THE ULTIMATE REFERENCE TO EVERY SPECIES WORLDWIDE WILD CATS ©F THED id you know that Buropean royalty once used cheetahs to hunt deer, or that eara- calsean capture binds by leaping sixcand a half feet Straight up inte the air from a standing starr? Have you ever wondered wherher domestic cats really do always land on their feet when they fall, 6 how Canada Iynx ean stalk their prey in the ‘winter without falling through the snow? Wild Cats of the World isa treasure trove’ol an- swvers tO these questions, and many others, for anyone who's interested in learning more about the worlds felids, including the ones with whom ‘ve share our homes. Mel Sunqoist and Fiona Sunquist have spent more than a decade gather ing information about cats from every available source, many of them quite difficult to find. ‘They draw on historical documents such as de- scriptions of hunts and observations by natural ists and travelers, as well as more recent infor mation in scientific journals,, archaeological research, reports from government. agencies, -and newsletters fram awide variety of organiza tions, Weaving information from these sources together with their own experiences observing. ‘wild cats around the world, the Sunguists have created the most comprehensive reference on felids available. Each of their accounts of the thirty-six species of cat contains a description of the cat, including hurtin interactions with it, as ‘well as detailed dation distribution, ecology and behavior, statusin the wild, and conservation ef forts, Many photographs, incluiling more than forty in Full color, illustrate these accounts. From the two-pound black-footed cat to the five-hundred-pound tiger, aid from the African serval with its satellite-dish ears to the web- footed fishing cat of Asia, Wild Cats ofthe Would has information that will fascinate and educate felid fans of any stripe (or spotWild Cats of the WorldWild Cats of she World MelS}00-C2066-FM 2/27/02 2:58 PM Page iv Mel Suni is asin pefewor in the Depart of Widlic Eealoy and Conservation atthe Univesity of Fx, Gainesville Fiona Sunguit a eitr fer interatina Wile Magarin. The Sagus ae coauthors of Tier Meo, published bythe Union of Cheapo Fs a 1988, The Sus ca be eachal vs loonie wail to wiMeanavoldtac con Fromtipieve arian yx (roo by Try Whitahe) The Univerty of Chie Pras, Cag 60657 “The Leivennty of Chiao Pas, Le, Lom © 200 by Fons Samui and Mel Sonus, Allighus reserve Pubs 2002 ated in Chis | Mtwowoserceascsoae2 12345 ISBN 0.226-72999.8 (cloth) Lito of Congres CataipingrioePublisatin Data ‘Sooguist Mes E ‘Wild cats ofthe world Mel Sunqut an! Fons Sanus, pom Anche itive feroces aie ISON 0-226-77998(loth ale pape) 1 Flue. 1. Suga, Foe I, Tie QU787.€23 863 2002 59978—detl 2001082771 ‘Thich piel on ake pager Material protegido por derechos de peO0-COOSE-7m 2/a7/0a 3158 OR Page y Contents Preface ix Introduction 1 What lu a Cat? ‘The Essence of Cats 5 Taxonomy of the Cat Family 4 Species Accounts ® 7 8 ic golden cat 32 ‘Chinese desert cat 57 Jungle cat 60. Sand cat 87 Black-footed cat Bs Wildeat 83. European wildcat 5, African-Asian wildcat 92 Domestic ea » ‘Faguacundi 43 Oeeloe bo Oncilla 130 Mangay 35 Serval ae Material protagido por derechos 6(O0-COOSE-7M 3/27/00 3:88 En Page vi oO Lynx Canada lynx Eurasian lynx Iberian lynx Bobeat Pampas cat Geoffroy’ cat Kodkod Anwlean mountain at Manut Leopard cat Flat-headed eat Rusty-spotted cat Fishing cat African golden eat Puma Clouded leopard Lion Jaguar Leopard Tiger Marbled cat Saow leopand Stuy and Conservation Field Research Techniques: Recent Advances Relocating Cats: History and Guidelines for the Future. ‘Conserving Felids in the Twenty-First Century 152 154 7 185 205 25 20 335 233 27 24 252 xs 285 305 a8 M43 a m7 or derechos de auto(00-CROSE- he 2/27/02 2158 TH Page wit @ APPENDIX CITES Listings Governing Trade in Wild Cats 409 APPENDIX 2 IUCN Red List: Conservation Status of Wild Cats au APPENDIX 3 Olfactory Communication in Felids 43 APPENDIX 4 ‘Vocal Communication in Felids ar APPENDIX 5 Reproduction in Felids “5 Arrenmix 6 List of Scientific and Common Names Mentioned in the Text 435 Photographic Credits “4 Index 43 Color photographs follow pages 134 and 342 Material protegido por derechos00-CROSE- FH 2/27/02 JED ON Dege vELS © Material protegido por derechos de aO0-CROSE- FE 3/27/00 2158 Om wage ie Preface “his honk arose from our lfelomyg fascination wiht "Tete ver the ys we hve elected thouds ‘of publications on ats, en topics ranging fom the structure of the domestic cats mide eat to accounts ‘of livestock depredation by tigers in Malaysia. When, sve first hegan assembling this informal cat library, che “bible” for people interested in nding snformation on ‘cats was C. A. W. Guggitbergls Wild Cats ofthe World Someone suggested that we write an updated version, ‘The lea evolved into a book thae woul summarize ‘what isknown about the ecology and behavior ofeach ‘ofthe thiney-six species of wild ‘The purpose of this book is literature on eats 59 that readers will have aoe only ‘current review of what is known about the ecology and behavior of each cat, but ale a comprehensive source of reference material. It was very important to tus that the book serve asa thorcugh guide tothe lter- anure for stxlents and others who needed to dig fue- ther. We have deliberately