<<

Medicine and Surgery of Camelids

Third Edition

COPYRIGHTED MATERIAL

1 General Biology and Evolution

The domesticated camelids of the world have had a The classifi cation of SACs has been controversial. signifi cant impact on Old and cultures. One system classifi es the , , and Populations of and South American camelids within the and vicu ña as a single (SACs), also called New World camelids, declined in in the genus . Another system classifi es all the latter part of the nineteenth and early twentieth SACs within the genus Lama. Others classify the llama centuries. Governments neglected them as being an and alpaca as of L. guanicoe guanicoe . Recent unimportant component of the life of indigenous DNA studies have determined that and guana- people and tried to replace them with other domestic cos are appropriately species of Lama , and that the . Only in the last few decades have these alpaca and vicuñ a belong in the genus Vicugna .12 animals been recognized as a valuable resource and The family was previously designated as efforts made to research their unique physiology and an infraorder, , under the suborder Ruminan- adaptation to hostile environments. 1,4 tia, 18,23,26 but the most authoritative and current classifi - cation gives Tylopoda suborder status. 8,10,17,22,25,28,29,30 Some modern taxonomists have deleted the suborder Linnaeus placed the llama, alpaca, and Old World designation and use only family names. This book will camels in a single genus, Camelus, in 1758. Other tax- follow the classifi cation for camelids as listed in Table onomists proposed separate genus status for SACs in 1.1 . the early nineteenth century, but none of this work Collectively, llamas, , , and vicu ñ as was accepted by the International Commission on are called South American camelids or New World Zoological Nomenclature. The genus name Auchenia camelids (NWC), although the term “ auquenidae ” is was proposed by Illiger for SACs in 1811 and is fre- often found in older South American literature. Both quently seen in print even today in the South Ameri- camels and SACs are included in the term “ camelid. ” can literature. However, Auchenia had been applied Alpacas and llamas exist only as domestic species. earlier to a genus of insects and thus was not a valid Guanacos and vicu ñ as are wild species. It is generally name for any other . In 1827, Lesson published accepted that the alpaca shares some characteristics an acceptable paper classifying the New World Cam- with the vicuñ a, e.g., incisor teeth with an open pulp elidae in the genus Lama. In 1924, Miller assigned the cavity and continuous eruption into adulthood. vicu ñ a to a separate genus, Vicugna.13,14,17 The systematic classifi cation of Old World camels GENERAL BIOLOGY has never been controversial. The one - humped The karyotype of all camelids is 2n = 74. All SACs () is named Camelus dromedarius (Figure 1.1 ) have produced fertile hybrids. Dromedary and Bac- and the two- humped camel (Bactrian) C. bactrianus trian camels also produce fertile crosses. 9 In Israel a (Figure 1.2 ). Recently there has been acceptance of a cross between an alpaca male and a dromedary female third species of camel, the wild of Mon- by artifi cial insemination produced a stillborn full- golia, Camelus bactrianus ferus 12,14,21 (Figure 1.3 ). term fetus. In the United Arab Emirates scientists have succeeded in producing four living OWC and NWC Medicine and Surgery of Camelids, Third Edition by Murray E. Fowler offspring (two males, two females). It has yet to be © 2010 Blackwell Publishing, Ltd. determined if the hybrids are fertile. See Chapter 22 3 4 Chapter 1

Table 1.1. Classifi cation of camelids and other artiodactylids.

Class — Mammalia Order — Artiodactyla Suborder — Suiformes — Hippos, swine, Suborder — Tylopoda — Camelids Old World genus and species Camelus dromedarius — Dromedary camel C. bactrianus — Bactrian camel C. bactrianus ferris — New World genera and species Lama glama — Llama L. guanicoe — Guanaco Vicugna pacos — Alpaca V. vicugna — Vicuna Figure 1.1. Dromedary camel. V. vicugna mensalis (Peruvian) V. vicugna vicugna (Argentine) Suborder — Ruminatia — , sheep, , , giraffes, , antelopes,

for illustrations (Dr. J. Skidmore, personal communica- tion, Dubai, United Arab Emirates, April 2009). Despite size differences, the anatomy of all species of camelids is similar. SACs have similar behavioral patterns (Chapter 3 ).26 The camels completed Pleisto- cene evolution in a semidesert environment in south- ern , the Middle East, and North Africa and developed sophisticated adaptations for dealing with heat and dehydration (Chapter 9 ). The SACs became adapted to South American habitats, especially the high altitude lands of the . Figure 1.2. Bactrian camel. Camelids have a complex, three- compartmented stomach. Gastric digestion is similar to, but not analo- gous with, digestion. The two suborders separated from each other 65 million years ago when primordial species were simple stomached. Both groups used fi brous forage and developed similar foregut fermentation systems by parallel evolution (Chapter 13 ). Camelids regurgitate and rechew ingested forage, as do , but are more effi - cient than ruminants in extracting protein and energy from poor quality forages (Chapter 2 ). It is important to understand that camelids are not ruminants, pseudo- ruminants, or modifi ed ruminants. See Table 1.2 for other differences between camelids and ruminants.5 SACs establish communal dung piles, and in large herds there may be multiple dung piles, as animals in a herd segregate themselves into smaller units. Llamas and alpacas that have been imported into the following extended quarantine are more likely to urinate and defecate at random. Camels defecate at Figure 1.3. Wild Bactrian camel from the Gobi in random. Feces are pelleted in both groups and used . Used by permission from George B. Schaller, for fuel by people who share their habitat. Camelids Wildlife Conservation Society. have a unique reproductive cycle (Chapter 17 ). General Biology and Evolution 5

