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BiologicalJoumal ofthe Linnean Society (l995), 54: 27 1-295.

With 3 figures

Evolution and present situation of the South American Camelidae
JANE C. WHEELER

Facultad de Medicina Veterinaria, Universidad Nacional Mayor de San Marcos, Apartado 41-0068, Lima 41, Peru
Received 2 3 Afarch 1994; acreptedfor publication I6 Septmbrr I994

This paper provides a review of South American camelid evclution, classification and present status. Particular attention is paid to the debate roncerning origins of the domestic alpaca and llama and the contribution of researrh on faunal remains from Andean archaeological sites towards resolving this issue. Changes in incisor morphology during the domestication process suggest that the alpaca may be descended from the vicuiia, while a comparison of fibre production characteristirs in preconquest and extant llama and alpaca breeds indicates that extensive hybridization hetwern the two species is likely to have occurred since European contact. The potential role of hybridization in the formation of extant South American camelid populations has not been studied, and may be the root cause of taxonomic disputes. ADDITIONAL KEY WORDS:-guanaco hybridization conservation. breeds
~ ~~

vicuna

llama

~

alpaca

-

domestication

subspecies

CONTENTS Introduction . . . . . . . . The wild South American Camelidae . . The guanaco I h a guanicoe (Muller, 1776) The vicuiia C‘icugna vicugna (Molina, 1782) Origin ofthe domestic forms . . . . The domestic South American Camelidae . The llama Lama glama (Linnaeus. 1758) . The alpaca Lamapacos (Linnaeus, 1758) . Hybridization . . . . . . . . The future . . . . . . . . . References . . . . . . . . .

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“I 273 273 277 279 284 284 287 289 290 290

INTRODUC’IION

The South American camelids are classified together with the Old World camels in the order Artiodactyla, suborder Tylopoda, and family Camelidae, but subdivided into Lamini and Camelini at the tribe level. Two New Word genera, Lama and Vicugna, and one Old World genus, Camelus, are recognized. Ruminant digestion in the Tylopoda evolved independently of, and parallel to, ruminant digestion in the suborder Pecora (Bohlken, 1960). The Camelidae are distinLguishedby: absence of horns or antlers, presence of true canines separated from the premolars by a diastema in both the upper and lower jaws, position
0024-4066/95/003271+25 $08.00/0 27 1

0 1995 The Linnean Society of London

272
T,mi.i:

J. C : . LVHEELER

1. Systematic classification of the extant South American Camelidae
Artiodactyla O~.\.cn, 18-18 ‘l‘ylopoda Illigcr? 181 1 C;imelidae Gray, 182 I Lamini Wcth. 1965

Ordcr Suborder Family Tribe

I m i a Cuvirr, 1800
C c n m d r r J 1 .innaeus, 17 58 I m i n I:riscli, 1775 (rcjcctcd ICZN Opinion 2.58) 1,noiin ‘lk-dmiaini3 180-1 (unused senior synonym) durlrrnia Illiger, 181 1 inec ‘I’hinihcrg, 1789) T h n i n Rafinesque, 181-1 I m i n Lesson, 11127 I m i n Gray, 1872

of the vcrtcbral artery conflucnt to the neural canal in the cervical vertebrae, anatomy of the rear limbs which permits the animal to bend its legs beneath the body and rest on its stomach, and thc presence of a nail covered digital pad rather than a hoof. In 1758, Linnaeus described the two domestic New Jliorld camelids as Camelus glanzn “Camelus peruvianus Glama dictus” (llama) and Camelus pacoc “Camelus peruvianus laniger Pacos dictus” (alpaca), placing them together in a single genus with the Old World dromedary and bactrian camels, Canze1u.r dromednrius and Camelus bactrianus. Thc two remaining New World species, thc wild guanaco and vicufia, were subsequently designated Canielus guanicoe by Muller in 1776 and Camelus oicugna by Molina in 1782. As early as 1775, Frisch proposed that the four New World species be placed in the genus Lnnza, but this work is not accepted by the International Commission on Zoological Nomenclature (Hemming, 1985a) and authorship of Lama is credited to Cuvier, 1800 (Hemming, 1985b). According to the accepted nomenclature as listed in Cabrera (1961), the vicufia was assigned to a separate genus, Kcugna, in 1872 by Gray. None the less, the citation of Gray, 1872 is in error as this author described the vicufia as Llama vicugna (1872: 101). The earliest reference to Vicugna is by Miller, who in 1924 proposcd the generic separation based upon the vicufia’s unique hypselodont incisors. Although this designation has not been universally recognized, recent molecular analysis bascd on mtDNA sequence data suggests that classification into two gcnera could be appropriate (Stanley, Kadwell & Wheeler, 1994). Four species of New World camelids survive today: , . pacos (Linnaeus, 1758), guaiiaco the llama, L. glanza (Linnaeus, 1758), alpaca I L. guanicoe (Miiller, 1776) and vicufia I’icugna uicugna (Molina, 1782) (Table 1);

1979 Cnme1op.54 Tribe Camclini \Vchh. AYofoli. I934 Camelina Harrison. 1985) (Table 2). Harrison. 1939 ~ h l ? f F / U J LillIl~l~llS. Genraiu. the other descendant of Henziazichena.CAhlELID EVOLUTION ?'AHI. 1875 Hmriniichoim H.fimily 'l'ribc Subtribe Subtribe CLunclopina Harrison. after Harrison. Cabrera. Two genera. 1930. 1985 Artiodactyla Owen. Palaeolama (1. 1758. 18-18 Tylopoda Illigcr. 1979 .$zs Harrison.5 Lamina Harrison. Webb. 1932.i Subtribc hlatthcw and Cook.5 Myr) (Harrison. The tribe Lamini. Cabrera. I800 I7cugna hfiller3 1924 Order Suhordcr Family Su1.as (10-4. 181 I Camclidae Gr'iy. 1989) which probably . Fossil remains of L. 1867 Imna Cuvier. bactrzanus Linnaeus. 1821 C:anielinae Gray. Systematic classification of the fossil South American Camclidae. 1974) evolved from Pliauchenia approximately 10 Myr.1 Myr) remained in North America (Harrison. 18. 1979 Jlegomnelu Frick.E 273 2. 1112 1 Lamini \\'ebb. The first of these. while some species of Hmiauchenia migrated to South America during the Pliocene/Pleistocene transition approximately 3 Myr. I929 G@ntocnmu/rt. 1979. is no longer considered to be an ancestor of Lnmn and Vicugtza. 1758 and domestic and wild Bactrian camels C. 1965. 1979) and Hemiauchenia (10-0. originated in the Great Plains of western North America between 9 and 11 million years ago (Myr) (Harrison. represented by fossils of the genus Pliauchenia.. I979 .1 Myr) (Webb.eidy. Goin & Ortiz Jaureguizar. Alfoeas.5-0. Only Lnmn and Kcugna survived the end of the Pleistocene period some 10 000 years ago. 1758 together with the Old World domestic dromedaiy Catnelus drotnedarius Linnaeus. 1974). 1985). 1930. 1880 Poloeolonra P. 1965.1 Myr).i Barbour and Schulz.\~qa$opii.ilf. gzianiroe are found in Argentine Pleistocene deposits (L6pez Aranguren. 1776) The guanaco is the largest wild artiodactyl in South America. 196. 1979. Gcnrais and Ameghino. It is from the latter genus that Lnnza and Vicugna evolved in South America approximately 2 Myr (L6pez Aranguren.i I. 1972. 1905) Ti/nnoplopu~Barbour and Schulz. and its descendant Camelops (4. 1965 hlegatylopina Harrison. 1979 Pliciiirhrnin Copt-. THE \Z'ILD S O U I H AhIERlCAN CAhlELIDAF The guanaco h n i a guanicoe (Muller. Webb. Menegaz.5-0. 1932.

