not really from the island of Bali; it is a long-haired version of the Siamese. Likewise, the Somali is a long-haired Abyssinian. The genetic differences between these breeds are very small, equivalent to that between nearby human populations such as the Italians and French. The next great leap in the evolution of the domestic cat began about forty years ago. Using modern reproductive technology, breeders began hybridizing domestic cats with other cat species such as leopard cats, servals, and jungle cats, attempting to create a cat that would have the friendly personality of a domestic tabby with the body of a wild cat. Most people believe the current model will be hard to improve upon.

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Black-Footed Cat FE LI S N I G RI P E S

The tiny black-footed cat is native to southern Africa, primarily South Africa and Namibia, but there are a few records from neighboring Zimbabwe, southern Angola, and Botswana. Almost everything we know of this cat’s biology and behavior comes from Alex Sliwa’s pioneering long-term study on a farm in central South Africa. The smallest cats in Africa, adult blackfooted cats weigh only 2.2 to 5.5 pounds (1 to 3 kg). They have a large, broad head with widely spaced, rounded ears, and a short tail marked with narrow stripes and a black tip. The hair on the soles of its feet and the toe pads are black, hence the name. Black-footed cats live in dry, open grasslands and semi-desert. Rain is almost non-existent, so they must obtain whatever moisture they need from their food. They spend the day resting in abandoned burThe tiny secretive black-footed cat is the rarest of the African felids. Fewer than 10,000 breeding individuals roam the desert grasslands of southern Africa.

rows, hollowed-out termite mounds, or under rock slabs, and emerge to hunt only after sunset. Despite the difficulties of trying to stalk prey where there is nothing much to hide behind, black-footed cats are remarkably successful hunters. Sliwa was able to watch radio-collared cats hunt and found that they attempted to catch something every thirty minutes, with a 60 percent success rate. During a night’s hunting, each cat makes a kill roughly every hour, killing between ten and fourteen small birds, lizards, and rodents. These tiny cats have an enormous appetite, and they eat a fifth of their own body weight every night. What they can’t eat they cache, or stow away, a behavior more common to canids than cats. Black-footed cats will take any animal they can overpower, including animals that are a third to a half of their own body weight, such as hares, mongooses, and bustards. Males kill much larger prey than females, but the smaller, quicker females are more successful at capturing larks, pipits, and other small 219

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The black-footed cat gets its name from the black toe pads and black fur on the soles of its feet.

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Female black-footed cats usually raise two litters of kittens a year. These one-day-old kittens have been safely hidden in a hollow termite mound.

ground-roosting birds, sometimes even grabbing a bird in midair. Sliwa identified three different hunting styles. In the “fast hunt,” the cat moves swiftly through and over the vegetation, flushing prey from cover. A second

style, “slow hunting,” involves a slow stalking movement, with the cat winding snakelike between grass tufts, turning its head from side to side, alert to movements and sounds. In the “sit and wait” hunt, the cat sits motion-

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Scent marking: Cats use scent marks to communicate with other cats Cats communicate with scent from glands on their feet, tail, forehead, cheeks, and chin. When a pet cat rubs its body along the sofa or its chin on the doorway, it is marking its territory, just as wild cats do. Scent contains information on the identity and social status of the cat that left the scent mark and the time the mark was made. Cats scent mark most often in areas where their ranges overlap with neighboring cats, presumably leaving the message: “I was just here.” These scent marks fade over time, and cats have to re-mark the scent posts throughout their range to let other cats know the area is occupied. Territorial male tigers move around their home ranges and renew scent marks roughly every three weeks. Interestingly, when a male dies, it usually takes about three weeks before a new male moves in to claim the now-vacant territory.

less beside a rodent den, waiting for signs of activity, its ears move constantly and its eyes open at the slightest sound. Black-footed cats are solitary, and the only social interactions are between females and their kittens, and between males and females during mating. Though small in size, black-footed cats roam large areas, regularly walking as far as 10 miles (16 km) in a night. Both males and females leave scent marks throughout their ranges as a sort of remote messaging system. Males scent mark more frequently than females, about ten to twelve times per hour. On the night before he mated, one male was seen to spray mark 585 times—an extraordinary feat for an animal that lives in an arid environment with little or no standing water. Black-footed cats also communicate vocally, and their “meow” has been compared to the roar of a tiger, only an octave higher. These cats live at low densities in their semi-desert environment, and the female is only receptive to a male’s advances for five to ten hours. This means that males and females must have a rapid way of finding one another when the female is ready to mate.

Under these living conditions, a loud longrange call is the most effective way for the pair to find each other. Females usually give birth to two litters a year. Pregnancy lasts about sixty-five days, nearly a week longer than the domestic cat. Black-footed cat kittens develop faster than domestic kittens and are able to crawl and hold their heads up on the day of birth. The tiny black-footed cat’s breeding habits are clearly adaptations to reduce their exposure to larger predators. A brief period of receptivity reduces vulnerability during mating, and a long gestation period, small litter size, and rapid development of young combine to shorten the time when the kittens are helpless. The black-footed cat is listed as Vulnerable on the IUCN Red List. Grazing and agriculture are degrading the areas where these cats live. And when ranchers lace carcasses with poison to kill livestock predators, blackfooted cats often die when they scavenge on the carcass. These cats are protected in most parts of their range, and hunting is prohibited in Botswana and South Africa. The Blackfooted Cat Working Group aims to advance

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Distribution of the black-footed cat.

and enhance the understanding and conservation of this Vulnerable felid. For more information on the project, field updates, new camera-trap photos or to make a donation to this work go to their website at black-footed -cat.wild-cat.org/.

status: IUCN Red List—Vulnerable weight: 2.2–5.5 pounds (1–3 kg) head-body length: 13.8–19.3 inches (35–49 cm) tail length: 3–7.5 inches (8–19 cm) litter size: 2 kittens

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The thick-set, heavily furred wildcat of Europe looks very much like a large domestic tabby cat.

Wildcats FE LI S SI LVE STRI S

The wildcats (Felis silvestris) of Eurasia and Africa have been the subject of intense and evolving taxonomic debate. Until 2007 Felis silvestris was most commonly divided into four races, based largely on physical appearance and distribution. The thick-set, heavily furred forest cats of Europe were known as the silvestris group, and the light-bodied steppe cats of Asia were the ornata group. The slim, long-legged lybica group lived in Africa, and catus, the domestic cat, was found throughout the world. The differences between these groups are very small, suggesting a recent common ancestor. Now new genetic information suggests that the Chinese mountain cat, once thought to be an entirely separate species, is another subspecies of silvestris, making five in all. The domestic cat F.s. catus is sometimes considered a sixth subspecies, although the split from F.s. lybica was extremely recent, only 10,000 years ago.