Table 1.2. Differences between South American camelids and ruminants.

South American camelids Ruminants

Evolution Evolutionary pathways diverged 40 million years ago Evolutionary pathways diverged 40 million years ago Blood Red blood cells elliptical and small (6.5 μ ); predominant RBCs round and larger (10 μ ); predominant WBC is white blood cell is neutrophil; leukocytes up to 22,000 lymphocyte; leukocytes up to 12,000 Foot Foot has toenails and soft pad. Second and third Foot has hooves and sole. Second and third phalanges are phalanges are horizontal. nearly vertical Digestive system Foregut fermenter, with regurgitation, rechewing and Same (parallel evolution) reswallowing Stomach — 3 compartments, resistant to bloat Stomach — 4 compartments, susceptible to bloat Dental formula — I 1/3, C 1/1, PM 1 – 2/1 – 2, Dental formula — I 0/3, C 0/1, PM 3/3, M 3/3 × 2 = 32 M 3/3 × 2 = 28 – 32 a Reproduction Induced ovulator Spontaneous ovulation No Estrous cycle Follicular wave cycle No follicular wave cycle Copulation in the prone position Copulation in standing position Placenta diffuse Placenta cotyledonary Epidermal membrane surrounding fetus No epidermal membrane on fetus Cartilaginous projection on tip of penis No cartilaginous projection on tip of penis Ejaculation prolonged Ejaculation short and intense Respiratory system Soft palate elongated; primarily a nasal breather Soft palate short; nasal or oral breather Urinary system Kidney smooth and elliptical Kidney smooth or lobed Suburethral diverticulum in female at external urethral No suburethral diverticulum orifi ce Dorsal urethral recess in male at junction of pelvic and Dorsal urethral recess in some species penile urethra Parasites Unique lice and coccidia; share some gastrointestinal Unique lice and coccidia; share GI nematodes nematodes with cattle, sheep, and goats Infectious diseases Minimally susceptible to tuberculosis; no known natural Highly susceptible to tuberculosis, bovine brucellosis, and bovine brucellosis; mild susceptibility to foot- and - foot - and - mouth disease mouth disease; rare clinical disease with other bovine and ovine viral diseases

a I = incisors, C = canines, PM = premolars, M = molars.

the Incas used them as pack animals.11 All guanaco SOUTH AMERICAN CAMELIDS subspecies share uniform coloration, with a dark The guanaco (Figure 1.4 ) has the broadest distribu- brown upper body, neck, and limbs; whitish fi ber on tion, both historically and currently, of the four SACs. the underside of the neck and belly; and a grayish to Four geographic subspecies of guanaco have been black face. described,6 ranging from sea level in Tierra del Fuego Vicu ñ a distribution is limited to the puna (Quechua at the southernmost tip of to 4,600 m for highland) life zone of the Andes (elevation 4,200 to in the Andes. The northernmost populations exist at 4,800 m). 4,6,26 The vicu ñ a (Figures 1.5 , 1.6 ) is the small- latitude 8 ° south in .3,11 Guanacos live in both est of the SACs and has the fi nest fi ber coat. It has a migratory and sedentary groups.4 Captive - born gua- cinnamon - colored coat, white underparts, a pale cin- nacos may be tamed and handled similarly to llamas; namon face, and a bib of white on the chest. There 6 Chapter 1

Figure 1.6. Argentine vicuñ a (short bib hair).

Figure 1.4. Guanaco.

Figure 1.7. Heavily fi bered huacaya alpaca.