solid circles represent archaeological sites containing llama and alpaca remains. C . WHEELER Figure 1.274 J. 1986). Bolivia (Hoffstetter. 1980). date to some 2 Myr (Webb. solid lines mark the extent of their distribution prior to 1532. Tonni & Politis. stippled areas represent the cxtent of present ranges. They are also present at Tarija. Inrma pacos.qrza oicugna. but did not spread into the high Andean puna ecosystem before the establishment of modern climatic conditions between 12-9000 years ago (Hoffstetter. These numbers rapidly declined during . Neither fossil nor recent guanaco remains have been found in Ecuador or Colombia.. solid circles represent Pleistocene and early Holocene fossil localities. Raedeke (1979) has estimated the prehispanic guanaco population to have been 30 to 50 million based on carrying capacity of the territory which they occupied. 1A). C. dashed linrs mark the extent of present ranges. 1974) (Fig. 1986) in strata recently dated 97-73 000 years BP (before present) (MacFadden et al. Distribution of the South American Camelidae. A. Lnma guunicoc and B. Lama glama and D. 1985. guanacos were found along the Pacific shore and into the high Andes from approximately 8" South latitude to Tierra del Fuego. 1983). Kcu. as well as east into the Paraguayan Chaco and across the pampas to the Province of Buenos Aires (Torres. Prior to European contact.

1952. Sexual dimorphism is absent except for the presence of large canines in the male. 1991).g.g. 1952).. 1960. population in decline 1991 Andean population 92 882 wild. 1982a). compared to 100 cm for the small northern guanaco L. During the nineteenth century the impact of indiscriminate hunting and commercial sheep rearing reduced the guanaco population to 7 million. cacsilensis from Calipuy. 1982. from the tip of the nose to the base of the tail. 191 (Raedeke. . See Table 3 for comparison with the other South American camelids. population stcady Status the European conquest. Herre. parvs 109-1 19 94. Tierra del Fuego and Argentina south of 35"s latitude. 185 (Cabrera & Yepes. guanicoe from Patagonia and Tierra del Fuego (Cabrera & Yepes. The chest.g. 1973). guanicoe) to a lighter brown with ochre yellow tones in the northern variety (L. Rottman & Taber. MacDonagh. endangered species 3 776 793 domestic. and at present an estimated 602907 survive (Wheeler. General characteristics of the South American Camelidae Characteristics -__ Withers height in cm Adult weight in kg Gestation in days Birth weight in kg Condylo-basal length of skull in mm Feeding habits Brhaviour guanaco L. cacsilensis). Fibre diameter varies from 16.5 pm to 24 pm and contains from 5 to 20% hair (Verscheure & Garcia. Raedeke. 1979. guanicoe and 90-100 cm for L. varies from 167 (MacDonagh. 1974). 1949).g.g. Peru (Kostritsky & Vilchez. glania ~ __~ alpaca I.g. L. 1776). Reported body length. eyes and borders of the ears. guanicoe small 100 large 110 120 small 96 large 12(t130 small ? large 345-360 small ? large 8 15 small 244 large 280 k 25 browser-graser polygynous migratory and sedentary 602 907 wild. Four poorly defined subspecies of guanaco have been described: the first. the head grey to black with white around the lips.5+ 7. ziicupa 70 90 40 55 342-345 4-6 225 grazer polygynous sedentary territorial llama I.g.UTION 275 TABLE 3. huanacus (Molina.CAMELID EVO1. 1980.3 330 350 130-150 348-368 8 16 250 browsergazer polygynous territorial 6-7 22 1 grazer polygynous territorial 281 I612 domestic. Franklin. guanicoe varies from I20 to 130 kg (Raedeke. vulnerable species vicufia V. compared to 96 kg for L. cacsilensis (Herre. 1949).g. Live weight for adult L. 1974).. Lama guanicoe guanicoe (Muller. 1954a) for L. 1979. cacsilensis (Kostritsky & Vilchez. belly and internal portion of the legs are more or less pure white. the second. 1960). Withers height of adult animals varies from 110 to 120 cm for L. All guanacos exhibit similar pelage coloration varying from a dark reddish brown in the southern populations (L.104 59. Miller. is said to occur in Patagonia.g. Carpio & Solari. 1979) and 210 cm (Dennler de la Tour.

g. A different interpretation of guanaco subspeciation has been proposed by Franklin (1982). darker animals (L. north-western Paraguay (Torres. Cunazza. cncsilensis starting some 6000 years ago (Wing. the third.g. Moore. and little information on zoogeographic variations in guanaco morpholoLgy is available. 1992. 1991. Wheeler. and a second isolated population on the eastern side between 18" and 55"s latitude. domestic llama remains appear in archaeological sites located between 22" and 24"s latitude.g. guanicoe) contrast markedly with the smaller. 1992). Osteological remains from Andean archaeological sites document the origin of llama domestication from the guanaco at high clevation localities within the range of L. None the less. L. although tamed animals were sometimes kept (Benavente. It is unclear if these remains represcnt independent domestication events or the introduction of domestic animals from the central Andes. is found between 8" and 20"s latitude in Peru and northern Chile (Hoces. guanicoe. Recent survey data compiled by Torres (1992) shows population clusters which may correspond to the four proposed guanaco subspecies. At Pampa Galeras (Ayacucho. uogliz ?] in south-eastern Bolivia (Villalba. This subspecies is the best known of all the guanacos (Franklin. A predominately high altitude. 1944. a local high altitude form from southern Peru. Chile (Hesse. with a third &.'76 . small northern Subspecies.g. L. 1984) have postulated the possible existence of an undescribed coastal subspecies of cguanaco. cacsilensis. with one population restricted to the western slopc of the Andes between 8" and 41"s latitude. L. is restricted to Chile. There is no evidence that the Patagonian guanaco was ever domcsticated. 1985) and on the eastern slope of the Andes from 19" to 30" or 31"s latitude (Puig. 1992). while in the Province of Jujuy. 1982. The characteristics which set each subspecies apart arc not fully detailed in thesc early works.J C. 1992). found on the eastern slope of the Argentine Andes between approximately 21" and 32"s latitude.g. 1983. 1991a & b) they are dated to 4500 BP. extends from approximately 32"s latitude on the eastern slope of the Chilean and Argentine Andes.g. and it is clear that these larger. Argentina llamas may be present by 3400 BP (Yacobaccio & Madero. then a stronger case might be made for subspecific status with L. 1992). T o the south. cardensir Lonnberg. 1992. In the Salar de Atacama. huanncus ?] is found on the western slope of the Andes between 22" and 28"s latitude (Cunazza. a second population cluster [L. Further south still. some authors (Ponce del Prado & Otte. 1985). 1988. Within each of these areas. Guanaco numbers have continued to decline since European contact. 1986. If such a geographic separation. Dransart. the large austral subspecies. 1984. WHEELOR 1782). 1992).g. guanicoe the south-eastern. is real. 1989). Raedeke. cacsilensir being the north-western form and L. 1982. voglii Krumbiegel.g. cardensis) found at the northern bound ary . throughout Patagonia to 55"s in Tierra del Fuego (Puig. 1992). 1979 among others). Peru) these animals descend to the coast suggesting that human disturbance has displaced them from this part of their range in other areas. 1913. The most extensively studied populations are located at the southernmost limits of guanaco distribution. presumably produced by the salt pans of southern Bolivia and the crests of the Andean chain. Cunazza.g. and the fourth. lighter coloured specimens (Lg. the range of L. In .g.