The Current Subspecies: • European wildcat (Felis silvestris silvestris) • Near Eastern wildcat (Felis silvestris lybica) • Eurasian wildcat (Felis silvestris ornata) • Southern African wildcat (Felis silvestris cafra) • Chinese mountain cat (Felis silvestris bieti) status: IUCN Red List—Least Concern weight: 6–20 pounds (2.5–9 kg) head-body length: 18–33 inches (46–85 cm) tail length: 10–15 inches (25–37 cm) litter size: 1–6 kittens

African Wildcats (Felis silvestris cafra and F.s. lybica) The African wildcats are two of five subspecies of wildcat grouped with the European and Asian wildcats, and the more recently identified southern African wildcat and Chinese mountain cat. Genetic studies show that these subspecies began separating some 230,000 years ago, with the most recent 225

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Distribution of the wildcat.

split being the emergence of the domestic cat from the African wildcat subspecies. Domestication of the African wildcat began about 10,000 years ago in an area known as the Fertile Crescent—modern-day Lebanon, Israel, Syria, Turkey, and Iraq—where humans first began to domesticate grains. Grain stores and human refuse attracted rats and mice, and these in turn created a new food source for wildcats. Over the years, the cats that were the most tolerant of people survived, leading to the modern domestic cat. African wildcats are slim, long-legged an-

imals with variably colored fur that is basically used for concealment. Their coat can be tawny gray-brown to reddish depending on where they live. Wildcats that live in the desert tend to be paler, while forest species are darker with bars or streaks that disrupt the outline of their bodies. Two main features distinguish African wildcats from domestic cats and hybrids. African wildcats typically have a rich red-brown color on the backs of their ears; domestic-wild hybrids sometimes retain a little red at the base of the ears but usually have dark gray or

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Asian wildcats have large litters. The mother will soon begin to introduce these month-old kittens to solid food, bringing beetles, rodents, and lizards to the den.

black-backed ears. The second striking characteristic of the African wildcat is its long legs. When sitting upright, the long front legs raise the cat’s body into an almost vertical position. This characteristic pose, which is almost impossible for domestic cats or hybrids, can be seen in the ancient Egyptian bronze mummy cases and tomb paintings. Even when walking, the African wildcat’s long legs and high shoulder blades give it a distinctive action; it moves more like a cheetah than a domestic cat. African wildcats are a fairly common species, with an ability to live in almost every type

of habitat. They do most of their hunting on the ground but can climb well and survive on almost any type of small prey. Rats and mice often make up as much as three-quarters of their diet, but they also eat birds, lizards, and the occasional hare. They usually begin hunting at sunset, are most active between 10:00 pm and midnight, then again around dawn. In winter they often hunt well into the morning hours. When it comes to reproduction, these adaptable cats can clearly adjust their breeding to fit local circumstances, and this reproduc-

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European wildcats have a broad head, with a flat face, and a long, thick coat. They look much more powerful and robust than the slender African wildcat.

tive flexibility allows them to take advantage of superabundant food when it is available. Females come into heat several times a year and, if conditions are right, can sometimes raise two litters in a year. The kittens nurse for a month and become mobile at four weeks. Females begin to take their kittens with them on hunting trips when they are three months old, and young are independent at six months

old, by which time their mother is either pregnant again or has a new litter. Like other wildcats, African wildcats are solitary: both male and female ranges overlap, and both sexes scent mark by spraying urine throughout their ranges. Both have a very loud call—a short, high-pitched “prau” cry that they use to attract members of the opposite sex.

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For at least 9,000 years, African wildcats have been living around people and villages. Even before they were domesticated, people valued them for their rodent-catching abilities.

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Throughout much of Europe, wildcats have been trapped as predators and hunted for their fur. Gamekeepers have historically put a lot of effort into eradicating these cats because they are thought to be major predators of pheasants and grouse.

Wildcats are one of the few species of cats that may have benefited from human activities in Africa, since agricultural development has increased the number of rats and mice. Ironically, considering the fact that

the Africa wildcat is its ancestor, the domestic cat may prove to be the ultimate threat to the wildcat. Throughout Africa, even in remote areas, it is becoming more and more difficult to find purebred African wildcats

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because they interbreed freely with domestic cats.

European Wildcat (Felis silvestris silvestris) The European wildcat looks very much like a large domestic tabby cat. A long, thick coat, broad head, and comparatively flat face make them look larger, but they typically weigh only 6 to 13 pounds (3 to 6 kg), about the same as the tall slim African wildcat. European wildcats are heavily hunted throughout their range, but they still occur in the north of Scotland, Spain, Portugal, France, Germany, Poland, Switzerland, Italy, and southeastern Europe. They live in mixed forests, swamps, rocky areas, and dense thickets, but they are not well suited to moving through deep snow and generally avoid regions that get heavy snowfall. Very little is known of the wildcat’s social habits. They are basically solitary and tend to avoid hunting in the same area even when their ranges overlap. Males and females associate during courtship and mating but otherwise are seldom seen together. Wildcats hunt almost exclusively on the ground at night. Studies in Scotland found that these cats constantly zigzag when hunt-

ing, backtracking and frequently investigating scrub and heather clumps, and they look under the lowest branches of pine trees by pushing their heads through snow buildups. During the heaviest blizzards, they stayed in their dens for as long as twenty-eight hours or moved to lower elevations. In most places this cat feeds mainly on small rodents, but in Scotland they hunt rabbits extensively. They are adaptable predators and can survive on many different foods. They occasionally eat insects, frogs, fruit, roe deer fawns, young chamois, fish, and weasels, and sometimes become significant predators of poultry. Historically, gamekeepers throughout Europe have put a great deal of effort into exterminating wildcats because they are thought to be major predators of pheasants, grouse, and rabbits. Wildcats are also caught in traps set for wolves and foxes. Currently one of the most important threats to the European wildcat is the fact that they hybridize so easily with domestic cats. In Scotland, for example, fewer than 400 purebred wildcats remain in the most remote highland areas, with an additional handful in the captive-breeding program. The key to the Scottish conservation plan is trapping and eventually eradicating the estimated 100,000 feral or farm cats roaming the Western Highlands.

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Sand Cat FE LI S M A RG ARI TA

The stocky, short-legged sand cat is highly specialized for life among the sand dunes and rocky outcrops of the Sahara and the Middle East. Long dense hair growing between its toes form a thick mat that covers the pads of the cat’s feet, insulating them from the hot sand and acting as a sort of cushion on which the animal walks. Because the long hair on the feet also obscures the pads, the tracks of a sand cat are indistinct and difficult to follow. Sand cats also have exceptional hearing. Enlarged ear chambers and big ears provide increased hearing sensitivity. With their specialized hearing, sand cats can hear sounds made nearly a third of a mile farther away than most other cats. Besides all these unusual physical characteristics, this odd little cat also has a distinctive way of moving—belly to the ground, it travels at a fast run punctuated with occaSand cats have exceptional hearing. They travel widely in search of food and keep in touch with one another via a short bark-like contact call.

sional leaps. If pressed, it can put on a sudden burst of speed, sprinting at 18 to 24 miles per hour (30 to 40 km/hr) for short distances. It can also disappear in plain sight: if danger threatens, it crouches flat beside a small rock or tuft of grass, chin on the ground, ears down, becoming almost impossible to see. Throughout its range, the sand cat must cope with extraordinary temperature variation. In summer in Turkmenistan’s Karakum Desert, air temperatures can exceed 122ºF (50°C), while the sand surface temperature can be 176°F (80°C). In winter the air temperature can fall as low as -13°F (-25°C). The sand cat handles these inhospitable conditions by retreating to a burrow during the worst of the heat or cold, and its thick coat offers some insulation. Sand cats can survive without freestanding water, and for most of the year they do not drink, obtaining enough moisture from their prey. Sand cats begin to hunt a little before sunset or just as it becomes dark and continue through the night. Their huge ears and 233

23 4   d o m e s t i c c a t l i n e a g e

Listening in the desert: Enlarged ear chambers allow desert-living cats to hear over long distances