Figure 1.5. Peruvian vicu ñ a (long bib hair). ance of Corriedale sheep . 6,16,19 The coat of suri alpacas consists of long fi bers with no crimp that hang are two geographic subspecies of vicuñ a: the Peruvian, down alongside the body in ringlets. A type of llama with long white on the bib, and the Argentinian, is now being bred that has suri fi ber characteristics. with shorter hair on the bib. The vicu ñ a was consid- Alpaca coloration varies from white to black with ered the property of the Inca kings, and only royalty intermediate shades and combinations. The alpaca is were allowed to wear garments made from the fi ber. the primary SAC fi ber producer of the Andean The two breeds of alpaca, huacaya (Figures 1.7 , 1.8 ) highland. and suri (Figure 1.9 ), have both become popular in the The llama is the largest of the SACs; however, United States. Alpacas are separated on the basis of among individuals there is marked variation in size, fi ber coat characteristics: 90% in Peru are of the huacaya overlapping sizes of other species. The llama has been breed. Huacaya fi ber is shorter than that of the suri a beast of burden since its . Two breeds breed and is crimped and spongy, giving it the appear- are recognized in Peru: the more woolly varieties are General Biology and Evolution 7

Figure 1.11. Llama.

Figure 1.8. Huacaya alpaca female.

called “ ch’aku ” in Quechua (Figure 1.10 ), and those with less fi ber on the neck and body are called “ q’ara ” (woolless) (Figure 1.11 ). Llamas were killed as sacrifi - cial offerings in the Inca culture, and the practice con- tinues to the present. 4 Both alpacas and llamas are slaughtered for in Andean countries. Lack of refrigeration necessitates immediate consumption, or the meat may be sun dried for storage. Dried meat is called “ charqui, ” from which the North American word “ jerky ” originated for a similar product. OLD WORLD CAMELIDS Dromedary camels are especially adapted to life in hot, arid areas of the world, notably the Middle East, North Africa, and , with a large population in Australia. have been used Figure 1.9. Suri alpaca. since ancient times for transport of people and goods, warfare, food, fi ber, and companionship. With the advent of modern transportation and the establish- ment of paved highways, camels have become some- what obsolete. However, country leaders have deemed it advisable to treasure some of the traditional uses of dromedaries and have fostered camel racing as a sport. The Bactrian camel is adapted to the cooler, arid of Mongolia, southern areas of the former Soviet Union, , and south . It is a beast of burden for carrying goods throughout its native lands and was the cargo carrier of goods along the Road from inner China to the Mediter- ranean. Bactrian camels have a heavy fi ber coat to cope with the cooler arid and provide the bulk of the fi ber used in the manufacture of Figure 1.10. Woolly - necked llama. garments. 8 Chapter 1

Approximate Ruminantia beginning of Epoch Epoch, millions Tylopoda Traguloidea Cervoidea Bovoidea of years before

Recent 0.01

Pleistocene 3

Pliocene 13

Miocene 25 Tayassuidae Camelidae Tragulidae Cervidae

Oligocene 37

Eocene 55

Figure 1.12. Suggested evolution of recent artiodactylids. Adapted from Romer 1966, Simpson 1980.

EVOLUTION

Camelid Camelid evolution began in over 82 million years ago in the early Eocene epoch. 32 Geologic and paleontologic time scales are estimates and subject to revision. Figure 1.12 provides a diagram of the rela- tionships of the various artiodactylid families since the Eocene epoch. Webb published a defi nitive work on the evolution of camelids based on paleon- tologic evidence.23 He suggested three major tribes, separated as early as the Eocene epoch. This family tree (Figure 1.13 ) is sketchy, with insuffi cient fossil records to trace lines accurately (postulates indicated Figure 1.13. Skeletons of various prehistoric camelids 8,23 by broken lines). Two tribes, Camelopini (Camel- removed from the La Brea tar pits in Los Angeles, ops) and (Camelus), evolved in North California. America only west of the Mississippi River. The tribe (Lama) was also found in Florida. 11,29 The Pleistocene epoch was characterized by a series of periods of extreme cold and glaciation in northern height of one of the early Pleistocene glacial periods, North America and Europe. 18 The last glacial retreat the sea level was lowered suffi ciently to expose a wide occurred about 10,000 years ago, marking the begin- land bridge. 2 Plant and animal species moved back ning of the recent epoch. It was during the Pleistocene and forth across this bridge; the camel line of Cameli- epoch that the camelidae fl ourished (Figure 1.14 ).7,9,19,22 dae migrated from North America into Asia to con- Many genera in the family Camelidae became extinct, tinue the evolutionary process, dying out in North for unknown reasons, before the recent epoch. America. Once in Asia, Camelus radiated through eastern Camel Europe (Rumania and southern ), the Middle Asia and Alaska are now separated by the 90- km East, and North Africa as far west as the Atlantic and (56- mile) - wide Bering Strait. However, during the as far south as Tanzania. It is likely that the dromedary General Biology and Evolution 9

Figure 1.14. Historic world distribution of camels.

