Evolution and Present Situation of the South American Camelidae-Biological Journal of the Linnean...

8/13/2019 Evolution and Present Situation of the South American Camelidae-Biological Journal of the Linnean Society 1995

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BiologicalJoumal of th e Linnean Society (l995),5 : 27 1-295. With 3 figures

Evolution and present situation of the SouthAmerican CamelidaeJANE C. WHEELERFacultad de Medic ina Veterinaria, Universidad Nacional Mayo r de San Marcos,Apartado 41-0068 Lima 41, PeruReceived 2 3 farch 1994; acreptedfor publication I6 Septmbrr I 9 9 4

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

~ alpaca domestication subspecies

CONTENTSIntroduction . . . . . . . .The wild South American Camelidae . .

The guanaco I h a uanicoe (Muller, 1776)The vicuiia C‘icugna vicugna (Molina, 1782)

Origin of the domestic forms . . . .The domestic South American Camelidae

.

Hybridization . . . . . . . .The future . . . . . . . . .References . . . . . . . . .

The llama Lama glama (Linnaeus. 1758)The alpaca Lamapacos (Linnaeus, 1758)

. . . . “ I. . . . . . . . . . . 273. . . . . . . . . . . 273. . . . . . . . . . .. . . . . . . . . . .. . . . . . . . . . .. . . . . . . . . . .. . . . . . . . . . .. . . . . . . . . . .. . . . . . . . . . .. . . . . . . . . . .

277279284284287290290289

INTRODUC’IIONThe 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 Wordgenera, 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 aredistinLguished by: absence of horns or antlers, presence of true canines separatedfrom the premolars by a diastema in both the upper and lower jaws, position

27 10024-4066/95/003271+25 $08.00/0 995 Th e Linnean Society of London


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272 J . C:. LVHEELERT,m i.i: 1 . Systematic classification of the extant South AmericanCamelidaeO r d c rSuborderFamilyTribeI m i a Cuvirr,

Artiodactyla O~.\.cn, 8-18‘l‘ylopoda Illigcr? 181 1C;imelidae Gray, 182ILamini Wcth. 19651800c n m d r r J 1 innaeus, 1758I m i n I:riscli, 1775 (rcjcctcd ICZN Opinion 2.58)

1 no i in ‘lk-dmiaini3 180-1 ( unus ed senior synonym)durlrrnia Illiger, 181 1 inec ‘I’hinihcrg, 1789)T h n i n Rafinesque, 181-1I m i n Lesson, 11127I 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 beneaththe body and rest on its stomach, and thc presence of a nail covered digitalpad rather than a hoof.

In 1758, Linnaeus described the two domestic New Jliorld camelids asCamelus glanzn “Camelus peruvianus Glama dictus” (llama) and Camelus pacoc“Camelus peruvianus laniger Pacos dictus” (alpaca), placing them together ina single genus with the Old World dromedary and bactrian camels, Canze1u.rdromednrius and Camelus bactrianus. Thc two remaining New World species, thcwild guanaco and vicufia, were subsequently designated Canielus guanicoe byMuller 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 ZoologicalNomenclature (Hemming, 1985a) and authorship of Lama is credited to Cuvier,1800 (Hemming, 1985b). According to the accepted nomenclature as listed inCabrera (1961), the vicufia was assigned to a separate genus, Kcugna, in 1872by Gray. None the less, the citation of Gray, 1872 is in error as this authordescribed the vicufia as Llama vicugna (1872: 101). The earliest reference toVicugna is by Miller, who in 1924 proposcd the generic separation based uponthe vicufia’s unique hypselodont incisors. Although this designation has notbeen universally recognized, recent molecular analysis bascd on mtDNA sequencedata suggests that classification into two gcnera could be appropriate (Stanley,Kadwell Wheeler, 1994). Four species of New World camelids survive today:the llama, L. glanza (Linnaeus, 1758), alpaca I,. pacos (Linnaeus, 1758), guaiiacoL. guanicoe (Miiller, 1776) and vicufia I’icugna uicugna (Molina, 1782) (Table 1);