included information fom the “gray” lnerature: unpublished reports, theses, dis- serrations, newsletters, and personal commanications. Where we have included mich information, we have sven the sources. oth the taxonomy and the onker of presentation ‘of the species accounts follow Wozenceafts clastic ‘cation in Mammal Species ofthe Wid (edited by Wil- son and Reeder). We have tried t fellow the same for- mt for exch species account so that topics are eaticr to find and crow-reference between specs. Suenmary tables and information on general topics such as scent ‘marking, vocaltations, and repwoduction are included as appendixes. Overall. our mnrent was to write a book that would be accesible to both scientists and the in- terened general publi ‘With our focus on ceology and behavior, there were many topics tac we did nce address. We did not have the expertise to include information on diseases and Parasites, reproductive physiology, captive research and management, and the rapidly evolving Feld of molecular gonetics. Information on status, recowery roxas, and actions recommended foe the conserva ‘Hon of fells are weated only briefly his book Readers are urged to consult Nowell and Jackson's Wild Car A Staras Surwey ond Consrastion Action Plan fora cent review of this material. Portions of this valuable comprchersive review are available at the IUCN Cat Specialist Group website at ep! Imxnionojeatfolk During this volume’s extremely lng gestation pe- nod we have had a great dal of hep from unde of generous people. ean McConville was an enorens Alp with all the editorial busines that need to be _xikesed. Liberian Kay Kenyon (Nationa Zoolog cal Park) and Nancy Matthews (Teanslation Publish- ing Prostam) at the Smithsonian Institution and Stephanie Haas and Veron Kisling atthe University of Flora helped tack down obscure references from actos the slobe. Gustav Peters conected ou dein tons and helped produce accurate tales of Fela vor calizations that would never have heen possible with cout his help. Warren Jolson kept uso the stig find narow wher it came t9 genetics and provided the diagram of fel lineages. Kerry Dresler patienely solved oar frequent computer problems. Stimulating dliecomtors with John Eisenberg speed ws to pres tw the finish. Other biologists, z0 sta an owes of privare collections unvelishl shared information on every aspect of Feld lives. Dale Jolson and Adan Freedman put together the figures. We would ike to thank che following indica for thee valuable con- ‘batons to the book: Bill and Penny Andeews, Mor- elo Arana, Sean Austin, Ted Bailey, Sonny Bass, Robert Bellen, Juan Beltcin, Ana Bennett, Lis Bens ett, Bill Beg, Francis Bisbal, Tomas and Cee Bohm, Lief Blomayist,.du Boehm, Anatole Bea sin, Urs Bretenmower and Chrixane Beeitermoser ‘Wanten, Rainer Brocke, Joan Burton, Pat Calls, 6 al protegido por derechos «i qutoO0-CO0EE-7H 2/37/02 I58 TH Page S ee ‘Tim Caro, Arturo Caso, Ravi Chellam, Sarah Chris- tie, Peter Crawshaw, Tor Dalimer, Erie Dinerstein, Tetsy Dresser, Siva Elgupillay, Louise Emmons, Laven Farrell, Dave Ferguson, Gail Foreman, Debra Forth: man, Neil Franklin, William Franklin, Helen Free- ‘man, Todd Fuller, Sam and Alicia Fulton, Ross Garcia-Peeea, John Gittleman, Lon Grassman, Judith, Hayter, Jeremy Holden, Rafnel Hoogesteija, Maurice Hoenocker, David Houston, Don Hunter, Agustin Iie arte, Masako Kava, Peter Jackson, Rasiney Jackson, Marti falkoty, David Jeany, A. JT Jobnsingh, Ullas Kaeant, Kae Kawanahi, Mohgesmed Khan, Ardeew Kitchener, Devra Kleiman, Gaty Koehler, Mike Konecny, Steven Laing, Paul Leyhausen, Fred Lindsey, ‘Olof Liberg, Ken Logan, Mark Ludiow, Dave Macht, Laurie Marker, Inés Maxit, Chuck McDougal, Jill Mallen, Heinrich Mendelisshn, Dennis Merit, Dale Miguelle, Brian Mille, Sriyanie Miththapala, Furi Misutams, Edganlo Mondo, LC. Moolman, Dieter Morsbach, Peter Noeton, Andees Nowaro, Kristin Nowell, Hideo Ohara, Steve O'Brien, Gea Ollriche, Tadeu Oliveira, Juraich Payne, Pablo Peeita, Kien Poole, Howaal Quigley, Pat Quillen, Alan Rabino- ‘wite, Rajan and Lynette Rajaratnam, Fateh Singh Rathore, Justina Ray, Kent Redford, Francisco Robles, Mask Rosenthal, jim Sanderson, Charles Santiaplli, George Schaller, Krystal Selunide, Daniel Seog: rnamillo, eho Seidensticker, Alan Shoemaker, Alex Shiwa, Brian Slough, James L. D. Srnith, Philippe Seal, Philip Stander, Chis and Tide Stuart, Claire Sunquist, Linda Sweanot, Anew Taber, Michael ‘Tewes, Valuuik Thapar, Ron Tillon, Buehars Tonkin Leyhausen, Ble Van Valkenburgh, Johar Visser, Clive Walker, Susan Walker, Dave Webb, Chis Wem: mer, Terry Whitaker, Vivian Wilson, Shigeki Yor suma, and Gunter Zesler And to Susan Abrams and all the wonderflly pas tient people at the University of Chicago Press, we thank you. al protegido por derechos «i qutoG2-CO0S5-IEF 2/27/02 2158 OH Page 2 Introduction 0d made the cat to give bumankind the pleasure of caressing the tiger. — Anon. sor ene of the mont facinating spect of cat Fatcg that tom the oq Hck fol ca tothe fve-wundeed-pound ger, cats ae all vaeiations ‘on a common theme. When you have a cat in your home, you live with the esence of tgs. Inded, che appeal of domestic cots may be that they are fst 9 sthisker away fro chee will relaives. Just how elose this relationship is, or how diferent the various spe- ‘ies. ae fom one