evolved from the Bactrian camel, although the hump(s) Table 1.3. Approximate number of years that certain may be an acquired characteristic of domestication. animals have been domesticated. Wild camels had become virtually extinct in North 14,000 Llama and alpaca 6,000 – 7,000 Africa before historic times (3000 B.C.). Only C. - 12,000 – 15,000 5,500 nus ferus now exists in the wild state in one small area 11,500 Dromedary camel 5,000 in the Trans- Altai on the border of Mon- Sheep 11,000 Bactrian camel 4,500 golia and China, with a limited population of fewer Cattle 9,000 4,000 than 1,000 animals.12,14,22 9,000 Cat 3,000 – 4,000 South American Camelids The fi rst SACs migrated to South America at the beginning of the Pleistocene epoch (approximately DOMESTICATION three million years ago) when an open land connection between North and South America developed.17,24 The Camels Isthmus of Panama was formed by volcanic eruptions The approximate time of domestication of a few from the ocean fl oor, forming a series of islands that animals is listed in Table 1.3 . The precise time and ultimately became interconnected (the Caribbean land location of domestication of the Bactrian camel is bridge). The major earliest SAC genus appearing in unknown, but it is thought to have occurred sometime South America was (Tanupoloma), prior to 2500 B.C. on the border of and which radiated throughout the fl atter regions east of on the east side of the Caspian Sea. 14 The name the Andes. During the middle Pleistocene, the genera Bactrian is derived from a place name, Baktria, on the , Lama, and Vicugna developed from the Oxus River in northern . Strangely, this is long - limbed, fl atland - adapted Hemiauchenia. These not the place of origin of the domestic two- humped genera had shorter limbs, which more easily adapted camel, nor is the species even found in this area at them to the mountainous Andes. present. Various species of Palaeolama migrated back to Domestic Bactrian camels had spread north into North America. Fossils associated with North Ameri- southern Russia by 1700 to 1200 B.C. and were in can Hemiauchenia have been found along the Gulf western by the tenth century B.C. Bactrian Coast and Florida (Figure 1.15 ). 28 camels were used in China as early as 300 B.C. as the 10 Chapter 1

Figure 1.15. Historic and current distribution of NWC in South America.

original Silk Route camels but were later replaced alpaca herding economies spread beyond the limits of by crossbreeds of the dromedary and Bactrian the puna and became important in the economy of the camels.14 Andean people from sea level to high mountain Domestication of the dromedary occurred prior to elevations.17,27 3000 B.C. in the . The term “ drom- The was dependent upon the llama and edary ” is derived from dromos (Greek for road) and alpaca for food, fuel, clothing, transport of goods (the thus is only directly applicable to the riding or racing wheel was not introduced to South America before the dromedary camel. However, dromedary is the name conquest), and religious ceremonies. All SACs were used throughout the world for this species, which the property of the government, and production of existed in historic times only as a domesticated animal. domestic species was rigidly controlled. The fi ber from Dromedaries were fi rst associated with nomadic vicu ñ as was for royal usage only. The maximum Semitic cultures and never became important until the numbers and broadest distribution of SACs developed rise of the Arabian culture.14 under Inca rule. After the Spanish invasion of 1532 and Dromedaries were reintroduced into North Africa the introduction of European breeds of , in the third century B.C. More were brought into Egypt numbers and distribution of SACs declined. However, during the Roman period, after the third century A.D., llamas and alpacas survived because they are essential but became important domestic animals only with the to Andean culture. They are the most reliable source Moslem conquests of Egypt in the seventh to eleventh of food, fi ber, and fuel in the high, cool Andean envi- centuries A.D. A comparison of camel characteristics ronment. SAC ownership is the primary source of is found in Table 1.4 . wealth for indigenous people. 17,31 In 1970 the Peruvian government initiated an agrar- Llamas and Alpacas ian land reform. The large ranches, owned by wealthy, The cradle of llama domestication is the Andean absentee landlords, were confi scated and returned to puna (elevation 4,000 to 4,900 m), probably around the campesinos or pastoralists who had been working Lake Titicaca, at approximately 4000 to 5000 B.C. the land. Some groups of campesinos formed coopera- Alpaca domestication probably occurred elsewhere, tives, and the better ones hired skilled people to perhaps near Telarmachay. Alpaca- type incisors have manage the cooperatives, some with herds as large as been found in middens at Telarmachay, dated at 4000 40,000 alpacas. During that time, however, most of the to 3500 B.C.15,16,20,31 Once domesticated, llama and llamas and 80% of alpacas in Peru were under the General Biology and Evolution 11

Table 1.4. Characteristics of camels.