Deserts are challenging places to live. Predators like sand cats and black-footed cats often have to travel 4 to 6 miles (8 to 10 km) in a night’s hunting because the sparse plant cover means prey like mice, gerbils, and snakes are few and far between. Each cat must hunt over a huge territory to find enough food. Because they hunt mainly at night, hearing has become the most important way to sense prey, potential predators, and the calls of other cats. But in terms of sound transmission, deserts have several characteristics that make them different from other habitats. In general, sounds travel farther in deserts. There are no trees to scatter or “attenuate” the sound or produce wind noise. At night when the ground is cool, temperature inversions create conditions that effectively channel sounds near the surface. The sand cat, black-footed cat, and Pallas’s cat live in open, arid habitats, and these cats have enlarged bony ear chambers. Other desert-dwelling species such as fennec foxes, springhares, and jerboas have similarly specialized ear morphology, with large external ears, and this specialized hearing apparatus is used for amplifying sounds and detecting vibrations. Springhares, for instance, are known to rest with the top of their head pressed against the ground, probably so they can pick up the vibrations of an approaching predator. Of the three desert-dwelling cats, the sand cat is the most specialized and has the most sensitive hearing. The position and size of the sand cat’s external ear increases the accuracy of location of the sound, and the structure of the middle ear is specialized to absorb more low-frequency sounds. In addition to listening for prey, these desert-living cats also use their sensitive hearing to keep in touch with one another. Both sand cats and black-footed cats have extremely loud, deep voices for such small cats. Their barking calls are long, impressively loud, and low in pitch—low-frequency sounds travel farther in dry open habitats. Males and females use their loud calls to find each other during the mating season, and territorial males probably call to inform rival males where they are, just as roaring lions do.

s a n d c a t   235

One of the few cats that dig burrows, sand cats are found among the sand dunes and rocky outcrops of the Sahara Desert and the Middle East.

enlarged ear chambers suggest that they rely on sound to find prey. Almost anything from gerbils to geckos and birds to beetles is potential prey to be stalked, pounced on, or chased. They search for prey while walking and listening and often travel long distances. One radio-collared male was followed as he walked 5 miles (8 km) in a single

night, and then returned to his burrow the following evening. In some areas, reptiles play a major role in the sand cat’s diet. Saharan nomads say these little cats are snake hunters and regularly kill horned vipers and sand vipers. The nomads call them “the cat that digs holes,” in contrast to the African wildcat, which does not dig.

23 6   d o m e s t i c c a t l i n e a g e

Distribution of the sand cat.

Throughout its range, the sand cat makes its home in burrows—either occupying abandoned burrows of foxes or porcupines, or enlarging gerbil and ground squirrel burrows. In cooler weather, before retiring for the day, they often spend time sunbathing at the entrance to their burrow. Sand cats growl and spit much like domestic cats, but some of their other vocalizations are quite different. They have a loud barking call that has been likened to the yelping noise made by a small dog. Males

use the barking call during the mating season, but in zoos both sexes bark. These loud calls help males and females to find each other in the desert, where individual cats live far apart. In the Sahara, nomads say that they see sand cats around dusk, just after milking time, drinking camel’s milk from storage gourds. The nomads also say that these cats quite often kill and eat chickens and are sometimes caught in traps set in front of chicken enclosures. Despite occasional conflicts, sand cats

s a n d c a t   2 37

are generally not persecuted in this part of their range; rather, they are respected by the local people—Muslim tradition has it that, like the eagle owls and hoopoes, sand cats were the companions of the prophet Mohammed and his daughter Fatima. Despite the fact that these beautiful little cats make highly attractive exhibits, the species is difficult to keep in captivity and you will rarely see them in zoos.

status: IUCN Red List—Near Threatened weight: 3–7.5 pounds (1.4–3.4 kg) head-body length: 15–20 inches (39–52 cm) tail length: 9–12 inches (23–31 cm) litter size: 2–5 kittens, but usually 3

23 8  d o m e s t i c c a t l i n e a g e

Jungle Cat FE LI S C H AUS

More than 2,000 years ago, the Egyptians venerated, tamed, and even trained cats, depicting them in wall paintings and statues. Some cats were used to protect granaries against rats and mice, while others were trained for hunting. Among the mummified cat remains that have been examined, scientists have identified two species: the African wildcat and the larger jungle cat. However, of 190 mummified remains examined, only three were those of jungle cats. While the Egyptians may have tamed the jungle cat and kept it for its rodent-catching ability, there is no evidence of true domestication of this species. It is, however, in the domestic cat lineage, along with the wildcat, sand cat, and black-footed cat. A more appropriate name for the jungle cat might be “swamp cat” or “reed cat”—as this cat is not usually found in dense jungle, The jungle cat is the most common small felid throughout Asia. These adaptable cats survive in cultivated areas near villages and croplands, where they hunt rodents.

but prefers tall grass, thick brush, swamps, and reed beds. Jungle cats are good swimmers and are clearly not averse to getting wet. Over much of the Indian portion of its range, the jungle cat lives in dry open forests and grasslands. They can survive where there is only sparse vegetation and limited water, and they sometimes settle in cultivated areas around villages. In many parts of Asia, the jungle cat is the small-cat equivalent of the jackal. The cat’s rodent-catching abilities, wide habitat tolerance, and adaptability to living alongside villages and agricultural crops make it the most common of the small felids. Jungle cats are larger and lankier than domestic cats. Like the caracal and the African wildcat, the jungle cat also has a plain unspotted coat, but adults often retain some of the kitten spots or stripes on their legs. Black jungle cats occur regularly in southeastern Pakistan and in India. Jungle cats are not as strictly nocturnal as many cats and are often seen hunting in the early morning and late afternoon. They make 239

The jungle cat’s unspotted, grayish-brown coat blends well in dry woodlands and reed beds, the typical haunts of this cat.

j u n g l e c a t   241

Distribution of the jungle cat.

high, nearly vertical jumps in attempts to capture birds and also use long, arching leaps to spring onto rats and birds in tall dense cover. Small mammals—principally rats, mice, and gerbils—are their primary food; one study found that each jungle cat eats three to five rodents per day. Birds rank second in importance. In southern Russia, waterfowl are the mainstay of the jungle cat’s winter diet. Where overwintering populations of waterfowl congregate in huge numbers on unfrozen rivers and marshes, the jungle cat hunts among reed beds and along edges of wetlands, searching for injured or weakened birds. Jungle cats also eat lizards and small snakes, and, in turn, are sometimes food for larger snakes. In Pakistan a jungle cat was found entwined by a cobra; both animals were dead, and there were signs of a fierce fight. In India the body of a jungle cat was recovered from the stomach of a python. In addition to this eclectic menu, jungle cats also eat fruit. In a study in southern Uz-

bekistan, the fruits of the Russian olive made up 17 percent of their diet in winter. Where they live next to villages, the cats regularly prey on chickens, ducks, and geese. The jungle cat has been heavily hunted and trapped, and it is considered threatened in eastern Europe. However, the cat’s generalist diet and ability to survive in a variety of habitats, including those altered by humans, makes its continued survival quite likely. status: IUCN Red List—Least Concern weight: 10–20 pounds (5–9 kg) head-body length: 23–30 inches (58–75 cm) tail length: 8–11 inches (21–27 cm) litter size: usually 3–4 kittens, as many as 6