another, can be discovered in what wwe have leamed over the years about the intrgans world of wil car Krorledge of the felils—the cat famsly—has inereasal siguicanay in the yast quartet century Treennytve years ago, when C. A. W. Guassbery snore his clase Wit Cats ofthe Wild the aye, the eadkod, and the Andean mouerain cat were known ‘nls from afew museurnskins ant an occasional stated specimen. The biology of ever easily rcoyptable species, such a the babat, ut, tiger, leopard, ehee- tah and ocelot, was virwally unknown, and nothing was known about what they needed in terms of space and focd. On the conservation front, hundreds of thousand corte eats were being tapped and thei kins tad in he woe’ fr mathe Tealay, weave basic information about the biology andbehavior of most cat species. Our increased know ‘dg has sven a greaereause for concern for some species and reasons tobe optimistic abou the sareval ‘of others. Information on reproduction and survival, ‘tuch ofr leaned irom captive nina, has shown ws that trapping and hunting willhave a heavy impact om many ofthe smaller spore cars. Species sch as the ccelot and marzay have very small liters and may beced only once every two years. Other secs sul as the ger, have turn! cut to he april seiient letheirhabitat and prey me protected, they wil have ‘even une low levels of poaching and hunting “Thou habitat ls, une, a ileal epg prey species threaten manyof the world’ cats, we have also mage some oujor conservation advances i the ‘past ewenty-five years. One ofthe most significant has been CITES, the Convention on International Trade in Endangered Species of Wild Fauna and Flora. In ef- fectsince 1975,CITESisaglobal treaty designed topro- tect plant and animal species fom unregulated trade. In the 1970s and 1980s a powerful combination of public interest and effective legislation succeeded ta shutting down the international trade in spotted eat skins. At the peak of the trade, nearly hal « mille, cat skins were traded on world markets each year To- day, internacional trade in pelts of sported cats has ale most disappeared, and trade in the remaining species is cleceeasing ‘This bonk i a compilation of what we know about the life histories, hwo, al ecology of the work's thirty-aix species of wild cas. I is intended asa refet- cence for al of those —cat lovers as well as professional scientists—who are fascinated by cats. In this book ‘you will nd informacion on each species, presented in sections on description, distribution, ecology and be- Ageior (which includes habitat, feeding evoleg; social organization, reproduction, and development) andl status in the wild. For species such as the tiger, lice, Jasvat, puma, bobeat, and the lynxes, the informa ton is fairly extensive; for others, such asthe Andean mountain at, bay cat, and marbled cat, our know cage remains limited. ‘Wehope this volume will serve asa stimulus for far- ther eseareh. As cat lovess, we represent a powerful constituency. Callectivelg. we can influence policy di- rect esearch funding, and help promote creative ways tosolve human-cat conflicts. Conservation ultimately depends on political will, which comes, in the end, from public interest. The more we know about these alluring creatures, the moge interested we become in their welfare and survival 6 Ma al protegido por derechos «i quto02-CO0S5-IEF 2/27/02 2158 mH vege? © Material protegido por derechos de aWhat Is a Cat?IOL-C2066 2/37/02 2155 OH Pages © Material protegido por derechos de aIL-OB0SS 2/37/02 2155 OH Pages The Essence of Cats MORPHOLOGY “The eat boy is, in essence, a reflection ofits diet. ATL ‘cats are meat eater, designed and specialized for cap ‘tweing and killing lve prey. They are sometimes called Aryperearmcores because they need a much higher peo portion of protein in the diet than almost any other rimmal. Fora domestic ent t remain healthy, ts dict ‘oust contaia 12 percent protein (by weight) for adult ‘eats and 18 percent for kittens. Dogs can survive on, much fess: ale dogs can gee by with only 4 percent protein. That is why dogs can survive and thrive on a ‘vegetarian dhet, while i is very diiull to create a eee ctorian dict that contains sufcient protein fora cat Cats have relatively short but powerful forelimbs that can he rotated, forepaws with Tong claws, a lithe a2. Pr ry) ea an bod length 09%) Frgwe Lor effet ond ly length etn a eth or wl lh lala flexible spine, and well-muscled hindlimbs, all of which combine to give them quickmess, agility and power, All cats are digitigrade; thar is, they walk on their toes. The soft toe pads distribute the weight over the balls of the fet, givingeats thet id walking mo~ tion. The cats skull is highly domed, the zygomatic arches are wide, the face w foreshortened, and stron sagital eres provide an anchor for powerul jaw ‘uscles. The mechanical advantage provided by the foreshortened face and powertul the bite force on the canine teeth. Excluding the shore-tailed bobeat and Iyex, most folids have tas that measure a least one-third to one- half oftheir head and body length (fg. 1). The mare sy, clout leopard, and marbled eat have exception- 302 5 —o— o gido por derechos de stoIL-O06S 2/37/02 2155 Om Page ee ally long tails, which are thought to funetion as bale rancing rou as they move about ia the tees. A similar fanction