Characteristic Dromedary Bactrian Wild Bactrian Camelus dromedarius Camelus bactrianus C. bactrianus ferus

Breeds/types 50 different breeds recognized Geographical differences Single type Draft type: heavy body, stocky legs Riding type: Slim body, long legs Racing type: Similar to riding Weight kg/lb 300 – 650* /661 – 1432 450 – 700/992 – 1543 450 – 690/992 – 1521 Weight of newborn kg/lb 26 – 45/57 – 100 35 – 54/55 – 120 ? Height at shoulder cm/in 180 – 210/71 – 83 180 – 195/71 – 77 180 – 200/70.8 – 78.7 Body length cm/in 120 – 200/40 – 80 120 – 200/40 – 80 140 – 156/55 – 61 Shape 1 fi rm, upright hump 2 large humps, may be 2 small, conical humps fl opped over Color Cream to tan to dark brown Cream to tan to dark Cream to gray – brown brown Unique anatomy Male has a soft palate No dulaa, ears 15 cm No dulaa, prominent toenail, diverticulum (dulaa) which small foot and fl at sole for may protrude from mouth rapid gait to escape predation. Able to drink salty water, face narrow, ears 10 cm /hair Diameter 20 – 50 μ Diameter 10 – 40 μ , long Short fi ber staple, primary source for camel hair garments Special adaptations Adapted to heat, aridity, and Adapted to cooler, arid Adapted to the of sparse vegetation environments northern China and Mongolia. Running speed 21.6 – 40.3 kph (13.4 – 25 mph) 15 – 20 kph (9.3 – 12.4 mph) 40 kph (24.9 mph)

* Males, castrated as juveniles, may grow taller and larger than 1,134 kg (2,500 lb).

control of traditional pastoralists4,6,17 , who maintained llamas are estimated to be between 100,000 and 120,000, small to moderate herds (thirty to 1,000) grazing on and alpacas are thought to number more than 150,000. communal lands. The large cooperatives were dis- Sporadic importation of llamas and alpacas from banded in 1988, partly as a result of the activities of into the United States began in 1984 after the U.S. the Sendero Luminosa (Shining Path) terrorists and government periodically recognized Chile as free of because the new president and his advisors the foot - and - mouth disease. Importations have also origi- cooperatives were not functioning as they should. nated in and Peru, but these must meet special Currently llamas and alpacas are raised by traditional quarantine protocols for animals deriving from coun- pastoralists. tries where foot - and - mouth disease is endemic. A few In the nineteenth century SACs were exported to animals have been imported from England, New other countries from South America as zoo animals. Zealand, Canada, and Australia. Peru enacted legislation in 1843 prohibiting the export A general overview of the similarities and differ- of live alpacas. Approximately seventy years ago, all ences among the four South American species is listed of the Andean countries banded together to prevent in Table 1.5 . Sizes of SACs are listed in Table 1.6 . Addi- exploitation of SACs by other countries. No legal tional biological information is provided at the begin- exportations occurred from then until the 1980s, when ning of other chapters. the ban was lifted from alpacas and llamas.4 North American llamas have expanded from the USES OF CAMELIDS small population imported from South America prior Camels are an important part of the culture of the to 1930 and a few animals that had been imported from nomadic peoples of Asia and the Middle East, supply- other countries. Current numbers of North American ing food (meat and milk), fuel (the fecal pellets), fi ber 12

Table 1.5. Characteristics of South American camelids.

Characteristic Vicu ñ a Alpaca Guanaco Llama

Types/breeds/ Peruvian — Apron or bib on Huacaya — with light May be as many as 4 subspecies South American breeds: Heavy subspecies/ frontal chest crimp. Fleece at right angles to or races neck fi ber (chaku, lanuda, races body. tapada) Argentine — No chest bib Suri — Fiber lacks crimp. Fleece Short neck fi ber (ccara, pelada). No hangs in ringlets. North American breed standards. Alpaca and Llama Show Association show classifi cation: Light, medium, and heavy wool divisions.

Conformation Topline Straight with rounded rump Straight with rounded rump Straight or slightly rounded Straight from withers to tail from withers to tail Head and neck carriage Alert Vertical Vertical Vertical Vertical Resting Vertical 15 degrees from vertical Vertical Vertical Ears Short, curved on both borders Short, spear - shaped, curved on Medium length, curved on both Long, inner border straight or both borders. Huacaya — sharp borders curved inward (banana ears) tip. Suri — rounded tip. Face Short Short Medium Long Pastern Sloped Almost vertical Sloped Sloped

Fiber Diameter in 10 – 30 (13 – 14) Huacaya — 16 – 40 (22) 18 – 24 Undercoat — 16 – 40 (26) microns (avg.) Suri — 16 – 35 (23) Guard hair — 40 – 150 (70) Quality Finest of all SACs. Staple short. Excellent. Staple long. Inner coat is excellent; juvenile Variable; inner coat may make pelts are used for fur excellent garments. Staple may garments. Staple short. be long or short. Characteristic Vicu ñ a Alpaca Guanaco Llama Coverage Uniform over body, head, and Topknot present. Fiber extends No topknot. Uniform over body, Fiber usually doesn ’ t extend much upper legs. No topknot. below the knee and hock. head, and upper legs. below the knee or hock or onto the face Guard hair The bib and lower body fi ber are Highest - quality animals have no Concentrated on lower parts of Numerous guard hairs. guard hair. body and limbs