it was only with the Pleistocene glacial retreat and establishment of the present Holocene climatic regime between 12-9000 BP that T4cugna moved into their present high elevation puna habitat (HoEstetter. The first. although a recent revision of some of these materials has led Mcnegaz et al. 1991). and in 1971 the government responded by declaring it an endangered species. 1930. 1985). Vicuiia remains have not been found in either palaeontological deposits (Hoffstetter. however. Separation of V.CAhlEI. In 1974.V. Dennler de la Tour (1 954a) called attention to the impending disappearance of the Patagonian guanaco if the hunting of yearling chulengos was not controlled and protected reserves established. None the less. vicugnn. however. 1986) or archaeological sites (Miller & Gill. mtDNA sequence data support a divergence of at least two million years between vicufia and guanaco (Stanley et a/. 1782) At present vicuiia distribution is limited to areas of extreme elevation between 9" 30' and 29"s latitude in the Andes (Fig. Palaeontological remains suggest. The vicuiia T4rugna uicugna (Molina. 1984. 1B). Recently the South American Camelid Specialist Group of the IUCN has urgently recommended increasing protection for the guanaco in general. This animal is not only highly endangered but also virtually unknown to science. inenralis was based on its smaller size relative to V. (1989) to conclude that the vicuiia evolved from the guanaco at the beginning of the Holocene. 1982). 1986) dated to between 97 and 73000 years ago (MacFadden et al. Bolivia include vicufia remains (Hoffstetter. Harrison. J'icugna vicugna uzcugnn (Molina.g. In 1957. Webb. By 1969. However. that the genus IScugna may have originated further east on the Argentine plains as early as two million years ago (Lopez Aranguren.. 1994). Cabrera. Argentina and Chile. 1782) is said to occur between 18" and 29"s latitude. 1932. including 250000 in Peru. V.g.II) EVOLU 1 ION 277 1954. and the recommendation should be taken seriously since archaeozoological and ethiiohistorical data demonstrate based on geographic distribution that L. nzensalis (Thomas.V. 1986. and two years later Jungius (1971) estimated a total of between 5000 and 10000 in Peru with another 2000 living in Bolivia. and th? relict northern L. 1985). No estimate of preconquest vicufia numbers is available.. 1976). In 1969. 1990) in Ecuador and Colombia. while the second. Grimwood found that the Peruvian guanaco population was on the edge of extinction. carsdensir is the anceytral form of the domestic llama (Wheeler. Pires-Ferreira & Kaulickc.V. Grimwood reported only 10000 in Peru. The present Andean population probably exceeds 93 000 thanks to rigorous protection programmes in the area (Wheeler. four in Chile and three in Peru. None the less. Koford calculated the total Andean vicufia population to he at most 400000. 1917) is reported between 9" 30' and 18"s latitude. 45 versus 57 mm for length of the molars and 70 versus 90 cm . 1974. these measures are insufficient and in Bolivia and Paraguay guanaco populations are not protected. Wheeler. the IUCN (International Union for the Conservation of Nature and Natural Resources) declared the guanaco a vulnerable species (Thornback & Jenkins. racrilensas in specific ('Torres. 1983) indicating that their range had expanded westward to the Andes by that date. At present the Lguanaco receives protection in 14 reserves in Argentina. Two subspecies of vicuiia have been described. 1986. and fossils from Tarija. 199 1).

52 pm (Carpio & Solari. Glade. The best studied vicuiia.V. the vicufia has made a remarkable recovery over the last 20 years. as well as covering the anterior portion of the rear legs.7 cm for males. 1982b) and the frequency of primary hair in the fleece is 2"/0 (Carpio & Solari. In 1957. mensalis.V. vicugna. Chilean.278 J. 1983). A third purported subspecies. mensalis. vicugna lacks the long chest hairs. WHEELER for withers height (Thomas. 1984. 1986. The head. V.V. 1991. Fleece fibre diameter is 12.52+ 1. sides and dorsal surface of the tail are a dark cinnamon colour. 1989). 1982). 1988. 1917).28 cm in adult animals and the long chest hairs reach 18 to 20 cm (Hofmann et al.. 1992. None the less. to vulnerable in 1972 (Thornback &Jenkins.4 cm for the males (Paucar et al.65 per mm2 (Carpio & Solari. Follicle density averages 78. In contrast with the continuing decline of guanaco numbers.5 to 6. sufficient phenotypic differences appear to exist between the northern (Peruvian. 1982). passing from endangered status in 1969.V. Koford. Recent survey data compiled by Torres (1992) shows vicufia populations spread throughout the high Andean puna ecosystem (at 3800 m above sea level and higher) from 9' 30' to 29"s latitude. 1950. the chest. This change is a direct result of conservation programmes. mensalis (1.2 kg and 36. Osteological remains from Andean archaeological sites point to the origin of alpaca domestication from vicufia at high elevation localities within the range of V. Villalba. Dennler de la Tour. 1992.V.2 cm in adult males (Carpio & Santana. interior surface of the legs and ventral surface of the tail. V. 1943. rising halfway up the sides to mid-rib height and all the way to the ilium crest. We have not encountered similar comparative statistics for V. belly.V. 1982a) and the average fleece fibre length is 3. Average coat length is 3.3 cm for females and 110.V. Grimwood. Total length measurements from the Paucar study of V. neck. 1982a). with average weights of 33. mensalis (Osgood. However. Bolivian) and southern (Argentine) vicufia populations to justify the existence of at least two geographic races.. 1954b. the northern V. 144 to 175 cm. Grimwood reported only 10000 in Peru. e h d n e Krumbiegel. Gilmore (1950) and Pearson (1951) also report greater total lengths.V. and two years later . Peru. The eyes and edges of the ears are outlined in white. No clearly defined geographic separation exists between the two proposed vicuiia subspecies and many authors ignore V. 1992. Hoces. is distinguished primarily by the long growth of hair on the chest. and heavier live weights. mensalis from Pampa Galeras (Paucar et al.. Gilmore. (1983) for 19 adult vicufias from the same local.5 cm for females and 90. C. 1944 has been described based on specimens found in German zoos. a recent study of 50 male and 50 female V. including 250000 in Peru. mensalis approximately 6000 years ago (Wheeler. Moore. 1984).2 kg respectively. 1969. 45-55 kg for V. 1992).V. In contrast. reports average withers height measurements of 86. and has a lighter beige pelage coloration with white covering a greater portion of the body. These figures contrast with the total length measurements of 137-181 cm reported by Hofmann et al. In 1969. with white covering the lower portion of the face. 1957).5 years old) from Pampa Galeras. back. without any obvious geographic clustering which might suggest two distinct subspecies (Cajal. Koford estimated the total Andean vicufia population to be 400000 at the most. 1984) are 96.