Acknowledgments

We have been privileged to work with cats in many different ecological settings, and with people who were willing to share their ideas, thoughts, and insights from their own work on cats. Our understanding of felids has benefited greatly from conversations and the occasional “wee dram” with the late John Eisenberg, Devra Kleiman, and Griff Ewer; what a wealth of intellectual capital we were able to tap. We have thoroughly enjoyed the friendship, the stories, and the many shared experiences with the tiger people, especially John Seidensticker, Ullas Karanth, Dave Smith and Francie Cuthbert, Hementa and Sushma Mishra, Chuck McDougal, Eric Dinerstein, Dale Miquelle, Linda Kerley, Kae Kawanishi, Peter Jackson, A. J. T. Johnsingh, Mahendra Shrestra, Ron Tilson, Philiip Nyhus, Valmik Tapar, Margaret Kinnaird, Tim O’Brien, Eric Wikramanayake, and Per Wegge. Rod Jackson, Tom McCarthy, Leif Blomqvist, and the late Helen Freeman were always willing to discuss the ecology and conservation issues surrounding snow leopards. Rajan and Lynette Rajaratnam introduced us to the cats on Borneo, and Lon Grassman, Sean Austin, Jeremy Holden, and Alan Rabinowitz filled in many gaps in our knowledge of South-

east Asian felids. We thank them all for their patience. It has also been extremely enlightening to interact with scientists working on cats in Europe and Africa, including Juan Beltran, J. Du P. Bothma, Chris and Tilde Stuart, Urs Breitenmoser and Christine Breitenmoser-Würsten, Helmut Hemmer, Marna Herbst, Luke Hunter, Paul Funston, Andrew Kitchener, Ted Bailey, Gus Mills, Laurence Frank, Justina Ray, Alex Sliwa and Gea Olbricht, Philip Stander, Philippe Stahl, Krzystof Schmidt, Olof Lieberg, and Lars Verdelin. We can’t image better tutors than Laurie Marker and Tim Caro to help us understand cheetah ecology. Similarly, our understanding of Central and South America felids has been enriched by conversations with Andres Novaro and Susan Walker, Daniel Scognamillo and Inés Maxit, Laura Farrell, Joe Roman, Mark Ludlow, Rafael Samudio, Anthony Giordano, Francisco Bisbal, Marcelo Aranda, Roland Kays, William Franklin, Marianela Velilla, Mario Di Bitetti, John Polisar, Anthony Novack, Tadeu de Oliveira, Eduardo Silva, Andrew Taber, Andy Noss, Agustin Iriarte, Rafael Hoogestijn, Arturo Caso, Michael Tewes, Howard Quigley, Alan Rabinowitz, Jim Sanderson, Louise Emmons, Peter Crawshaw, Kent Redford, Lyn 243

24 4  a c k n o w l e d g m e n t s

Branch, Brian Miller, and the late Edgardo Mondolfi and Tomas Blohm. The collegiality of Martin Jalkotzy, Gary Koehler, Ken Logan, Linda Sweanor, Maurice Hornocker, Chris Belden, Darrel Land, Dave Onorarto, Madan Oli, Jeff Hostetler, Roy McBride, and the late Dave Maehr and Ian Ross is greatly appreciated. They were always willing to share information and to talk about pumas. We also appreciate the cooperation of Kim Poole and Brian Slough regarding requests for information on Canada lynx. We greatly appreciate the intellectual contributions of Jill Mellen, Kristen Nowell, Chris Wemmer, John Gittleman, George Schaller, Eduardo Eizirik, David Wildt, Gustav Peters, Blaire Van Valkenburgh, Todd Fuller, Ji Mazak, Stephen O’Brien, Warren Johnson, and Carlos Driscoll to our understanding of felid ecology, behavior, morphology, and genetics. What an education it has been. Thanks also to Pat Callahan and William Swanson at the Cincinnati Zoo for information on the behavior of cats in captivity. Their observations added some nice touches to the stories.

We would like to thank the following for generously contributing their photographs for this book: Dharmendra Khandal, Ullas Karanth, Eleanor Briggs, Rod Jackson, Mohammad Farhadinia, Andy Hearn and Joanna Ross, Laurie Marker, Andres Wilting and Azlan Mohamed, Fernando Vidal, Linda and Peter Hoch, Johanna Turner, Jeremy Holden, Sebastian Kennerknecht, Tadeu de Oliveira, Alex Sliwa, and Jim Sanderson. We also greatly appreciate the contribution of Dale Johnson for her professional touch on the figure and the range maps. We also want to thank Betty Romano, Bev Ritter, Frank Cellon, Claire Sunquist, and Travis Blunden for always being there, for keeping us on track, and being a constant source of encouragement. Lastly, we want to thank Terry Whittaker, who graciously and professionally handled all the logistics of finding the right pictures for the species accounts, and for his patience. We are sure he would have rather been doing something more enjoyable, like taking more of his own marvelous pictures.

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Pallas’s Cat Barashkova, A., and I. Smelansky. 2011. “Pallas’s Cat in the Altai Republic, Russia.” Cat News 54: 4–7. Brown, M., and B. Munkhtsog. 2000. “Ecology and Behaviour of Pallas’s Cat in Mongolia.” Cat News 33: 22. Fox, J. L., and T. Dorji. 2007. “High Elevation Record for Occurrence of the Manul or Pallas Cat on the Northwestern Tibetan Plateau, China.” Cat News 46: 35. Murdoch, J. D., T. Munkhzul, and R. P. Reading.

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Additional Readings

Bradshaw, J. 2013. Cat Sense: How the New Feline Science Can Make You a Better Friend to Your Pet. New York: Basic Books. Breitenmoser-Würsten, C, E. Hofer, K. Vogt, and U. Breitenmoser, eds. 2011. “Cats of the World— Snapshots.” Cat News, Special Issue No. 6. Brown, D. E., and C. A. López González. 2001. Borderland Jaguars. Salt Lake City: University of Utah Press. Divyabhanusinh. 2005. The Story of Asia’s Lions. Mum­bai: Marg Publications. Hornocker, M., and S. Negri, eds. 2010. Cougar: Ecolo­gy and Conservation. Chicago: University of Chicago Press. Hunter, L., and G. Hinde. 2005. Cats of Africa: Behavior, Ecology, and Conservation. Baltimore, MD: Johns Hopkins University Press. Karanth, K. U. 2011. The Science of Saving Tigers. Hyderabad, India: Universities Press. Logan, K. A., and L. L. Sweanor. 2001. Desert Puma: Evolutionary Ecology and Conservation of an Enduring Carnivore. Washington, DC: Island Press. Long, R. A., P. MacKay, W. J. Zielinski, and J. C. Ray, eds. 2008. Noninvasive Survey Methods for Carnivores. Washington, DC: Island Press. Loveridge, A. J., S.W. Wang, L.G. Frank, and J. Se-

idensticker. 2010. “People and Wild Felids: Conservation of Cats and Management of Conflicts.” In Biology and Conservation of Wild Felids, ed. D. W. Macdonald and A. J. Loveridge, 161–95. Oxford: Oxford University Press. Macdonald, D. W., and A. J. Loveridge, eds. 2010. Biology and Conservation of Wild Felids. Oxford: Oxford University Press. Sanderson, J. G., and P. Watson. 2011. Small Wild Cats: The Animal Answer Guide. Baltimore, MD: Johns Hopkins University Press. Seidensticker, J., S. Christie, and P. Jackson, eds. 1999. Riding the Tiger: Tiger Conservation in Human-Dominated Landscapes. Cambridge: Cambridge University Press. Seidensticker, J., and S. Lumpkin. 2004. Cats: Smithsonian Answer Book. Washington, DC: Smithsonian Books. Sunquist, M., and F. Sunquist. 2002. Wild Cats of the World. Chicago: University of Chicago Press. Tilson, R., and P. J. Nyhus, eds. 2010. Tigers of the World: The Science, Politics, and Conservation of Panthera tigris. 2nd ed. London: Elsevier. Wison, D. E., and R. A. Mittermeier, eds. 2009. Handbook of the Mammals of the World. Vol. 1, Carnivores. Barcelona: Lynx Edicions.