has been attributed to the long tls of sow Teopards, pumas, and cheetahs, but for these species the tal is prewamably an aid to moving about in pre= ipitous terrain or making rapid curs while pursing prey. Teeth With a shortened face comes a shortened ja with less space for teeth. Cats have fewer teeth than other ear- ssivores—ustlly 28 639, compared wih 42 for dogs aad bears. The teeth are specialized fo different tasks Small inciots are used for holding and nibbling, and de rear molars (or camassials) move against each ‘other like scissor blades for sbicing and cutting. The larg, strong, somewhat fattened canines are used for stabbing and delivering the killing bite. Ethologist Paul Leyhausen suggests thar the ear’ canine teeth fit benweea the neck vertebrae of prey like a “key in a Jock.” Leybausen also gests chat the many nerves at the base ofthe canine teeth allow eats to eel” forthe zap between the prey’s vertebrae before biting down. As the cat bites, ehe tooth inserts itself between the vvertobeae like a wedge, forcing themn part ana break {ng the spinal cord eee as ‘wrt Figne 2. Ding of retain mechan erie ins Claws While most car species are terrestrial, many have tes tained the ability to climbs aided by the sharp, curved, retractile claws that abso serve to hold prey. Among the Carnivora, only the cats use their fotepavs to re- strain prey with a clasping motion prior to the elie ery of a billing bite. Cats are well known for thes sharp cles, which remain fully retracted when they ate resting o# walking. Claw rettaction & passive. Ate tached tothe terminal hone of each toe, the claws ate hheld hack and off the ground by a retractor ligament. ‘Ta extend its claws, the cat contracts the dana sol ventral muscles of the toe; the ligament streehes, and the claw protrudes (fg. 2). Though the cheetah, ser- val, and slacheadel c2¢ are sometimes described 28 having noneetroctle clws, they actually have the sane claw retraction mechanists ne other cat species ‘The difference is that their claws protrae beyond the fur, and cheetah claws lack the sheaths that cover the claws of oer cats Body Size Felids vary enormously in size, sometimes even within the same species (Ags. 3 and 4). The sice disparicy among leopard, for example, once Ted taxonomists to conclude that forest leopands and savanna leopards ‘were diferent species. Similar se diferences ate also found amor Fumes, jaguar, stu tigen, all species with Heoal geographic distributions. Male jaunts Venezuela, which average 104 kilograms, are neatly twice as heavy as males in Belize, which average 56 Kilograms. Which species ithe smallest is still open to discussion, with the black-footed cat. kodked, and rustyspotted eat all candidates fo the tie ‘An examination of the numbers and sizes of fell species that occur in lange geographic regions shows that Southeast Asia has en species, South America hasten species, and Afri has eight (fas 5,6, and 7) “The greatest diversity occurs in the tropical zones the Teas in the temperate sone, Color Panerns ‘Most felis are blotched, spotted, or striped. Tigers are the only large caes marked with distinct transverse stripes: most of the other lage cats are marked wth rosettes. The cheetah’ coat pattern of erp, dark goes ‘s quite distintive. The uousual striped marking pat- ‘tem of the king cheetah is repeetedly due to a single recessive gene. In some smaller cat species the dark markings ate fused eo form hands, streaks, or stripes, al protegido por derechos «i qutoMAACINGS FASTA BSR gE Ee ees ee hall re nee eso cspoctally along the hack: and across the fimbs; in oth- crs the pattern is small dack sts, snd sil ther small cats are enly faintly spotted. Clouded leopards and ‘marble eats have blotched coats. The bon and puma fre more or less uniformly colored, although che imanes of male lions may he quite distinctive. The young ofboth lion and poma are spotted. The caracal ley has anpatterned fur except for dark spots above theeyes and shadowlike markings on the cheeks. The only other cat of uniform colo is the jaguarundi, which has a reddish moeph and a blackish morph Melanism (black pigmentation) appears to be com- oe frm nr 906 m7 ‘ae 5, Number of pucks al ts wi cats in Soubent Ast me Faye 6. Number of resin tse wld in Sth Ame _mom im some species (¢. serval, leopard, jaguar, ko kod, angle ext, Geoffroy eat, oncilla) and rare ot «x= tuemely Localized: in several others (c.g, bobcat, loved leopand, puma, cheetah. ‘White spors or bars are found on the hack of the fears in many species, although in almost as many species the ear spots are poorly defined or absent (©. wildcat, jungle eat, Chinese desert cat, sand eat, Dack-footed eat, African golden eat, Asiatic golden «at, bay cat, caracal, jaguarundi, manul, puma, rusty spotted cat, on). Their function is unknown, al- though one suggestion isthar the ear spots, along withMAACNGS TATA BSR gS oe 8 —— Levene the white on the underside ofthe til tp of some spe- ‘ies, serve as “follow me” signals to the young. which may be especially important in low-light conditions. ‘The prima function of coloration i concealment, ‘Cats that live in che desert tend to he light and sandy. colored. Forestliving species are darker and marked with spots or streaks that disrupe che outline of their bradies, concealing them in dappled