Peru (1 1" 11's latitude and 75" 52'W longitude). 1985). 1981). The present situation appears to have improved remarkably. Wheeler. Located 170 km north-east of Lima. 1981. Faunal materials from archaeological sites (Wing. and extended periods of drought occur. the Pampa Galeras population reached a crisis brought on by a prolonged drought. Moore. Archaeozoological data from one of these sites. while during later periods the frequency of camelid remains increased dramatically suggesting a shift to the utilization of domestic animals. cacsilensis) and vicufia (V. 1834).CAMELID EVOLUIION 279 Jungius (1971) calculated a total of between 5000 and 10000 in Peru with another 2000 living in Bolivia.6% to 27. 1991).8"C. 1988. 1976. Annual precipitation averages 500 to 1000 mm and is normally restricted to the months from November to March. Rabinovitch. were the primary prey of early human hunters. the total Andean population reached 92 882 (Wheeler. Argentina and Chile. the Peruvian population had risen to 62000 (Franklin. During this period pregnancy rates decreased from 85% to 58% and recruitment dropped from 76% to 27% (Otte & Hofmann. Hernandez & Cajal. 1986).28% growth rate was registered. This is due to their mobility during periods of drought and their ability to convert the dry ligneous puna grasses into a source of stored protein which can be utilized for human consumption. conservation efforts at the Pampa Galeras reserve began in 1967 with 1753 vicufias. Both guanaco (L. overgrazing and over-population.. 1984).6% (Sanchez. and in 1991. approximately equal numbers of camelids and deer were hunted. 1986. 1984). This high growth rate has subsequently been repeated when Chilean and Bolivian reserves were established (Rodriguez & Torres. at 4420 m above sea level. During the first years.g. Mean annual temperature is 4. ORIGIN OF THE DOMESTIC FORMS T o date the earliest evidence of camelid domestication comes from archaeological sites located between 4000 and 4900 m elevation. with an average daily variation of greater than 20" and frost occurs 330 nights of the year. 1989) indicate that during the earliest occupation of this zone 12 000 to 7500 years ago. although the timing is irregular and unpredictable. have produced the most extensive evidence concerning this shift to date (Wheeler. together with the huemul deer Hippocamelus antisensis (d'orbigny. 1984. but exact census data are lacking due to recent security problems in the area. Telarmachay Rockshelter. Palaeoclimatological data indicate that no significant climatic changes have . 1986. 1982). No agriculture is practised in the area today. In Peru. 15 years after the establishment of a protection programme. and a negative 1 1. 1982) while adult mortality increased from 5. mensalis) have inhabited this tundra environment for approximately 12 000 years and. Abortion rates rose dramatically and pregnancy was delayed from once a year to once every two years (Mtnard. and grazing ungulates represent the most reliable food resource. Wheeler et al. In 1982. an unexpected annual population increase of 21% was registered at Pampa Galeras (Sanchez.V. 1984. In 1978-79 however. Telarmachay is situated near the absolute upper limits of crop growth potential. in the puna ecosystem of the Peruvian Andes.

280 J C \YHEEI. up to 7Oo/o of each year's young may be lost before two months of age due.7% to 81. 1986). Around 6000 years ago. camelid remains increased from 64. In the Andes. together with changes in dental morphology.5 (Raedeke. the frequency of foetal and neonatal camelids increased markedly to 56. and continued to rise until it reached 73. Ramirez. outbreaks of enterotoxaemia are associated with unsanitary corralling practices during the wet season birth period. as well as by the presence of a critical number of captive or domestic animals.. to specialized hunting of guanaco and vicufia approximately 7200-6000 years ago. camelid remains gradually increased from 64. Instead. It has not been possible to determine if these shifts were associated with body size reduction as has been documented for other domestic ungulates because species specific characters for separating postcranial hones are lacking. while deer remains diminished from 34. During the preceramic period. 1991. 1986.6% of the faunal assemblage. in part.7"/0 to 88. 1986) revealed a 8200 year long occupational sequence and recovered more than one metric ton of animal bones fiom the preceramic levels.1°/0) of the bones coming from fetal or neonatal animals (Wheeler. These figures suggest either the development of specialized hunting of neonates. The epizootic nature of enterotoxaemia is to some extent controlled by climatic conditions that permit sporulation of the bacteria. Between 9000 and 6000 years ago. then to control of early domestic alpacas and llamas by 6000-5500 years ago.80/0. .Oo/o of all camelid remains in the deposits dated to 3800 years ago (Wheeler. but rather. 1979: 199) to 30% (Franklin. determination of early camelid domestication at Telarmachay is based upon an increase in the frequency of both camelid and neonatal camelid remains. 9000 to 1800 years ago.. This shift was not caused by decreased availability of deer in the zone. but closely correspond with mortality rates experienced by llama and alpaca breeders today. 1986). no human hunting) mortality rates of 4. 1978: 42) which have been recorded for the guanaco and vicufia. Similar epidemics are not known to occur in the wild camelids. They far exceed expected frequencies for both the foetal/neonatal age group and the natural (i. unpublished data).20/0 to 9. however. or the appearance of other mortality inducing factors in the environment. 1991).7% of the total faunal sample. Archaeozoological analysis of these materials produced evidence of a shift from generalized hunting of guanaco. Five seasons of excavation at Telarmachay Rockshelter (Lavallee et al. personal communication). At present.3'10 to 37. to failure of passive immune transfer (Garmendia et al. and finally.2% of the total (Wheeler. with-just over one third (35. to the establishment of a predominately herding economy beginning 5500 years ago (Wheeler. These figures are consistent with a hunting economy because between 35'/0 and 40'/0 of animals in contemporary guanaco and vicufia populations fall within this category (Franklin. Thus. 1986). vicufia and huemul deer 9000-7200 years ago. by a change in animal utilization patterns from generalixd to specialized hunting and cventual domestication of the camelids. 1987) with resulting mortality from Clostrr'dium fxrfringenf Type A eiiterotoxaemia and other pathogens (Leguia. 1978.ER taken place in this area over the last 10000 years (Van der Hammen & Noldus.e. the ever greater dependence upon camelids in the diet during this period suggests increasing specialization in guanaco and vicufia hunting. 1986). an economically unviable strategy.