Color Color pattern similar to that of 22 solid colors recognized, Basic body color light to dark Numerous solid colors from white guanaco, but the basic body ranging from white to black. reddish brown (cinnamon) to black. Multicolors (pinto, color is a yellowish light Multicolored fl eeces also above, whitish hair below appaloosa) also seen. brown. The white in front of produced. (countershading). White the rear limbs may extend to extends up behind the foreleg the top of the back. Argentine and in front of the rear leg, subspecies has long white around the perineum, inside guard hairs on the chest, but of legs, and up the bottom of this is extended into a bib the neck. Front and outside of (8 – 14 in./20 – 35 cm) in the upper limbs are body Peruvian subspecies. color or grey. Head, face, and ears are dark grey to black, with the darkest color over top of the head and bridge of the nose.

Teeth Incisors long, narrow (sides Incisors elongated and continue Incisors broad, spatulate - shaped Incisors similar to those of guanaco parallel) and continue to grow to grow into adulthood. and do not continue to grow. throughout life. Enamel Enamel present only on labial Enamel surrounds the crown present only on labial side of side of incisors. of the incisors. the incisor teeth.

Feeding strategy Grazer/browser Prefers to graze succulent forage Uses broad range of habitat for Browser by preference but grazes in marshes and moist places grazing and browsing grasses, even dry, harsh species but also utilizes drier grasses and shrubs 13 14 Chapter 1

Table 1.6. Weights and sizes of South American camelids.

Characteristic Vicu ñ a Alpaca Guanaco Llama

Weight (lb/kg) Adults 99 –121/45 – 55 121 – 200/55 – 90 220 – 265/100 – 120 250 – 550/113 – 250 Birth weight 9 – 13/4 – 6 13 – 20/6 – 9 18 – 33/8 – 15 18 – 40/8 – 18 Height at withers Adults (in./cm) 34 – 38/86 – 96 30 – 38/76 – 96 43 – 45/110 – 115 40 – 47/102 – 119 Some imports 38 in.

Figure 1.16. Dromedary camel as a riding camel for tourists. Figure 1.18. Racing dromedary camels in the United Arab Emirates.