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Illustration Credits

Photos by Terry Whittaker: pp. ii–iii, viii, 4, 13, 16, 18, 25, 28, 31, 38, 40, 42, 44, 51, 60, 62, 70, 72, 74 (second image from bottom), 76, 80, 84, 88, 92, 94–95, 98, 100, 102, 104, 105, 106, 108, 110, 111, 118, 124, 134, 157, 174, 180, 182–83, 184, 186, 188, 189, 190, 191, 198, 200, 202–3, 218, 220, 224, 227, 228, 230, 232, 234, 235, 238, 240 Adapted from Stephen J. O’Brien and Warren E. Johnson, “The Evolution of Cats,” Scientific American (July 2007), p. 70, Fig. “The Cat Family Tree”: p. 2 Photo by Rian van den Berg/ Shutterstock.com: pp. 6–7 Photo by Mogens Trolle/Shutterstock.com: p. 8 Photo by Alta Oosthuizen/Shutterstock.com: p. 10 Photo © Luiz Claudio Marigo/ naturepl.com: pp. 20–21 Photo by Vadim Petrakov/Shutterstock.com: p. 23 Photo by Eduardo Rivero/Shutterstock.com: p. 23 Photos by Dharmendra Khandal: pp. 30, 34, 86, 89 Photo by vanchai/Shutterstock.com: p. 30 Photo by visceralimage/ Shutterstock.com: pp. 32–33 Photo © Andy Rouse/naturepl.com: p. 36 Photo by Rodney Jackson: p. 46 Photo © Andy Rouse/naturepl.com: p. 48 Photo by Johan W. Elzenga/ Shutterstock.com: p. 50 Photo by Jeremy Richards/ Shutterstock.com: pp. 52–53 Photo by Dennis Donohue/ Shutterstock.com: pp. 54–55 Photo by Daleen Loest/Shutterstock.com: p. 57 Photos by Andrew Hearn and Joanna Ross: pp. 63, 71, 74 (marbled cat), 192 Photo by Sebastian Kennerknecht: p. 68 Photos by Eric Isselee/Shutterstock .com: pp. 82, 214 Photos by Fernando Vidal, Fauna Andina: pp. 114, 117

Photos by Jim Sanderson: p. 120 Photo © iStockphoto/Tim Abbott: p. 122 Photos by Tadeu de Oliveira: p. 126 Photo © Gabriel Rojo/naturepl.com: p. 128 Photo by Volodymyr Burdiak/ Shutterstock.com: p. 132 Photo by Erik Mandre/Shutterstock.com: p. 135 Photo by Wolfgang Kruck/ Shutterstock.com: p. 136 Photo © Jose B. Ruiz/naturepl.com: p. 138 Photo by FloridaStock/Shutterstock.com: p. 142 Photo by Keith Williams: p. 144 (top) Photo by nialat/Shutterstock.com: p. 144 (bottom) Photo by Andre Klopper/ Shutterstock.com: pp. 148–49 Photo by Scott E. Read/Shutterstock .com: p. 146 Photo © Peter Blackwell/naturepl.com: pp. 154–55 Photos by Stuart G. Porter/Shutterstock .com: pp. 158–59, 160 Photo by mdd/Shutterstock.com: p. 152 Photo by Gail Johnson/Shutterstock.com: p. 161 Photos by S. R. Maglione/Shutterstock .com: pp. 164, 166–67 Photo by Tom Tietz/Shutterstock.com: p. 168 Photo by Peter Hoch: p. 173 Photo © Visuals Unlimited/naturepl.com: p. 176 Photo by Huangdi: p. 185 Photo © Nick Garbutt/naturepl.com: p. 196 Photo by Jeremy Holden/FFI: p. 199 Photo by Jane Burton/naturepl.com: p. 204 Photo by Damien Richard/ Shutterstock.com: p. 206 Photo by pics4sale/Shutterstock.com: p. 208 Photo by Sari O’Neal/Shutterstock.com: p. 209 Photo by Artem Loskutnikov/ Shutterstock.com: p. 211 Photo by Linn Currie/Shutterstock.com: p. 213 Photo by Seiji/Shutterstock.com: p. 216 Photo by Alex Sliwa: p. 221 261

262  i l l u s t r a t i o n c r e d i t s Photos © iStockphoto/EcoPic: pp. 224, 229 Adapted from Carlos A. Driscoll et al., “The Near Eastern Origin of Cat Domestication,” Science 317 (July 2007), fig. 1 (A), p. 520: p. 226 Photo by HTO: p. 6 Photo by Pierre-Jean Durieu/ Shutterstock.com: p. 9 Photo by EcoPrint/Shutterstock.com: p. 11 Photo by Michael Sheehan/ Shutterstock.com; p. 12 Photo by Sharon Morris/Shutterstock.com: p. 24 Photo © Tony Heald/naturepl.com: p. 35 Photo by Villiers Steyn/Shutterstock.com: p. 56 Photo by Mohammad Farhadinia: p. 156 Photo courtesy of Cheetah Conservation Fund: p. 158 Photo by maryo/Shutterstock.com: p. 194

Photo by Diane Henry: p. 210 Photo courtesy of WCS India Program: p. 43 Photo © Yves Lanceau/naturepl.com: p. 116 Photo by Zhiltsov Alexandr/ Shutterstock.com: p. 178 Black leopard spots photo by Eric Isselee/ Shutterstock.com, leopard spots photo by bluehand/Shutterstock.com, Jaguar spots photo by worldswildlifewonders/Shutterstock. com, cheetah spots photo by Jerome Scholler/ Shutterstock.com, clouded leopard spots photo by Vladimir Sazonov/Shutterstock .com: p. 74 Photo by Scott Granneman: p. 215 Photo by Vishnevskiy Vasily/ Shutterstock.com: p. 212

Index

Note: Page references in italics indicate illustrations. Abyssinian cat, 217 Acinonyx jubatus. See cheetah Africa: Chobe National Park, Botswana, 8; Kalahari, 5, 11, 49–52; lion populations in, 10, 14, 14; Nairobi National Park, 53; Namib desert, 11; Namibia, 158; Selous Game Reserve, 14; Serengeti, 81 African golden cat (Caracal aurata), 92–96; Asiatic golden cat and, 77, 93; coat of, 93; color phases, 93; diet of, 93; distribution of, 96 African wildcat, 225–31, 229; breeding habits of, 228; distinguishing features of, 226–27; distribution of, 226; domestication of, 226; domestic cat and, 230–31; habitats, 227; hunting and, 227; vocalizations of, 228–30 allergies, pets and, 209 Amur leopard, 51, 51 Anatolian shepherds, livestock predation and, 158 Andean cat (Leopardus jacobita), 120–23; build of, 121; conservation of, 123; diet of, 122, 123; distribution of, 123; as Endangered species, 123; fearless nature of, 121; habitat of, 121–22; pampas cat and, 129; range of, 121–22; viscachas and, 122, 122–23 Andean Cat Alliance, 123 Asian wildcat, 227 Asiatic cheetah, 156, 156 Asiatic golden cat (Pardofelis temminckii), 76–78; African golden cat and, 77, 93; diet of, 77; distribution of, 78; habitat of, 77 Balinese cat, 217 Bastet (Egyptian cat goddess), 211, 211 bay cat (Pardofelis badia): on Borneo, 68, 69; conservation of, 70; discovery of, 69; distribution of, 71; trapping of, 69–70 bay cat lineage: Asiatic golden cat, 76–78; bay cat, 68–71; marbled cat, 72–75 Bengal cat, 111, 195