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CAhlELID EVOLUTION 273?'AHI.E 2. Systematic classification of the fossil South American Camclidae,

after Harrison, 1979, 1985OrderSuhordcrFamilySu1,fimily'l'ribcSubtribe

Subtribe

TribeSubtribc

Artiodactyla Owen, 18-18Tylopoda Illigcr, 181 ICamclidae Gr'iy. 1821C:anielinae Gray, 1112 1Lamini \\'ebb. 196.5Lamina Harrison, 1979

Pliciiirhrnin Copt-, 1875Hmriniichoim H. Gcnrais and Ameghino, 1880Poloeolonra P. Genraiu, 1867I m n a Cuvier, I800I7cugna hfiller3 1924

CLunclopina Harrison, I979.ilf;, zs Harrison, 1979Cnme1op.i I,eidy, 18.54

Camclini \Vchh. 1965hlegatylopina Harrison, 1979

.\~qa opii.i hlatthcw and Cook 1905)T i / n n o p l o p u ~Barbour and Schulz, I934

Camelina Harrison, 1979J l e g o m n e l u Frick, I929G@ntocnmu/rt.i Barbour and Schulz, 1939~ h l ? f F / U J LillIl~l~llS,758

together with the Old World domestic dromedaiy Catnelus drotnedarius Linnaeus,1758 and domestic and wild Bactrian camels C. bactrzanus Linnaeus, 1758.

The tribe Lamini, represented by fossils of the genus Pliauchenia, originatedin the Great Plains of western North America between 9 and 11 million yearsago (Myr) (Harrison, 1985) (Table 2). Two genera, AYofoli;as (10-4.5 Myr)(Harrison, 1979) and Hemiauchenia (10-0.1 Myr) (Webb, 1965, 1974) evolvedfrom Pliauchenia approximately 10 Myr. T he first of these, Alfoeas, and itsdescendant Camelops (4.5-0.1 Myr) remained in North America (Harrison, 1979;Webb, 1965, 1974), while some species of Hmiauchenia migrated to SouthAmerica during the Pliocene/Pleistocene transition approximately 3 Myr. It isfrom the latter genus that Lnnza and Vicugna evolved in South Americaapproximately 2 Myr (L6pez Aranguren, 1930; Cabrera, 1932; Webb, 1972;Harrison, 1985). Palaeolama (1.5-0.1 Myr), the other descendant of Henziazichena,is no longer considered to be an ancestor of Lnmn and Vicugtza. Only Lnmn andKcugna survived the end of the Pleistocene period some 10000 years ago.

T H E \Z'ILD SOUIH AhIERlCAN C A h l E L I D A FThe guanaco h n i a guanicoe (Muller, 1776)

The guanaco is the largest wild artiodactyl in South America. Fossil remainsof L. gzianiroe are found in Argentine Pleistocene deposits (L6pez Aranguren,1930; Cabrera, 1932; Menegaz, Goin Ortiz Jaureguizar, 1989) which probably


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CAMELID EVO1,UTION 275TABLE. General characteristics of the South American Camelidae

CharacteristicsWithers height in cm-__

Adult weight in kg

Gestation in days

Birth weight in kgCondylo-basal lengthof skull in mmFeeding habitsBrhaviour

1991 Andean populationStatus

guanacoL. guanicoe

vicufiaV. ziicupasmall 100large 110 120small 96large 12(t130small ?large 345-360small ?large 8 15small 244large 280 k25browser-graserpolygynousmigratory andsedentary602 907wild,vulnerablespecies

70 90

40 55

342-345

4-6

225

grazerpolygynoussedentaryterritorial92 882wild,endangeredspecies

llamaI,. glania109-1 19

_ _ ~

130-150

348-368

8 16250

browsergazerpolygynousterritorial3 776 793domestic,populationstcady

alpacaI,. parvs94- 10459.5+ 7.3330 350

6-7

22 1

grazerpolygynousterritorial281 I612domestic,population indecline

the European conquest. During the nineteenth century the impact ofindiscriminate hunting and commercial sheep rearing reduced the guanacopopulation to 7 million, and at present an estimated 602907 survive (Wheeler,