Crisci & Vidal-Rioja (in press). the remains of permanent incisors with the same morphology as deciduous vicufia incisors appear in the Tclarmachay deposits (Wheeler. Vicufias have rootless hypselodont parallelsided permanent incisors with enamcl covering thc entire labial surface. and root-forming deciduous incisors with enamel covering thc upper lahial surface only (Miller.UTION 28 I Although it is not always possible to distinguish hctween the boncs of a terminal eleven and one half month foetal camelid and those of a neonate. however. go to waste. By 6000 BP. Antonius (1922). whereas those from the earlier levels were primarily foetal. 2B) (Thomas. Steinbacher (1953). . Weber. 1986). 1916. 1982. 1982. Other authors have concluded that both domestic camelids descend from the guanaco. 1904. but the presence of both large and small neonates suggests that this may have been the case. other researchers have come to different conclusions about their ancestry based on the study of living animals. In contrast to the data from Telarmachay and other Andean archaeological sites which indicate that the llama is descended from the guanaco and the alpaca from the vicufia (Fig. Meat produced by the often massive die-off of camelid neonates does not now. 1925. 1924. Guanacos have rooted deciduous and permanent spatulate incisors with an enamel covered crown (Miller. tooth wear studies indicate that the majority of Telarmachay specimens dating from 6000 to 3800 years ago were neonatal. unpublished data). an opinion subsequently supported by Ledger (1860). Peterson. 1891 . Darwin (1 868). Hilzheirner. 1928. Additional support for this interpretation comes from the study of bone distribution across the GO00 to 3800 year old living floors which indicates that newborn camclids were brought into the shelter whole and processed for consumption. The resultant pattern is very similar to that created by conteniporary traditional herders who utilize dead llama and alpaca neonates for food. L61inberg. The evidence from Telarmachay suggests an ancestral relationship which may explain the apparent retention of juvenile vicuiia dental traits in thc adult alpaca. Frisch attributed the origin of the llama to the _guanaco and the alpaca to the vicuiia. 1991). 199 1. 19 1 3. 191 3. Although contemporary alpacas with spatulate llama incisors have been reported by Kent (1982). Identification of the species which was brought under domestication at Telarmachay is based upon incisor morphology. with enamel covering only the upper labial surface (Wheeler. 1991. 1952). since these are indi~tin~guishable from guanaco incisors. Faige (1929). it is unclear if these are hybrids. Akimushkin (1 97 1) and Semorile. Krumbiegel (1941. and apparently did not then. Wheeler. Capurro & Silva (1 960). These permanent teeth match the dentition of many extant Peruvian alpacas in which both the deciduous and permanent incisors are root forming and parallel sided. 1953. 1982. Brehm. 2A). unpublished data). 1924). presumably taken ill utero through the h~intingof pregnant females (Wiceler. 1976. Cook.CAMELID E\ OI. In 1775. and permits the hypothesis that mortality induced by disease rather than by intentional butchery was the cause. 1952. and the vicufia was never domesticated (Fig. This shift from predominantly foetal to neonatal remains coincides with the significant increase in frequency of foetal/neonatal remains described above. Frechkop (1955). Herre. It cannot be determined if animals with llama type incisors also appeared in the 6000 BP deposits. Prior to domestication (9000 to 6000 BP) it is estimated that nine vicufia were huntcd for every guanaco based on incisor type and frequency.

1990) attributes llama ancestry to the guanaco. Finally. Bates. and that the guanaco and vicufia were never domesticated. glama. Model derivcd from study of extant animals in which both the llama and alpaca descend from the guanaco and the vicuiia was never domesticated.282 J.. Conclusions about llama and alpaca ancestry have. been based upon morphological changes produced by the domestication process. WHEELER A GUANACO VICUNA LLAMA ALPACA 1 I I B. 1983. vicugna fossils in Argentina. in large part. 1957) examined alterations in the mesotympanal area of the skull related to a decrease in llama and alpaca hearing acuity. and Piccinini et al. and reported an overall reduction in cranial capacity of both domestic species relative to the guanaco. 1963. GUANACO LLAMA z VICUNA ALPACA Figure 2 . 1961. but has deduced on the basis of shared morphological and behavioural traits that the alpaca originated from hybridization between the llama and vicuiia (Fig. Model derived from study of extant animals in which the llama is a domesticated form of the guanaco and the alpaca results from crossing the domestic llama with the wild vicufia. 1981/82. 1988. In the 1930s. 2C). L. 1975. based on the discovery of 2 Myr Plio-Pleistocene L. 1986. pacos. Herre & Rohrs. Jiirgens et al. 1976. 1982. Kruska. Rohrs. L. Hemmer (1975.. 1990). CUANACO VICUNA NOT DOMESTICATED ALPACA 1LLAMA - C. Kleinschmidt et al. B. During the 1950s. 1973. 1965.. C. Herre & Rohrs. 1953. A. Model derived from archaeozoological evidence in which the llama is domesticated from the guanaco and the alpaca from the vicuiia. 1952. Rohrs. This position is no longer considered a possible alternative. guanicoe and V. C. Herre and Rohrs (Herre. 1973. 1957. . Evolutionary models of llama and alpaca domestication. Herre & Thiede. Fallet. Zeuner. Pires-Ferreira. with subsequent hybridization between the two domestic forms producing the wan. Lopez Aranguren (1930) and Cabrera (1932) suggested that llama and alpaca evolved from presently extinct wild precursors.

suggesting that “the alpaca is a mixture of both lines. they found the vicufia cranium to be the smallest of all living Lamini. 1983). None the less. follicle structure. None the less. Russel & Stanley. concluded that this species was never brought under human control. 1992. the llama” (1990: 63). It has also been suggested that the vicuiia was never domesticated because it is more territorial than the guanaco (Franklin. Russel & Redden. 1984). which in turn were smaller than the guanaco suggesting that the latter is the only ancestral form. Although data concerning behaviour of the guanaco in this region are lacking. and concluded that the absence of transitional characteristics between vicufia and alpaca fleeces eliminates the former from consideration as an ancestral form. where vicufia and guanaco ranges overlap and llama and alpaca domestication occurred (Wheeler. 1982. Hemmer. These contradictory data on size reduction are almost certainly a product of sampling. coloration) in living camelids. based on the assumption that llamas have been selected exclusively for use as pack animals while alpacas have been bred for fibre production. 1952) and Steinbacher (1953) argue that the alpaca is the domestic vicufia based on unique shared behavioural traits which are said to differ from those observed in the guanaco and llama. concludes that while some alpaca behaviour patterns match those of the vicufia.CAMELID EVOLUTION 283 In contrast. 1974). 1983). 1976: 26). secondary/primary ratio. it is possible that the limited sedentary territorial organization observed in some Patagonian groups plays a more important role in these less extreme climatic conditions. alpaca. papers by Jerison (1971) and Hemmer (1990) report the ratio of alpaca brain size to body size to be smaller than in the vicuiia permitting a different conclusion about origins of the domestic forms. Wheeler. in part. This deduction is. Further to the north. based on the premise that domestic animals are smaller than their ancestors. on the other hand. llama and . a more benign climate and a constant food supply permit the characteristic sedentary social organization of the vicufia (Franklin. and. fibre length and diameter. this assumption is open to question because it is based upon study of guanacos located at the southernmost extreme of their range where seasonal migration in response to severe climatic changes is essential for survival (Franklin. None the less. Research on camelid behaviour has produced contradictory hypotheses concerning llama and alpaca origins. Krumbiegel (1944. as well as an extra fine fibre alpaca which is transitional between the vicufia and a second prehispanic fine fibre alpaca breed (Wheeler. Analysis of haemoglobin amino acid sequences in vicuiia. [produced] by crossbreeding of captured vicuiias with the only initially available domestic animal. 1982. Herre and Rohrs consider the llama and alpaca to be “races of the same domestic species bred for different purposes’’ (Herre. Fallet (1 96 1) found the llama to be an intermediate evolutionary stage between the wild guanaco and the specialized fibre-producing alpaca. new data on preconquest llama and alpaca breeds in Peru have revealed the prior existence of a fine fibre-producing llama. others are intermediate between those of vicufia and guanaco. submitted). Research on the relationship of brain size relative to body size by Kruska (1982) also found the vicufia to be smaller than alpaca and llama. Based on the study of pelage characteristics (skin thickness. as neither subspecific variation in the wild forms nor the possibility of hybridization between the domestic animals were considered in any of the studies.