light. Cats ving i mountainous teria often have light-colored or grayish coats. SENSES Eyes (Of al their senses, cats probably depend mest oi vie sion tobelp them capture prey Because cats hunt prey that i active both during the day and at night, their ves must e able to function ina range of onions ‘rom bright sunlight to almost complete darkness. Several adaptations allow a cateeyes to fametin c= ceafally in all ight comsitions Lange cyes with Lange pupils generally have good light-gathering abilities. Cts eyes ate extremely lage in relation to theie body size: the eve of a domestic cat, for example, is only slightly smaller than the eve ofa human. Like other imal «cat can regulate the arwaint of light dat eaters ie eye by dilating o eon traeting the pupil. On a sunny day, a cat's pupils con tract t0 a vertical shit or pinprick, while in dim light the pupils dilate toa large citcle of oval, sometimes appearing to take up the whole eye. oper a fa a ft 2 Frou 7. Number of pci ad stes Fld cin Aen. There are ro types of light-sensitive cells in the ‘retina of mammals: rods function n law levels of light and do not detect color cones are used in color vision. Good light-gathering ability 1s aswociated with an abundance of rods, and noe suspeisingly, rods predom- inate in cas eyes. However, cars also have a cone-rich patch in the center of the retina, 0 we know thar chey also have the ability to dicem color, at least under daylight conditions, Experiments have shown that cats can see green, blue, and possibly ted. Scientists speculate that not only do cats see fewer colors than sumans, but the colors they do see are much less satur rated than those we see ‘The ents sensitivity to light is abo heightened by a seructure known as the tapetum lucidum, a special re= flcctive layer beneath the retina, Caught in the bead- lights ofa car, a car’ eves shine back with a yellowish green light. This eyeshing, as i is called, comes from the rapstum. The minror-like rapetumm reflects the light hack through the retina to give the sensory cells a second chance to respond. It has been estimated that a eat light sensitivity fs nearly six times that of a human. Cars have the most highly developed binocular vision of all the carnivores: Their eyes ate set well forward and relatively high on the skull, allowing. them roaccurately judge distances while leaping foo ‘branch to beanch oe pouneing on pees Cats also have anextensive fel! of peripheral vision, which, strangely enough, may be partially responsible for their repus-IL-O2066 2/37/02 2158 ON Tage 18 ee tion for aloofness. Because its peripheral vision isso spond, a resting eat focuses itseyes infrequently. The re- sule i the cat's typical wide-eyed, staring énto-space Took that some people find so unsettling Whiskers Cats have specaled toulvscastive his, clled vib fisae oe whiskers om either se ofthe sie, around the eyes, Felow the chia, and on the ests. Whis- kers provide a sort of vision by touch, a tacole ehied eye tha serial to cats hunting at night. Sensitive cough to detect mince chase ae currents mew {ng around objects, whiskers permit even Blindfo ‘eats ro avoid obstacles without teaching them Stuer than other body hats and embeded more deeply in the ski, whisker are extremely sensitive to movement. They testin tiny nce of aid, esting hike svstraw sna soda bole. When anything brushes the big, the information i pased down wo aech suplyot serve endings that ie the sae ‘A hunting eat holds its whiskers out on either ste ofits fae ike a fan Just before pouring om ts pres: the catalufts its whiskers forward, extern them like at in font of is south, When the eat mikes cone tact with its res the whiskers tell i exactly which vey the animal is dosing in che final splirsecond. A cat eaying a fresly caught rouse wraps its whiskers :aound the prey sensitive tay ewitch that would in dicate tha the mouthful might squirm fee. Hearing Cats cat heat the 65~70 KH range, well abowe the fuuuars Haat of 15~20 KH. Cats do nor produce ul- teasonic cals, 19 heir ability 0 detect these highe frequency souads is probably rele wo hunting, Row dent ulteasouna communication occurs a the 20-50 KH: range, so small cats ane well equipped to detect the sounds of thle prex. Cats use their outer ears (pin- rae) as directional ampli o increase thee hearing sensitivity and pinpoint the location of sounds. The servily renowned for using ats huge dishlike eas to “sound hunt” fo small maramals, the extreme ex- ample ofthis abil ‘The auditory bullae of eats ako aid in deresting sounds. Ie manuals, the bla ae iat bony poe jections of the sll that protect the osiles of the tile eae. Each bulla has two ail cavities a outer chamber formed by the tympanic bone and an inner chamher formed by the entorympanic hone The balloe ae highly mosied (ited) in sone pe- «ics in connection with specialized modes of life, par- ticularly dhe detection of lew-frequeney sounds Offaction The allactory capabilites of eats ate no well under stood. Although there are occasional sefetences to cats tracking prey by scent they ane generally come cially. Craig Packer, wbo studied Liss in the Serer set, suggests tha where females live at high densitics inopen habeas and hunt large pre, they should form sroups, He argues that vultures and hyenas quickly find farge kill in open