Bengal tigers, 37 black-footed cat (Felis nigripes), 218–23; breeding habits of, 222; build of, 219; conservation of, 222–23; distribution of, 223; habitat of, 219; hearing of, 234; hunting technique of, 219, 221–22; litters, 221, 221; range of, 219; scent marking by, 222; as solitary cats, 222; vocalizations of, 222 Blackfooted Cat Working Group, 222–23 black panthers, 169 black pumas, 169 bobcat (Lynx rufus), 146–50; breeding habits of, 148–49; build of, 148; Canada lynx and, 143–44, 147; diet of, 147; distribution of, 150; habitat of, 147; kittens of, 150; trapping of, 148, 150 boma (livestock enclosure), 15 Bornean Clouded Leopard Programme, 63; bay cat conservation and, 70; flat-headed cat and, 197; leopard cat and, 195; marbled cats and, 73, 75. See also conservation Borneo: bay cat on, 68, 69; clouded leopards on, 63 breeding: captive-breeding program of Iberian lynx, 140–41; hybrid cats, 111, 113, 195, 217; selective breeding of domestic cat, 217 Calvin Klein Obsession perfume, 73, 100 camera trapping, 43, 190 Canada lynx (Lynx canadensis), 133, 142–45; bobcats and, 143–44, 147; build of, 143, 144; distribution of, 145; hunting technique, 145; populations, 144–45; snowshoe hare and, 143–45, 144; snow tracking and, 145; specialized hunting and, 143; trapping of, 145 cannibalism, pumas and, 170–71 captive-breeding program of Iberian lynx, 140–41 captivity: fishing cat in, 187, 189; flat-headed cat in, 197; Pallas’s cat in, 185; rusty-spotted cat in, 201; snow leopard in, 44; tigers in, 31 capybara, 23 263

26 4  i n d e x caracal (Caracal caracal), 86–91; appearance of, 87, 89; build of, 89; diet of, 87, 89, 91; distribution of, 90; hunting and use of, 87; hybrid cats and, 111; jump of, 90; livestock predation by, 91; lynx and, 87; range of, 87 Caracal aurata. See African golden cat Caracal caracal. See caracal Caracal lineage: African golden cat, 92–96; caracal, 86–91; serval, 80–85 Caracat, 111 CatBib, 210 catnip, 73 cave lions, 7 chamois, 136 Chanel No. 5 perfume, 100 Chausie, 111 cheetah (Acinonyx jubatus), 152–63; Asiatic, 156, 156; build of, 153, 155; claws, 153; conservation of, 158, 162–63; diet of, 155–56; distribution of, 162; hunting technique of, 156; kills and, 156, 160; king, 157, 157; lions and, 162–63; litters of, 161, 161–62; livestock predation and, 158; markings of, 74, 160–61; other big cats and, 155, 162; pumas and, 165; range of, 155; speed of, 153, 155; vocalizations of, 161–62 Cheetah Conservation Fund, 158 Chinese mountain cat (Felis silvestris bieti), 225; distribution of, 226 Chobe National Park, Botswana, Africa, 8 CITES laws. See Convention on International Trade in Endangered Species laws clouded leopard (Neofelis nebulosa), 60–65; on Borneo, 63; build of, 61; canine teeth, 61, 62; coat of, 61, 63; diet of, 61; distribution of, 65; saber-toothed cats and, 62; species of, 63–65; on Sumatra, 63; in trees, 61; vocalizations of, 64 coat patterns: spotted, 74; tabby, 157, 217. See also specific species color morphs: black cats and, 24, 24; inbreeding and, 12, 38; of jaguarundi, 175, 176; white lions and, 12; white tigers and, 38 communication: ear tufts and, 133; physical, 70; purring, 178; scent marking as, 222; tail, 216 conservation: of Andean cat, 123; of bay cat, 70; of black-footed cat, 222–23; camera trapping and, 43; of cheetah, 158; of flat-headed cat, 197; of Florida panther, 43; of Iberian lynx, 140–41; of jaguar, 26; of leopard cat, 195; of lions, 15; of snow leopards, 45; of tigers, 31, 35 Convention on International Trade in Endangered Species (CITES) laws, 109 Critically Endangered species, Iberian lynx as, 139. See also Endangered species

cross-species mating, 178 cubs: infanticide and, 58, 170–71; males cats raising, 58 Cyprus, domestic cats on, 208–10 designer cats, 111. See also hybrid cats DNA of scat, 122 domestic cat (Felis silvestris catus), 206–17; African wildcat and, 230–31; breeds of, 213, 213–14, 217; cardboard boxes and, 208; coat colors of, 214, 217; in Cyprus, 208–9; in Egypt, 209–11; European wildcats and, 231; feral, 215, 215; history of, 207–10; nautical cat associations, 211–13; persecution of, 216–17; popularity of, 214; selective breeding of, 217; significance of, 208–9; spread of, 211–14; wildlife predation and, 210, 212; witchcraft and, 216–17 domestic cat lineage: black-footed cat, 218–23; domestic cat, 206–17; jungle cat, 238–41; sand cat, 232–37; wildcats, 224–31 ear position in cats, 70 ear tufts, 133 ecotourism, 14–15 Egypt: Bastet and, 211, 211; domestic cats in, 209–11 Eizirik, Eduardo, 24 Eloff, Fritz, 11 Endangered species: Andean cat as, 123; fishing cat as, 190; jaguar as, 19; snow leopard as, 46–47 Eurasian lynx (Lynx lynx), 132–37; build of, 133–35; chamois and, 136; diet of, 135, 135, 136; distribution of, 137; ear tufts of, 133; humans and, 136; hunting technique of, 135–36; livestock predation by, 137; range of, 135; reintroduction of, 136–37; roe deer and, 135 Eurasian wildcat (Felis silvestris ornata), 225; distribution of, 226 European jaguar, 17 European rabbit, 140 European wildcat (Felis silvestris silvestris), 225, 228; distribution of, 226; domestic cats and, 231; hunting of, 230, 231; hybridization, 231; range of, 231 eyesight, 201, 204 falling from great heights, 116 Felis chaus. See jungle cat Felis margarita. See sand cat Felis nigripes. See black-footed cat Felis silvestris. See wildcats Felis silvestris bieti. See Chinese mountain cat Felis silvestris cafra. See Southern African wildcat Felis silvestris catus. See domestic cat Felis silvestris lybica. See Near Eastern wildcat Felis silvestris ornata. See Eurasian wildcat