. but information on molecular biology is limited. 1985. Hesse. In northwestern Argentina. Gentz & Yates. (1987) and Saluda-Gorgul. 1989). Although it is often assumed that the Lake Titicaca region was also a centre of llama domestication. Recent studies of the fibre from mummified ninth and tenth century llamas and alpacas suggests that post-conquest hybridization has modified the genetic makeup of living populations (Wheeler et al. 1994). 1986. After domestication in the Peruvian puna between 7000 and 6000 years ago (Wheeler. and it is thought that domestication may have occurred independently in both this region (ibid. WHEELER guanaco from Hannover Zoo. as have more recent data from ribosomal genes (Semorile et al. a single cranium of L. 1965. Germany. It is impossible to estimate the size of this preconquest llama population. 1990). 1986) indicate that all four species of the South American Camelidae have the same 211 = 74 karyotype. 1982). (1986). 1988).. 1991a). 1968. a fact which may well explain the diversity of conclusions about their ancestry. in press). Some 2400 years later they were being bred on the north coast of Peru (Shimada & Shimada. evidence of llama rearing has been recovered at sites located in the cloud forest on the eastern slope of the central Andes. the llama has adapted to a wide range of environments (Fig. (1988). 1C). Penedo et al. 1986). Hollander & Franklin. Like the guanaco. alpaca. earlier research on blood and muscle samples from llama. Vidal Rioja et al.. the llama was moved to the lower elevation interAndean valleys and into northern Chile where their remains have been found in archaeological sites dated to 3800 years ago (Wing. vicuiia. Larramendy et al. in press). 1977. Under Inca rule (1470-1532) llama distribution reached its furthermost expansion as pack trains accompanied the royal armies to southern Colombia and central Chile.. 'I'HE DOMESTIC SOUTH AMERICAN CAMELIDAE The llama Lama glama (Linnaeus. 1991. relevant data are lacking from early archaeological sites in Bolivia (Browman. Stahl. Reigadas. 1985) and in Ecuador (Wing.. but it clearly must have exceeded present numbers for early Spanish administrative documents record the virtual disappearance of these animals . and Piccinini et al. C. 1992). Jurgens et al. as well as in the dry Osmore drainage of south coastal Peru (Wheeler. 1984. Dransart. Other researchers utilizing immunological. Jaworski & Greger (1990) have examined satellite DNA. glama has been dated to 3400 years BP with stronger evidence for herding at 1450 BP (Yacobaccio & Madero. 1992).. 1960) indicated a llama-guanaco and alpaca-vicuiia subdivision. Taylor et al. 1984. (1990) to the conclusion that the vicufia was never domesticated.) and northern Chile (Hesse. guanaco and alpaca x vicufia hybrids at Santiago Zoo (Cappuro & Silva. 1986. However. Miller & Gill. Shortly thereafter. 1982."84 J. electrophoretic analysis and protein sequencing have found it impossible to draw conclusions about llama and alpaca ancestry (Miller. 1992. Cytogenetic studies (Capanna & Civitelli. 900-1000 BP. and research analysing the full mitochondria1 cytochrome b gene sequence in all six Camelidae has documented hybridization among the domestic South American camelids (Stanley et al. led Kleinschmidt et al. Wing. 1988. 1991. 1758) The llama is the largest of the domestic South American camelids and resembles its ancestor in almost all aspects of morphology and behaviour.

based on the measurement of up to 1600 fibres per animal. compared to 32.2 with a between sample standard deviation of 1. The reduction of both fibre diameter and variation in the fine fibre llama fleece was certainly produced by selective breeding for a single-coat through modification of the primary hair to resemble secondary undercoat fibre. The uniform coloration and fineness.CARIEIJD EVOLUTION LLAMA ALPACA 285 PRECONQUEST BREEDS COARSE FIBRE FINE FIBRE FINE FIBRE EXTRA FINE FIBRE Q'ARA CH'KU INTERMEDIATE WAk PACOWARI LLAMAWARI SURI HUACAYA INTERMEDIATE Figure 3. revealed the possible existence of both a fine fibre and a coarse fibre breed (Wheeler ef nl. In recent years the llama population has remained relatively stable. 3). 1977). 1992.. Average fleece diameter of the former was found to be 22.2) pin for the latter. within a century of contact (Flores Ochoa.. Research on the physical appearance of the El Yaral llamas. Because Andean civilization was nonliterate. submitted).7 (SDk4. an archaeological site in the Moquegua valley of southern Peru (Rice. as well as the absence of visible hairs in the El Yaral fine llama fleeces are ideally suited for textile production.. Associated with the pre-Inca Chiribaya culture. 1992.. Pi)st-doniestica1ioii llama and alpaca evolutioii sho\ring the relationship betwccn preronquest hrccds and cxtant phenotypes produccd b y a hrcalidown in controlled hrceding and hybridization. these animals had been sacrificed by a blow between the ears and immediately buried beneath house floors where thcy became naturally mummified due to the extreme aridity of the environment.8 pm. as well as analysis of skin and fibre samples taken at 11 different locations across the body. has provided a first view of preconquest breeds (Wheeler et al. and contrast markedly with the multicoloured double-coat of the coarse fibre breed. knowledge of pre-Spanish llama and alpaca herding practices must be reconstructed from archaeological remains. Wheeler et nl. totalling 3 776 793 in 199 1 (Wheeler. 1991). 1993). Wheeler et al. submitted) (Fig. The recent discovery of 900-1000 year old naturally desiccated llamas and alpacas at El Yaral. An additional evidence of specialized breeding is the accelerated fibre growth rate recorded for El Yaral fine llamas .