habitats, and rather chan shar ing such kills with these competicors, st would be- Ihoowve lions to share their kills with female relatives. ‘Coming at the problees from another direction, Timm Caro, who studied cheetahs on the Serengeti plains, ansues that most felids do not form groups because they typically live m areas where there # not enouch lange prey available to support a gu. Because domestic cats live in such a whe variety of habitats and under such a beoad spectrum of cee und stances, one might hope that they would peovide some insights into the costs and benefits af soeiairy. Though there are no examples of groups of domestic cats living on natural prey, there are several well: known studies of groups of domestic eas living around dock yard garbage dumpsters, fish offal dumps or fattn- yarch. The attiicial food sources at ches sites were ‘nel, predictable resource patches that supported sev ral females. At the dockyard site, cats lived in Fanily of 45 females. At the fxrnyaed ste, detailed observations showed that groups were highly seevctured in terms of individu relaconships, and several social clases could be distinguished, Some females gave birth sn communal dens, formed nursing coalitions, and jointly defended kitens against infan- rickdal makes, Females formed matrilineal subgroups, ‘consisted of amicable, tightly knit associations of several genceations of relatives descensdel fom the same mother. As with lionesses in pede, the matei- line « the basic social unic of geoup-living domestic Related tigress living in adjoining ranges on the preyerich foedplam of Chitwan tepresent another ex- pression of matrilineal organization, but one produced by different resource characteristics and dispersion. It fs not too diicul to imagine the cucumnstances under which thee neighboring sisters, mothers, and aunts might Gnd ve advantageous to get together «0 form a pide, I the habitae were open and supported abun- dant lage prey, and if there were serious competition rom vultuces, byenas, and other scavengers for ills, t ‘might create conditions that would favor the forma tion of groups There are many diferent rearons why cats form groups. Male lions and male cheetahs form coalitions to gain access to females. Female lions and domestic cats fotm groups to enhance reproductive success and territorial defense. But we sill do not completely n- derstand all the complesities ofthe isve—to beable 40 tease apart the factors that promote gtoup forma ‘ion, we will need information on other cat species ‘hat measures up tothe exquisitely detailed staies of lions and cheetahs in the Serenget. Lions in lodia's Gir Forest illustrate just how complex the isu ts fa the Gir, prey densities are low, lange prey is earee, and lions have no natural competitors for kills. Gir Forest lions eat many of the same species of ungulates that tigers eatin other parts of India Prey density, prey size, lack of comperitors, and habitar conditions would sug get that Gir lions ought to be solitary like ters, hut they are not. Like lions everywhere, Gir Hons live in prides of two to eleven females. There seemsto be ao ‘obvious reason for Gir lionesses to associate with ope another, even in small groups. Why do Gie lions sill behave like lions, even when they ive in “tiger” abi tat, feeding on “tiger” prey? Cleary, wee are sill miss ing a piece of the puss, al protegido por derechos «i quto22-0066 2/37/02 21S ON Page 16 Taxonomy of the Cat Family he systematics of felis has been the subject of ng and bitter debate among taxonomists. Since 1858, when Severtozaw divided the Felklac, of cat family, into five genera, there have been at east cen diferene clasitication schemes proposed, in which the rumbers of genera have ranged from two to as many as twenty-three, There i lest disagreement about the fouraber of fei species, with numbers ranging ftom thirtystx t thirty-nine. These various classification schemes have been based on an array of mornholost- cal behavioral, and genetic characteristics, including vocalizations, shape of the pupil, tooth number, tooth shape and size, cranial dimensions, foot and nose mor- phology, hybridization records, karyotype, ancl most recently DNA analysis 2543818 “Much of the confusion stems from the fact that al- most all the cats, with the exception of the cbeetab, make theit living in a similar fashion. They stalk, pounce, of rush and Kill theie prey using basically che same movements, teeth, and chs, and this means char eheir morphology is wery similar. Despite the fact that felis have the greatest range in body sie amen the Camivery, ringing from dhe sevenchundeed- pound tiger to the owonpound naty-spotted eat, they show litle variation in moepholegy: For practical reasons, cis volume follows the tax- ‘onomy tad out by Wozencraft? This taxonomic at- rangement is also wed by the Convention on Inter tonal Trade in Endangered Species of Wild Fauna and Flora (CITES), the World Conservation Monitoring Center (WCMC), and the Wild Cats Status Survey tnd Conservation Plan." MOLECULAR GENETICS The new and rapidly expanding Gell of molecular ge- netics promises to finally esolve the problem of fel phylogeny. Generic material fom diferent species.can bheanalyzed with a variety of molecular tests to deter- sine how cloncly eclated specie are to ome another Species