Camel racing has become a popular sport in , Saudi Arabia, Qatar, , and the United Arab Emirates (Figure 1.18 ). Adult female camels over seven years of age are generally used for major racing. Male camels are less easily managed. Younger camels may participate in short races but the major races are 4 to 10 km. The track is a huge oval and camels are quickly out of sight of the central grandstands. The race is viewed by remote television from vehicles that accom- pany the racers on a paved track on the inside of the race course. Figure 1.17. Dromedary camels as draft animals. The camels are ridden by lightweight jockeys (for- merly by children) sitting behind the hump. They are (clothing, ropes), leather, transportation (packing, now sometimes ridden by mechanical robot jockeys. riding; Figure 1.16 ), and racing. Special breeds of The start of major races is a raucous affair with as dromedaries were developed for riding and became many as fi fty camels at the starting gate. The camels important in the mobilization of military expeditions. move out at a gallop and may reach speeds up to Camels are also used for draft purposes, pulling plows 40.32 kph (25.05 mph) for the fi rst 2 km. Then they and wagons (Figure 1.17 ). Camels were used heavily settle down to a fast pace at an average speed of in Australia prior to the development of modern vehi- 32.4 kph (20.1 mph). cles for packing and pulling heavy and awkward items Camels may change gaits (pace to gallop, and vice into the Outback. The feral camels of Australia are versa) during the course of the race, much like the descendants of escapees. Camels also were used in the fl ying change of leads in a galloping horse. The camel United States as part of a military experiment in the gallop is slightly faster than the pace and is often used hot, desert environment of the Southwest. The Camel in the dash to the fi nish line. Wagering (betting) is not Corps was soon disbanded for various reasons. allowed in Moslem cultures. Prize money is awarded General Biology and Evolution 15 down to the tenth place. The top prizes in major races breeding and showing. The use of their excellent fi ber may be elite 4- wheel - drive sport utility vehicles. is currently a cottage industry, but has the potential Llamas have a long history of interaction with for commercial production. Andean people, supplying meat, leather, and fi ber (garments, blankets, rope, costales [burlap bags]); REFERENCES transporting goods; and serving as sacrifi cial animals.4 1. Bravo , P.W. 1991 . South American camelids and their In North America, llamas fi ll numerous niches, includ- crosses . Llamas 5 ( 8 ): 97 – 100 . ing, but not limited to, breeding, showing (Figure 2. Bravo , P.W. , and Fowler , M.E. 2001 . Order Artiodac- 1.19 ), parades, companion animals, packing (Figure tyla, family Camelidae (guanacos, vicunas ). In Biology, 1.20 ), driving, and fi ber (cottage industry). Well- Medicine and Surgery of South American Wild Animals. trained, quiet llamas are ideal for taking to schools and Ames, Iowa : Iowa State University Press , pp. 392 – 401 . convalescent hospitals and for interacting with emo- 3. Farb , P. 1964 . The Land and Wildlife of North America. tionally and physically disadvantaged children and Life Nature Library . New York : Time , p. 11 . adults. Another important niche is the use of llamas to 4. Flores Ochoa , J. , MacQuarrie , K. , and Porties , J. 1994 . guard sheep and goats from predation by . Gold of the Andes, Vol. I — The Andean Camelids and Alpacas are the premier fi ber - producing animals in Man ’ s Relationship with the Camelids, pp. 1 – 288. Vol. the Altiplano region of the Andes. They are also har- II— Europe, Modern Day Herders, Myths, Business, Barcelona, Spain : Francis O. Patthey Pub. pp. 1 – 352 . vested for meat in South America, and the leather is 5. Fowler , M.E. 2008 . Camelids are not ruminants . In used to make ropes. The pelts of crias are used to make Fowler , M.E. , and Miller , R.E. , eds. Zoo and Wild fi ne rugs and wall hangings; they are sewn together in Animal Medicine— Current Therapy 6th ed. Saint intricate patterns and designs. In North America, Louis : Elsevier , pp. 3375 – 385 . alpacas serve as companion animals and are used for 6. Franklin , W.L. 1982 . Biology, ecology, and relationship to man of South American camelids . In Mares , M.A. , and Genoways , H.H. , eds. Mammalian Biology in South America . Linesville, Pa. : Pymatuning Laboratory of Ecology, University of Pittsburgh Spec. Publ. 6 , pp. 457 – 89 . 7. Gahlot , T.K. , Tibary , A. , Wernery , U. , and Zhao , X.X. 2001 . Selected Bibliography on Camelids, 1991 – 2000 . Bikaner, India : Camelid Publishers . 8. Grubb , P. 1993 . Artiodactyla . In Wilson , D.E. , and Reeder , D.A. , eds. Species of the World , 2nd ed. Washington, D.C .: Smithsonian Institution Press , pp. 317 – 18 . 9. Harris , J.M. , and Jefferson , G.T. , eds. 1985 . Treasures of the Tar Pits . Los Angeles Natural History Museum . 10. Harrison , J.A. 1979 . Revision of the Camelidae (Artio- dactyla, Tylopoda) and description of the new genus Alforjas. Paleontol. Contrib., Pap. 95, University of Kansas, pp. 1 – 27 . Figure 1.19. Showing alpacas. 11. Kurten , B. , and Anderson , E. 1980 . Pleistocene of North America. New York : Columbia University Press . 12. Li , G. , Wang , N. , She , Q. , and Zhang , J. 2008 . Research on distribution, living environment and protection of wild Bactrian camel Camelus bactrianus ferus . J. Camel Pract. Res. 15 ( 2 ): 211 – 217 . 13. Marin , J.C. , Zapata , B. , Gonzalez , B.A. , Bonacic , C. , Wheeler , J.C. , and Casey , C. 2007 . Systematics, taxon- omy and domestication of alpacas and llamas: new chromosomal and molecular evidence . Revista Chilena de Hsoria Natural 80 ( 2 ): 121 – 140 . 14. Mason , I.L. 1984 . Camels . In Mason , I.L. , ed. Evolution of Domestic Animals . : Longman . 15. Miller , G.S. , Jr . 1924 . A second instance of the develop- ment of rodent - like incisors in an artiodactyl . Proc. U.S. Nat. Mus. 66 , Arctic. 8, No. 2545 . 16. Murphy , W.J. , Pezner , P.A. , and O’Brien , S.J. 2004 . Figure 1.20. Packing llamas on trek. Trends in Genetics 20 ( 12 ): 631 – 639 . 16 Chapter 1