i n d e x   265 Felis silvestris silvestris. See European wildcat feral cats, 215, 215 fishing cat (Prionailurus viverrinus), 186–91; build of, 187; in captivity, 187, 189; distribution of, 187, 191; as Endangered species, 190; genetics of, 189; habitat of, 187, 189–90; hybrid cats, 111; population declines of, 190; specialized hunting and, 187; swimming ability of, 187, 190, 191, 198; trapping of, 190 flat-headed cat (Prionailurus planiceps), 196–99; build of, 197; in captivity, 197; conservation of, 197; distribution of, 199; swimming ability of, 197–98, 198 Florida panther: black, 169; conservation of, 43; range of, 170; vehicle collisions and, 170. See also puma Freer, Rachel, 115 Geertsema, Aadje, 84 gemsbok, danger of hunting, 11 genetics: of cat species, 2; of fishing cat, 189; melanism and, 24, 24; of puma, 165; white tigers and, 38 Geoffroy’s cat (Leopardus geoffroyi), 108–13; coat of, 109; diet of, 109, 112–13; distribution of, 112; hunting technique, 113; hybrid cats, 111, 113; poaching of, 109 Global Positioning System (GPS) radio collars, 22 Global White Lion Protection Trust, 12 GPS radio collars. See Global Positioning System radio collars guar, 30 guarding dogs, 158 guiña (Leopardus guigna), 114–19; black, 115; decline of, 118; diet of, 115; distribution of, 119; livestock predation by, 116, 118; range of, 115 harimau-dahan. See clouded leopard hearing, specialized, 81–83, 234 homing cats, 167 human encounters with cats: Eurasian lynx and, 136; jaguars and, 22; leopards and, 53, 57–59; man-eating cats, 22, 57–59; pumas and, 171, 173 hunting: caracals used for, 87; Eurasian lynx and, 137; of European wildcat, 230, 231; infanticide and, 58; males and, 58; of Pallas’s cat, 185 hybrid cats, 217; Bengal cat, 111, 195; Caracat, 111; Chausie, 111; Jambi, 111; Safari cat, 111, 113; Savannah cat, 111; Stone Cougar, 111 hyenas, cheetahs and, 162–63 Iberian lynx (Lynx pardinus), 133, 138–41; build of, 140; captive-breeding program, 140–41; conservation of, 140–41; as Critically Endangered, 139; distribution of, 141; kittens of, 140; rabbits and, 139–40; specialized hunting and, 139–40 inbreeding, color morphs and, 12, 38

India, Sanjay Gandhi National Park, 52 infanticide: lions and, 58; pumas and, 170–71 Issoire lynx, 133, 139 IUCN Red List, 63 Jackson, Rodney, 45 jaguar (Panthera onca), 16–27; black, 24, 24; build of, 18–19; conservation of, 26; diet of, 18–22, 23; distribution of, 26; European, 17; human encounters with, 22; killing technique of, 19; leopard differentiated from, 17; livestock predation by, 22, 26–27; markings of, 74; melanism in, 24, 24; in North America, 19; physical characteristics of, 17–18; prey selection, 22; range of, 19, 22; roar of, 22; skin trade of, 18; society of, 22; in South America, 17; strength of, 17; survival of, 26; swimming ability of, 22, 198; vocalizations of, 22 jaguarundi (Puma yagouaroundi), 173–77; build of, 175; color morphs, 175, 176; diet of, 175; distribution of, 177; hunting technique of, 175; range of, 176 Jambi, 111 jungle cat (Felis chaus), 238–41; adaptability of, 241; diet of, 241; distribution of, 241; domestication of, 239; habitat of, 239; hunting technique of, 239, 241; hybrid cats and, 111; physical characteristics of, 239, 240 Kalahari desert, Africa: leopards in, 49–52; lions in, 5, 11 Karanth, K. Ullas, 43 king cheetah, 157, 157 Koshkarev, Eugene, 46 Kruger lions, 12 landing on feet, cats and, 116 leopard (Panthera pardus), 48–59; adaptability of, 52, 57, 59; Amur, 51, 51; climbing skills of, 54, 54, 56; desert, 49–52; diet of, 49, 52; distribution of, 59, 59; female, 56; humans and, 53, 57; hunting technique of, 50, 50–52; in India, 52; jaguar differentiated from, 17; in Kalahari, 49–52; man-eating, 57–59; markings of, 49, 74; in Nairobi National Park, Africa, 53; physical characteristics of, 49; in Sanjay Gandhi National Park, India, 52; secretive habits of, 57; as solitary cat, 52; in trees, 54, 54, 56–57 leopard cat (Prionailurus bengalensis), 192–95; build of, 193; conservation of, 195; diet of, 193; distribution of, 195; habitat of, 194; hunting technique of, 194–95; hybrid cats, 111, 195; Iriomote cat population, 194; range of, 193; swimming ability of, 193–94, 198 leopard cat lineage: fishing cat, 186–91; flatheaded cat, 196–99; leopard cat, 192–95; Pallas’s cat, 180–85; rusty-spotted cat, 200–204

26 6  i n d e x Leopardus colocolo. See pampas cat Leopardus geoffroyi. See Geoffroy’s cat Leopardus guigna. See guiña Leopardus jacobita. See Andean cat Leopardus pardalis. See ocelot Leopardus tigrinus. See oncilla Leopardus wiedii. See margay Leptailurus serval. See serval liger, 178, 178 lion (Panthera leo), 5–15; cave, 7; cheetahs and, 162–63; conservation of, 15; diet of, 8; distribution of, 14; ecotourism and, 14; habitats of, 5, 11; hunting technique of, 8–10; infanticide and, 58; Kruger, 12; liger and, 178, 178; livestock predation by, 15; male, 9; in Namib desert, 11; populations of, 10, 14; prides, 5; range of, 10; roar of, 13, 13; Serengeti, 7; social system of, 5; tigers and, 5–6; tigon and, 178; white, 12 livestock predation: boma and, 15; caracals and, 91; cheetahs and, 158; Eurasian lynx and, 137; guard dogs and, 158; by guiña, 116, 118; by jaguar, 22, 26–27; lions and, 15; pumas and, 169; ranching practices and, 26–27; by snow leopard, 45 Lynx canadensis. See Canada lynx Lynx lineage: bobcat, 146–50; Canada lynx, 142–45; caracal and, 87; Eurasian lynx, 132–37; Iberian lynx, 138–41 Lynx lynx. See Eurasian lynx Lynx pardinus. See Iberian lynx Lynx rufus. See bobcat man-eating cats, 22; leopard, 57–59 “Man-Eating Leopard of Rudraprayag,” 59 marbled cat (Pardofelis marmorata), 72–75; build of, 73; distribution of, 75; range of, 73; tail of, 74; in trees, 73 margay (Leopardus wiedii), 102–7; agility of, 102; build of, 102; coat of, 106; diet of, 103–4; distribution of, 107; litters of, 104; markings of, 104–6, 106; ocelot and, 103–6; tree climbing, 103, 104, 105 markings, 74. See also specific cats matrilines, 56; pumas and, 171 melanism, 24, 24 Mohamed, Azlan, 73 molecular scatology, 123 Mukherjee, S., `87 Nairobi National Park, Africa, 53 Namib desert, Africa, 11 Namibia, Africa, 158 National Cancer Institute’s Laboratory of Genomic Diversity, 24 Near Eastern wildcat (Felis silvestris lybica), 225 Neofelis diardi. See Sunda clouded leopard