that have heen separate fr along time show areater divergence ia theit DNA. However, because the cat family evolved fairly recently, and because many spsces separated fom ene another within a short period of ime, the dilerences are small and sometimes dificult t detect. The molecular genetic stead they are designed fora short, explosive sprint Aides by enlarged lungs, heart, aden glands, and ral puseges, cheetabs aceclerate rap, ataining ssspoel of 75 kilometers pee hour jst two seconds. Doing sprints thee respiatoey rate quickly climbs fiom 69 to 150 breaths per minut, and eat produce tion increases more than ifiyfokl.Cheetahsstore about 90 percent of the heat they produce while printing, by companion, Affican hunting dogs and domestic dogs store only about 20 percent ofthe heat they po sce during un, lescen likely that the length ofthe cheetah’ spent i determined by the amount of heat its kody can store before its temperature reaches adan- sous level —protably 40—41°C. In an experimental teal, cheetah refused to run when thee hay temper- ures reached 405°C. Since the speeds attained by cheetahs exceed those recorded forall ungulates, the outcome of any chase vwould seem ro be a foregone eanclusion. Homever, tele are adept 3 sigzaging, and to follow thes yeeycheetahs ave to make rapa ust high specs Dring these high-speed mancavers, cheetahs se theirtailsand feet to maintain balance and speed. The long til used ay a balancing ongan, and the claws are -exendad daring achase to increase traction-Cheetah claws ae shorter and straighter than those of other feds and thete are 90 cla sheath forthe claws to retract into, soeven when they are retracted the claws axe easy to $e, This fact account for the comeion ‘misperception that chectas have nonrtratile claws Ikea dogsVardaydescribesthe clausofhisret chee- tah m half-etrctable, “hut note stifly seta those of 2 dog." The pov are soull and compact, and the webbing herween the digits is much les than in other felis, allowing the toes to be spread widely. Further ‘more, the longitudinal ridges on the hand, pointed puso the feet function like eleatson a running shoe, eoviling eaction an rip ducing fast ears. DISTRIBUTION The cheetah was once wily distributed throughout Europe, Asia, Altica, and Noeth America. Four hun- Fir 10. Disha the cheetah dred years ag it was common in western and central Inia south ofthe Ganges, where its age closely mite toed that of is major prey the Hackbock. Ta the Mile Ease t was found throughout Palestine, Syria, Irog, Iran, Afghanistan, and the Sind." In Africa, the species ranged through all suitable habitats except desert aol topical foret, fom the Mediterranean 10, the Cape of Good Hore? Texas, the cheetah has almost disappeared from ‘Asia tis extinct in Indi, but seal populations may still exist in the former Soviet Union and inthe elon ‘where the intemational houndaries of lan, Afghani- stan, and Pakistan micet Cheetahs were reported to occur in the Khosh Yeilash Widife Refuge in ‘northeastern Iran," and there have been several re- cent sightings in tha general area 9° In Afric, the cheetahs distribution has been frxs- mented and greatly reduced, and major reductions have occured even since 1975. Though cheetahs in- hhabie a broad section of central, caster, and southern ‘Africa their major tronghols are in enstern and southem Africa, principally Kenya, Tanzania, and South Wese Africa/Namibia. An estimated 300500 survive inthe Sahara, most of theen en Mali, Niger, and (Chad. In East Africa the cheetahs distribution over- laps that of Thomsoris and Grant's gaelles and the serenuk, ond in southern Africa, that of the impala roaimocierd —h— erial pr gidoDOL-COO6S 2/37/02 28H ON Tage 6 ee ECOLOGY AND BEHAVIOR The cheetah i an animal of open country, aad in many parts of ts geographic range, this ca is steongly associated with grassy plains, open woodlands, and semidesert. A few cheetahs still survive among the sandy plains and rocky hills ofthe Sahara, where they spend the day among the rocks and beneath bushes, avoiding the intense eat of the sun. Cheetahs also have on the savannas and semiand rangelands of Kenya. They can hve equally well in areas wich more ‘cover; in the Masai Mara Reserve of Ease Africa they tise moderate cover for both husting and resting. Feeding Ecology Cheetahs hunt mainly by day, bue the hours vary slightly from area to area. In the Sahara, whete day ‘ime temperatures can reach 43°C, cheetahs do most ‘of their hunting at night and in the relatively cool hhoursafterdaybreak.