and emphasis was placed on breeding pure brown. The impact of such catastrophic mortality upon camelid genetic diversity and breeding practices has yet to be fully explored. 1965. 1991). the total llama population is estimated to be 3 776 793 (Wheeler. To the south. This region became a refuge for native livestock and herders. Small groups are found near Pasto. a memory assistance device made of knotted camelid fibre cords. these documents do provide details about Inca husbandry. 1982). Despite their European perspective. and it is known as qawra by Aymara speakers (Dransart. administrative documents record the disappearance of approximately 90% of the domestic camelids (Flores Ochoa. in press). Most llamas in Peru. and their descendants continue to inhabit the same marginal lands today. 1975. T o the south they extend to 27" in central Chile. These records describe the use of llamas as pack animals for the Inca army. WHEELER relative to contemporary animals (Wheeler et al. especially in Argentina. The name llama comes from Quechua (Flores Ochoa. Today. 1989). paid either in domestic camelids or in money obtained from their sale. resulted in depletion of the herds. the 'woolly' llama is more common and has a greater density of fibre on the body which extends forward between . as well as on quality fibre production for the state controlled textile industry (Murra.286 J. Colombia (1"N latitude) and Riobamba. Brotherston. 1975). but the most important production zone is located between 11" and 21"s latitude at elevations of 3800-5000 meters above sea level. goats. black and white animals for sacrifice to specific deities. Expansive state and shrine herds were managed by the llama camayoc.. The prolonged Spanish civil wars and heavy tribute levies. members of a hereditary caste of herding specialists. The growth curve and live weights of the llamas from El Yaral. Communally and individually owned herds also existed. at least three varieties of llama are recognized. 1982). Native Andean stockrearing was largely destroyed by the arrival of the Spanish. the former heartland of their distribution (Murra. as their grazing lands were usurped for the production of sheep. Ecuador (2"s latitude). our most detailed data on preconquest camelid breeding practices came from written documents of the colonial period. llamas and alpacas had practically disappeared even in the Lake Titicaca basin (Flora Ochoa. 1988). Prior to discovery of the El Yaral mummies. 1978. are very similar to those of contemporary llamas raised in the puna. submitted). Detailed data on size and colour of flocks were kept utilizing the quipu. By 1651. Coastal and highland valley herds were the first to disappear. but make no mention of fine fibre producing llamas. as well as 80% of the human population (Wachtel. cattle and pigs. 1991b). Bolivia and northern Chile are of the 'nonwoolly' phenotype characterized by sparse fibre growth on the body and the absence of fibre on the face and legs. and other Chiribaya culture sites of the same region. as well as their special interest in pack animals for use in transporting ore. Within little more than a century of the conquest in 1532. This may be due to the general failure of the early Spanish writers to distinguish between llamas and alpacas. 1977). Although specific llama breeds do not exist. In the puna this process was somewhat slower because both the Spanish and their livestock found the harsh climate and extreme elevation inhospitable. Introduced livestock diseases may also have played an important role in this process. C. and their age at death reflects controlled stockrearing with elimination of undesirable animals from the herd (Wheeler.

. the primary value of the llama presently lies in its use as a pack animal rather than as a fibre producer. but it is not clear to what extent selection is made for these characteristics today. In both areas llamas with intermediate phenotypes are also recognized. ranging from 32.0+ 15.7 pm (6) for ‘woolly’ cha’kus (Vidal. abundant fibre) and dolichomorphic (narrow. 17. Nevertheless. 1992. 1972.. 1985). Shimada.5f 17. The variability of present day llama fibre is related to an increase in hairs and general coarsening of the fleece. 3).). 1988) and Aymara (Dransart. sparse fibre).9 pm (6) pm (6) for intermediates. excavations in the central Peruvian puna have placed its origins between 7000 and 6000 years ago (Wheeler.9 pm (9) to 35. which probably began at the time of the Spanish conquest. submitted) (Fig. 30. Evidence of alpaca rearing at coastal sites in southern Peru dates from 900 to 1000 years ago (Wheeler et al. A different classification has been proposed by Cardozo (1954: 61) who divides llamas into brachymorphic (round. raising the possibility that more than three varieties of llama exist. 17. 1991b) speaking herders suggests that earlier management strategies may have been directed at producing animals with specific fibre types. Precolumbian alpaca remains have not been reported in the faunal materials from archaeological sites in Chile .1 f 12. 1988) and t’awrani in Aymara (Dransart. 1984. and it was from this region that the alpaca was subsequently moved to lower elevation interAndean valleys 3800 years ago (Wing. short profile. 1967). 1758) The alpaca is smaller than the llama and resembles the vicufia in many aspects of morphology and social organization. archaeological evidence is not presently available to evaluate this hypothesis (Browman. with wide variation in fibre diameter and a strong tendency to hairiness. Contemporary llamas lack the phenotypic uniformity associated with true breeds. Recent research on the fibre characteristics of Argentine llamas has identified the existence of seven distinct fibre types in the population (Frank & Wehbe. The woolly type is known as ch’aku in Quechua (Flores Ochoa. For this reason.5 pm (0) to 30. elongated profile. Fleece quality is uneven. 1986). The vast majority of llamas are held by traditional Andean pastoralists who utilize elaborate classification hierarchies based on colour. while the nonwoolly type is called q’ara in both languages (ibid.5 Ifr. The existence of these systems among both Quechua (Flores Ochoa. The alpaca Lama pacos (Linnaeus. 1D). 1991b).5) 18.CAMELID EVOLUTION 287 the ears and grows from inside the ears but is absent on the legs. Increased hairiness is produced by lack of controlled breeding. and Flores Ochoa (1988) indicates that the primary breeding criteria used by Quechua speaking herders in southern Peru is to divide llamas into ‘allin millmayuq’ and ‘mana allin millmayuq’ or fine and coarse fibre animals. Wheeler et al. and crossing between the two prespanish llama breeds from El Yaral could account for the entire range of fleece variation observed in todays animals (Fig.8 for coarse ‘nonwoolly’ q’aras. 1989). and 27. 1994). in press. Pelage coloration varies from white to black and brown passing through all intermediate shades with a tendency to spots and irregular colour patterns.6 pm (0) to 29. and llamas with wild guanaco coloration occur. Wheeler.5 Ifr. fibre and conformation characteristics to describe their animals. Although the Lake Titicaca basin of southern Peru and Bolivia has long been considered the focus of alpaca domestication.

where they have been recently reintroduced in Cajamarca.. The former have fleeces averaging 23. Spanish documents record their rapid decimation and displacement to remote. The Spanish conquest had a disastrous effect on both llama and alpaca populations. Representatives of two possible preconquest alpaca breeds have been found among the 1000 year old El Yaral mummies. known in the literature by their Quechua names as suri and huacaya or wakaya. with a greater tendency to uniform coloration than in the llama. and crosses of suri with suri produce some huacaya offspring. Fine fibre and extra fine fibre alpacas were distinguished. The suri is not known among the Aymara herders of Chile who refer to their huacayas simply as allpachu or alpacas (Dransart. Indced. suggesting that rigorous breeding selection for fine quality fibre was being practiced (Fig. It is impossible to estimate the number of preconquest alpacas. 1991b). alpaca distribution extends from approximately 8"s latitude. At prescnt.0 pm) (Wheeler et al. Crosses between huacaya and huacaya produce a certain percentage of suri offspring.6 (SD IfI 1. 1992. 1991) and 26. 1963) respectively. Although no artificial selection is made. 1977). are coarser.8 pm (Carpio. although they are found in limited numbers in these regions today. fibre diameter variation both within and across the fleece was remarkably low. contemporary Andean huacaya and suri fleeces average 31. while the latter fleeces average 17. as introduced stockrearing practices pushed the survivors into the marginal. an estimated 90% of all alpacas are huacayas (Novoa.9 pm). personal communication) or Ecuador (Miller & Gill. and in 1991 the total Andean alpaca population was estimated to be 2 81 1 612 (Wheeler. The fleece of both phenotypes varies from white to black and brown. Occasionally animals with intermediate wood characteristics are seen. In comparison to the preconquest El Yaral alpacas. may have a tendency to .0 pm (Von Bergen. Today. paqocha in Quechua (Flores Ochoa. Both groups had lustrous fibre ranging from wavy to crimped and dense to very dense. Argentina (Elkin. Wheeler et nl. Recent data indicates that over the last 25 years alpaca numbers have fallen significantly in Peru. crimped fibres which give it a spongy appearance similar to that of a Corriedale sheep.9 (SD IfI 1. 1990). from 3 290 000 in 1967 to 2 510 912 in 1986. passing through all intermediate shades. 75% of all alpacas. C.81f16.288 J. submitted). 1988) and allpachu in Aymara (Dransart. 1989). in the vicinity of Lake Poopo. The suri has long straight fibres. Hairs were visible in three of the four animals. 1982). Alpacas with wild vicuiia coloration occur. 199 1b). 3). with small populations located further to the south in northern Chile and north-western Argentina (Fig. to 20"s latitude. organized in waves which fall to each side of the body in much the same manner as a Lincoln sheep. extreme high elevation regions of the Andes (Flores Ochoa. 1991). but were not significantly coarser than the undercoat fibres. are held by traditional herders (Novoa. interAndean valleys and most of the puna. 1D). WHEELER (Dransart. based on physical appearance and average fibre diameter (1600 fibres measured per animal). 1982). Two alpaca phenotypes. 1991a & b.. Hesse. Bolivia.2k3. while the huacaya has shorter. Massive mortality accompanied the displacement of alpaca herds from the coast. are recognized but these do not breed true. extreme high elevation pastures where they are found today (Flores Ochoa. and these have been named chili by Cardozo (1954). 1989).