Neofelis nebulosa. See clouded leopard nepetalactone, 73 night vision, 201, 204 North America, jaguar in, 19 Nyhus, Philip, 31 O’Brien, Stephen, 24 ocelot (Leopardus pardalis), 98–101; coat of, 99, 106; diet of, 99; distribution of, 101; hunting technique of, 99– 100; kittens of, 100; margay and, 103–6; markings of, 74; range of, 99; swimming ability of, 198 ocelot lineage: Andean cat, 120–23; Geoffroy’s cat, 108–13; guiña, 114–19; margay, 102–7; ocelot, 98–101; oncilla, 124–27; pampas cat, 128–30 oncilla (Leopardus tigrinus), 124–27; diet of, 125; distribution of, 127; species of, 127 Otocolobus manul. See Pallas’s cat Packer, Craig, 5 Pallas’s cat (Otocolobus manul), 180–85; breeding habits of, 184–85; build of, 181; in captivity, 185; coat of, 181, 184; diet of, 184; distribution of, 185; habitat of, 181; hearing of, 234; hunting of, 185; hunting technique of, 181, 184; pikas and, 184, 185; populations of, 185; predators, 185 pampas cat (Leopardus colocolo), 128–30; Andean cat and, 129; diet of, 129; distribution of, 130; species of, 130 Panthera leo. See lion Panthera lineage, 3–65; clouded leopard, 60–65; jaguar, 16–27; leopard, 48–59; lion, 5–15; snow leopard, 40–47; tigers, 28–39 Panthera onca. See jaguar Panthera pardus. See leopard Panthera tigris. See tigers Panthera uncia. See snow leopard Pardofelis badia. See bay cat Pardofelis marmorata. See marbled cat Pardofelis temminckii. See Asiatic golden cat peccaries, 23 perfume and cats, 73 physical communication, 70 physical similarities between cat species, 1–2 pikas, 184, 185 poaching: of Geoffroy’s cat, 109; of snow leopards, 45–46. See also hunting Prionailurus bengalensis. See leopard cat Prionailurus planiceps. See flat-headed cat Prionailurus rubiginosus. See rusty-spotted cat Prionailurus viverrinus. See fishing cat puma (Puma concolor), 164–73; adaptability of, 169, 173; black, 169; build of, 165; cannibalism by,

i n d e x   267 170–71; cheetahs and, 165; continent-crossing, 166; diet of, 169; distribution of, 172; female, 171; fighting and, 170–71; Florida, 169; genetics of, 165; habitats of, 169; human encounters with, 171, 173; infanticide and, 170–71; kills, 169–70; livestock predation and, 169; male, 171, 173; matrilines and, 171; populations of, 168; range of, 167, 169; relocation and, 167; vocalizations of, 170; wildlife underpasses and, 170 Puma lineage: cheetah, 152–63; jaguarundi, 173–77; puma, 164–73 Puma yagouaroundi. See jaguarundi purring, 178 ranching practices, livestock predation and, 26–27, 45 relocating cats, 167 roe deer, 135 Russian Far East: Amur leopard in, 51, 51; tigers in, 29, 33, 37 rusty-spotted cat (Prionailurus rubiginosus), 200–204; in captivity, 201; distribution of, 204; hunting technique of, 201 saber-toothed cats, 62 Safari cat, 111, 113 Sanbona Wildlife Reserve, South Africa, 12 sand cat (Felis margarita), 232–37; burrows dug by, 235, 235–36; diet of, 235; distribution of, 236; habitats of, 233; hearing and, 234; hunting technique of, 233–35; physical characteristics of, 233, 234; vocalizations of, 234, 236 Sanderson, Jim, 121 Sanjay Gandhi National Park, India, 52 Savannah cat, 111 scat: DNA identification of, 122; molecular scatology, 123 scent marking, 222 selective breeding of domestic cat, 217 Selous Game Reserve, Africa, 14 Serengeti, Africa, servals in, 81 Serengeti lions, 7; manes of, 9; population of, 14 serval (Leptailurus serval), 80–85; build of, 81, 83; diet of, 83; digging, 83; distribution of, 85; doglike characteristics of, 85; hearing of, 81–83; hunting technique of, 83; hybrid cats, 111; leaping, 82, 83; mothers and kittens, 84; range of, 85; specialized hunting and, 81, 84–85 Siamese cat, 217 Siberian tigers, 37 Siegfried and Roy, 31 snow leopard (Panthera uncia), 40–47; breeding habits of, 44; in captivity, 44; conservation of, 45; diet of, 42; disposition of, 44–45; distribution of, 47; as

Endangered species, 46–47; livestock predation by, 45; physical characteristics of, 41, 42; poaching of, 45–46; range of, 46; vocalizations of, 42, 44 Snow Leopard Conservancy, 45 Snow Leopard Trust, 45 snowshoe hare, 143–45, 144 Somali cat, 217 South Africa, Sanbona Wildlife Reserve, 12 South America, jaguar in, 17 Southern African wildcat (Felis silvestris cafra), 225–31 specialized hunting: Canada lynx and, 143; fishing cat and, 187; Iberian lynx and, 139–40; serval and, 81, 84–85 species: cross-species mating, 178; genetics of cat, 2, 24, 24, 38; groups of related, 2; physical similarities between cat, 1–2 Stone Cougar, 111 sugar blindness, 125 Sumatra, clouded leopards on, 63 Sunda clouded leopard (Neofelis diardi), 63; distribution of, 64 Swanson, Bill, 185 swimming cats: fishing cat, 187, 190, 191, 198; flatheaded cat, 197–98, 198; jaguar, 22, 198; leopard cat, 193–94, 198; ocelot, 198; tigers, 198 tabby coat pattern, 157, 217 tail communication, 216 tapetum lucidum, 204 tigers (Panthera tigris), 28–39; Bengal, 37; in captivity, 31; conservation of, 31, 35; diet of, 30; distribution of, 39; female, 37, 56; flexibility of, 37; habitats of, 37, 39; hunting technique of, 29; killing technique of, 34, 37; liger and, 178, 178; lions and, 5–6; male, 37; physical characteristics of, 29, 34; range, 29, 37; in Russian Far East, 29, 32, 37; Siberian, 37; size of, 37; in snow, 29; stalking, 34; stripes, 29, 30; swimming ability of, 198; tigon and, 178; value of, 35; white, 38, 38; young, 37 tigon, 178 Tilson, Ron, 38 TNR programs. See trap-neuter-return programs touch-sensitivity, 118 trap-neuter-return (TNR) programs, 215 trapping: of bay cat, 69–70; of bobcat, 148, 150; of Canada lynx, 145; of fishing cat, 190. See also hunting tree-climbing cats: clouded leopards, 61; leopards, 54, 54, 56–57; marbled cats, 73; margay, 103, 104, 105; physical characteristics of, 105 United States, tigers living in, 31

26 8  i n d e x Varaday, Desmond, 74 vehicle collisions with cats, 170 vibrissae, 118 viscachas, 122, 122–23, 129 whiskers, 118 white lions, 12 white tigers, 38, 38

wildcats (Felis silvestris), 210, 224–31; African, 225–31; Asian, 227; distribution of, 226; divisions of, 225; European, 231; subspecies of, 225 wildlife predation: domestic cats and, 210, 212; feral cats and, 215 wildlife underpasses, 170 Wilting, Andreas, 73 witchcraft, cats and, 202, 204, 216–17