17. Novoa , C. , and Wheeler , J.C. 1984 . Llama and alpaca . Allen , W.R. , Higgins , A.J. , Mayhew , I.G. , Snow , D.H. , and In Mason , I.L. , ed. Evolution of Domestic Animals . Wade , J.F. , eds 1992 . Proceedings of the First International London : Longman , 116 – 28 . Camel Conference, Dubai, UAE, Feb. 2– 6, 1992. New- 18. Nowak , R.M. , ed. 1991 . Walker ’ s Mammals of the market, England: R and W Publications. World , Vol. II , 5th ed. Baltimore : Johns Hopkins Uni- Fowler , M.E. 1986 . Camelids . In Fowler , M.E. , ed. Zoo and versity Press , pp. 1334 . Wild Animal Medicine , 2nd ed. Philadelphia , W.B. Saun- 19. Romer , A.S. 1966 . Vertebrate Paleontology , 3rd ed. ders , pp. 969 – 981 . Chicago : Univ. Chicago Press . Fowler , M.E. 1995 . Restraint of camelids . In Fowler , M.E. 20. Safl ey , M. 2001 . Alpacas, Synthesis of a Miracle . Hills- Restraint and Handling of Wild and Domestic Animals , boro, Oregon , Northwest Alpacas . 2nd ed. Ames , Iowa State University Press , pp. 279 – 287 . 21. Schaller , G. 1994 . Tracking the Gobi ’ s last wild bears Fowler , M.E. 2000 . Restraint and handling of camels . Journal and camels. Int. Wildl. 25 ( 1 ): 18 – 23 . of Camel Practice and Research. 7 ( 1 ): 77 – 90 . 22. Simpson , G.G. 1945 . The principles of classifi cation and Gahlot , T.K. 2000 . Selected Topics on Camelids . Bikaner, a classifi cation of mammals. Bull. Am. Mus. Nat. Hist. India: Sankhla Printers . 84 : 258 – 65 . Gahlot , T.K. 2004 . Selected Research on Camelid Physiology 23. Simpson , C.D. 1984 . Artiodactyls . In Anderson , S. , and and Nutrition. Bikaner, India : Camelid Publishers . Jones , S.K. Jr. , eds. Orders and Families of Recent Gauthier Pilters , H. , and Dagg , A.I. 1981 . The Camel — Its Mammals of the World. New York: John Wiley and Evolution, Ecology, Behavior and Relationship to Man . Sons , pp. 563 – 88 . Chicago : University of Chicago Press . 24. Simpson , C.D. 1980 . Splendid Isolation . New Haven, Higgins , A. , ed. 1986 . The Camel in Health and Disease , Conn. : Yale University . Press . London : Bailliere Tindal . 25. Springer , M.S. , Murphy , W.J., Eizirik , E. , and O’Brien , Hoffman , E. , and Fowler , M.E. 1995 . The Alpaca Book , S.J. 2003 . Placental mammal diversifi cation and the Herald, California : Clay Press . Cretaceous - Tertiary boundary . PNAS 100 ( 3 ): 1056 – Manefi eld , G.W. , and Tinson , A.H. 1996 . Camels — A Com- 1061 . pendium. Series C, No. 22, The T.G. Hungerford Vade 26. Strahler , A.N. 1973 . Introduction of Physical Geogra- Mecum Series for Domestic Animals . Sydney, Australia : phy , 3rd ed. New York : John Wiley and Sons , pp. University of Sydney Post Graduate Foundation in Veteri- 356 – 57 . nary Science. 27. Tomka , S.A. 1992 . Vicu ñ as and llamas — parallels in Schwartz , H.J. , and Dioli , M. , eds. 1992 . The One - humped behavioral ecology and implications for the domestica- Camel in Eastern Africa . Weikersheim , Germany : Verlag tion of Andean camelids. Hum. Ecol. 20 : 407 – 33 . Josef Margraf . 28. Vallenas , P.A. 1970 . (Commentary on the position of Smuts , M.M.S. , and Bezuidenhout , A.J. 1987 . Anatomy of the South American camelids in systematic classifi cation) Dromedary . Oxford, England : Clarendon Press . Comentarios sobre la posici ó n de los cam é lidos Tibary , A. , and Anouassi , A. 1997 . Theriogenology in Cam- sudamericanos en la sistem é tica . Bol. Extraordinario elidae. Abu Dhabi, United Arab Emirates, Veterinary 4 : 128 – 41 . Research Center. Mina, Abu Dhabi, U.A.E.: Abu Dhabi 29. Webb , S.D. 1965 . The osteology of . Bull. Los Printing and Publishing Company . Ang. Cty. Mus. Sci. 1 . Wernery , U. , and Kaaden , O. - R. 1995 . Infectious Diseases of 30. Webb , S.D. 1974 . Pleistocene Mammals of Florida . Camelids. Berlin : Blackwell Wissenschafts - Verlag . Gainesville : University Florida Press . Wilson , R.T. 1984 . The Camel . London : Longman Group 31. Wheeler , J.C. 1995 . Evolution and present situation of Ltd . the South American Camelidae. Biol. J. Linn. Soc. Yagil , R. 1982 . Camels and Production and 54 : 271 – 95 . Health, Paper 26. Rome, Italy: Food and Orga- 32. Zeuner , F.E. 1963 . A History of Domestic Animals . nization of the United Nations. London : Hutchinson . Yagil , R. 1985 . The Desert Camel, Comparative Animal Nutrition, Vol. 5 . Basel, Switzerland : Karger . SELECTED GENERAL MEDICINE REFERENCES ON CAMELS Al - Ani , F.K. 2004 . Camel Management and Diseases . Amman, Jordan : Dar Ammar Book Publisher .