although it is slightly larger and less gracile than its wild progenitor. and considerable variation is reported in published statistics on fibre diameter. Crosses between the wild and domestic South American camelids produce fertile offspring. or alpaca x vicufia hybrid. 199 1b). The most probable cause of coarsening and hairiness in both huacayas and suris would be through hybridization with the coarse fibre llama breed. 3). such a process would not have affected only the alpaca gene pool. These hybrids are classified as llamawari or llama-like and paqowari or alpaca-like by Quechua speakers (Flores Ochoa. but this is said to rapidly increase in subsequent generations. Traditional herders recognize the existence of llama and alpaca crosses. 1968) and can. but comparison with preconquest animals suggests that it has been extensive and that breeds of fine fibre-producing llama and alpaca have likely disappeared in the process (Wheeler et al.3 to 17. 1992). a not improbable scenario amid the chaos and destruction of the conquest. 1977). Since this study included samples from the northern populations of both genera.. 1991b). First generation crosses are easily recognized. The extent to which contemporary llama and alpaca populations have been affected by this process has not been determined. llama and alpaca all possess the same 2n = 74 karyotype (Capanna & Civatelli.CAMELID EVOLUTION 289 hairiness.. the findings may possibly support Miller’s (1924) creation of the genus Kcugna. the region where they are sympatric in parts of their range.. produce fertile hybrids (Gray. The pacovicufia. Research on the fixation of phenotypic traits from generation to . has received considerable attention for its potential as a fine fibre producer. 1965. The origin of this degeneration almost certainly lies in the Spanish conquest. 1988) and wik‘ufia in Aymara (Dransart.3 pm for five first generation crosses. 3). Preliminary research on the cytochrome b gene sequence has found no evidence of hybridization between guanaco and vicufia (Stanley et a/.. but it is not always possible to identify hybrid animals based upon phenotype alone because it is likely that hybridization has been an ongoing process since the time of the Spanish conquest. Some coats containing up to 40% hair have been reported for both living varieties. (1990) report fibre diameters ranging from 13. Carpio et al. These are referred to by the generic terms wari in Quechua (Flores Ochoa. but a breakdown in controlled breeding between the fine and extra fine El Yard breeds would not alone account for the variation observed today. 1954). vicufia. as well as the generic term wakayu for any llama x alpaca offspring (Dransart. 1994). and are of uneven quality. under human influence. HYBRIDIZATION The guanaco. but do not normally occur in nature. however. Taylor et nl. The El Yaral mummies indicate the possibility that extensive crossbreeding between alpacas and llamas may have occurred since the Spanish conquest and has played a much more important role in the formation of today’s livestock than has been realized (Fig. Hybridization has been confirmed through DNA analyses (Stanley et al. The pacovicufia phenotype closely resembles that of the vicuiia. 1994) (Fig. Clearly. Aymara speaking herders use waritu and wayki for llama and alpaca phenotype hybrids.

Proceedings of the <oological Sorietv of London 134: 207-215. 1922. 1989. and generally exhibited the guanaco phenotype. if feral populations existed at that time. The uialking larder patterns of dumesticatiun. and they lived and reproduced without problem. although such information is essential for ensuring their future. No observations on changes in body size and fibre quality were recorded. 1971. The current consensus of opinion in the central Andean region is that no such populations exist today. although some had white blotches on the head and upper part of the neck. ed. . llama and alpaca populations have never been fully investigated. Stummesge. 1975. Reinbek Rowehlt. T H E FUTURE The present status of the South American camelids is the product of a largely unknown past. In the case of the wild camelids we need to be sure that we are protecting all genetic variants of each species. vicuiia.Sudanierika-Flora und Fauna. ’The uialking larder pattms of domestication. Brotherston G. London: Unwin Hyman Ltd. Bates M. Jena. Buenos Aires: Editorial Cartaga. there is an urgent need to identify relict populations of genetically pure precolumbian llama and alpaca breeds in order to insure both their preservation and the possibility of a return to high quality fine fibre production. the potential influence of genetic bottlenecks and/or hybridization in the formation of contemporary guanaco. and others are almost entirely white.290 J. The possibility that feral llamas and alpacas exist and might have crossed with wild camelids has not been fully explored. Argentina. MacDonagh (1940) has described a group of guanaco and llama hybrids living in a feral state in the Province of Cordoba. pastoralih and predation. . Remarks on the stomach and the systematic position of the Tylopoda. and much remains to be done before its potential as a fine fibre producer can be evaluated. Browman DL. Antonius 0. 1985. 1916. Even so. M@aciones. Xerez observed that domestic llamas were sometimes so numerous some escaped to the wild. C. Lcipzig. WHEELER generation of alpaca x vicuiia hybrids is lacking. Benavente MA. Bihiographilches Institut. although there is evidence to suggest that they may have played an important role. London: Unwin Hyman Ltd. and in 1555 Zarate recorded that once each year some llamas were released into the wild as an offering to the gods (Murra. Origins and development of Andean pastoralism: an overview of the past 6000 years. Andean Pastoralism and Inca ideology. These animals were the product of natural crosses. pastoralisnr and predation. Bohlken H. In light of the increased movement of both wild and domestic camelids throughout the Andes and the beginning of exportation in 1983. 1960.schichte der Haustiere. as well as llama and alpaca breeds. The most basic questions concerning genetic variation and the systematic classification of presumed guanaco and vicuiia subspecies. REFERENCES Akimushkin J. and not just increasing the numbers of potentially genetically impoverished subgroup/s. Die Suugetiere. Brehm A. 256-268. It is unclear. 1978). The behaviour of these feral hybrids was considered to be virtually identical with that of the guanaco. Boletin del h h e o Regional Araucania 2: 37-52. remain to be answered. In 1534. ed. however. In: Clutton-Rrock J. To name but two historic factors. 240-255. In: Clutton-Brock J. 1989. Reflexiones en torno a1 proceso de domesticacion de 10s camelidos en 10s valles del centro y sur de Chile.

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