* Io the Serengeti, cheetabshunt slightly later in the morning and cashier in the after- ‘noon than lions and hyenas, presumably as stratesy to avoud these other large carnivores In the Masai Mara region of southwestern Kenya, cheetahs are ac- tive mainly beeween 0730 and 1000 Ihours and again Fhenvcea 1630 and 1930 hours, they commonly Fest during the milday period." ‘Cheetahs have been seen hunting at night in the Secengetd and ir. Namibia,*™* but thete 9 general lack of information on theit nighttime activities. am ily groups generally spend the night resting in open grassy areas and are usally found in the same place in che morning where they were last seen i the eve> ‘ing, whereas males and javenile groups sometimes 1 ta aoe daring the igh. ‘ons, leopards, and hyenas all represent a chreatto ‘cheetahs. These larger and ore aggressive carnivores Kail adult cheecahs and cubs and steal cheetahs kills, 40 the timid, etining cheetah keeps its distance and maintains 4 carefel lookout for these other predators. Even when resting, the cheetah s alert to danger, and sentry" duty s shared among family members. If they are resting 00 top of a moun, cheetahs sean the sur- oundings simply by raising the hend. but an tall vee- tation they ston their haunches. In the Mar a total ‘of twelve minutes of every daylight hour was devoted to looking owt for danger. Not included in this este imate & tine devoted to looking out for other earnic vores while feeding, the situation in which most ago istic encounters eccue'* A hungry cheetah has ¢wo major ways of finding smc. Wotching frm a wet site, the eat may ace a potential hunting opporunty, When this happens, the cheetah unslly waits forthe animal to get chose ‘enough, then laanches into a chase, Cheetahs also find prey by walking slowly through the grassland look- ing o by canning the aten fmm alow Branch o¢ tere inte mound. To pt clone enough to attempt a chse, a cheetah may approach send of ples openly a et within 6D to 70 meters bette the eles ee, the cheetah may spring a them. But wherever there ivany cover at all the cheetah makes full we of 10 seal o choc 9 pore 4 us prey. Head lowered to shoulder level, Bay i a semtroach, i aleemately creeps or rns onward, then freee, maneuvering to within 30 meters or until it is close enough to the _gazelles to launch a short, fast chase. Unlike wild dogs or hyense which vend to selec thei prey during the hate portion ofthe hune, cheetahs seem to fc on ce animal before the chase begins and rarely atch targets ducing the chase * Tn areas with len cover, chestahs sometimes a roach prey ata ro, them within 200 meters or so ZIiesmotigh ped A ft appmach is alaooccssion- ally sein hans small Thomo cele, pas, or juvenile wildest. The fight response of prey arrears c be an important component of the chee- tans hunting technaque, as those prey that stand their ground of remain motionless are unlikely to be anced” Chectabs elect galls based on proximity and level of yiglance: more vigilant gael ate less vu rnerable to cheetahs because they fend fo ace mone rapidly when the cat hes its nal rsh More vis Tee gulls probably alo increse their weviver ship by detecting cheetahs ine the chase bens, hecame cheetahs abundoned neal thee-quates cf the hunts in which they were detected during the sealing hase ‘Wherever chert ate found, there are one o more ecient ples oe gselle-like antelope inthe 20-80 hilogre ange. These cats feed mainly on mediumatied unguliter and meno heirs weichs less than 40 kilograms. The list includes the impala, ‘Thomon’s and Grant's celles, singh, reek, waterbuck, kude, steenbok, daiker, warthog, and wil detest Spinghok are the dominant pcy in he Kala hari of South Africa” In the Serengeti, Thomson's ‘gazelles are the primary prey, accounting for 91 per cent of cheetah kills, although the frequency of hares in the diet increases when the migratory gazelles sre 6 M al protegido por derechos «i qutoDOL-COO6S 2/37/02 28H ON Tage aE scarcer abuent Te the bush and savanna worlds cf Kruger National Park, impale ae the dominant ey, constituting 68 percent of 2,532 kills, but chee- tah show a preference for eedbuck.” Impalas figure eominentlvinhedietofcheetahsfromthe Transvaal, although in many areas f Botswana and South West ‘Aica/ Navi, the bulk: of dhe dit i springbo. bn Naiebi National Path, impalas, Gene’ gazelle, and Thomson’ guellesaeconnt for alert 75 peteentofthe ‘cheetahs kills In almest all states t0 dae, fons a half-zown icles make up more than 50 percent cheetah Kill, even though young and adolecent Ungulaes constitute oply a small proportion of the rey population * Tn dhe Seteruet, prey preferences depend ta large ‘excar on which and how many cheetahs are doing the hanting. Male coalitions bunt the 80-kogran wildcheest, while single males waslly hunt Thoma. seller Subadule sibling groups, husting together without theie mother, chonse Thomson’ gazelle, ba solitary subsculs hune mote hares and infant Thom son's celles * Tn some hunts there appears ta be a degree of cor eperation, although whether its deliberate o singly fortitos 6 not cleat. Burney describes five hunt ing epiedes in which some degre of common effort seemes! apparent. fn ne ase a young male cheetah chased waterbuck calf towaed his tothe and a sib- Ting waiting in ambush. Another ime a family of che- tahsapreated to systematically search apatch of cover into which an impala fawa had run. A third form of ‘eonperative hunting involved typeof relay, in which sfecent faly merabers took upthe chase deperaing chow the prey turned ‘Atthe endef the chase, the cheetah trips or knocks the peey down and strangest. While ening slightly Ihindorbesde it, che cheetah reaches out and hooks the prey with the lige, strongly curved dewelam, ‘homing the prey of balance. Alemativel srike ‘oralpto the nin sith theforepane topples the query, and befote the animal can recover the cheetah has i by the throat. Cheetahs kill large prey by scans tion. Once a cheetah has its rey By the throat cst ally les dows andl maintains the stranglehold for fice tninutesorsa. The eats gripelosesoff che trachea, and