All guanacos exhibit similar pelage coloration varying from a dark reddishbrown in the southern populations (L.g. guanicoe) to a lighter brown with ochreyellow tones in the northern variety (L.g. cacsilensis). The chest, belly andinternal portion of the legs are more or less pure white, the head grey toblack with white around the lips, eyes and borders of the ears. Fibre diametervaries from 16.5 pm to 24 pm and contains from 5 to 20% hair (Verscheure

Garcia, 1980; Carpio Solari, 1982a). Sexual dimorphism is absent exceptfor the presence of large canines in the male. Withers height of adult animalsvaries from 110 to 120 cm for L.g. guanicoe from Patagonia and Tierra delFuego (Cabrera Yepes, 1960; Franklin, 1982; Herre, 1952; Raedeke, 1979;MacDonagh, 1949), compared to 100 cm for the small northern guanaco L.g.cacsilensis (Herre, 1952). Reported body length, from the tip of the nose to thebase of the tail, varies from 167 (MacDonagh, 1949), 185 (Cabrera Yepes,1960), 191 (Raedeke, 1979) and 210 cm (Dennler de la Tour, 1954a) for L.g.guanicoe and 90-100 cm for L.g. cacsilensis from Calipuy, Peru (KostritskyVilchez, 1974). Live weight for adult L.g. guanicoe varies from I20 to 130 kg(Raedeke, 1979; Miller, Rottman Taber, 1973), compared to 96 kg for L.g.cacsilensis (Kostritsky Vilchez, 1974). See Table 3 for comparison with theother South American camelids.

Four poorly defined subspecies of guanaco have been described: the first,Lama guanicoe guanicoe (Muller, 1776), is said to occur in Patagonia, Tierra delFuego and Argentina south of 35 s latitude; the second, L.g. huanacus (Molina,

1991).


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'76 ,J C. WHEELOR1782) , is restricted to Chile; the third, L.g. cardensir Lonnberg, 1913, a localhigh altitude form from southern Peru; and the fourth, L.g. voglii Krumbiegel,1944, found on the eastern slope of the Argentine Andes between approximately21 and 32 s latitude. The characteristics which set each subspecies apart arcnot fully detailed in thesc early works, and little information on zoogeographicvariations in guanaco morpholoLgy is available. The most extensively studiedpopulations are located at the southernmost limits of guanaco distribution, andit is clear that these larger, darker animals (L.g. guanicoe) contrast markedly withthe smaller, lighter coloured specimens (Lg. cardensis) found at the northernbounda y .Recent survey data compiled by Torres (1992) shows population clusterswhich may correspond to the four proposed guanaco subspecies. A predominatelyhigh altitude, small northern Subspecies, L.g. cacsilensis, is found between 8 and20 s latitude in Peru and northern Chile (Hoces, 1992; Cunazza, 1992). AtPampa Galeras (Ayacucho, Peru) these animals descend to the coast suggestingthat human disturbance has displaced them from this part of their range inother areas, None the less, some authors (Ponce del Prado Otte, 1984) havepostulated the possible existence of an undescribed coastal subspecies of cguanaco.Osteological remains from Andean archaeological sites document the origin ofllama domestication from the guanaco at high clevation localities within therange of L.g. cncsilensis starting some 6000 years ago (Wing, 1986; Wheeler,1984, 1991; Moore, 1988, 1989).

To the south, a second population cluster [L.g. huanncus ?] is found on thewestern slope of the Andes between 22 and 28 s latitude (Cunazza, 1992),with a third &.g. uogliz ?] in south-eastern Bolivia (Villalba, 1992), north-westernParaguay (Torres, 1985) and on the eastern slope of the Andes from 19 to30 or 31 s latitude (Puig, 1992). Within each of these areas, domestic llamaremains appear in archaeological sites located between 22 and 24 s latitude.In the Salar de Atacama, Chile (Hesse, 1982; Dransart, 1991a b) they aredated to 4500 BP, while in the Province of Jujuy, Argentina llamas may bepresent by 3400 BP (Yacobaccio Madero, 1992). It is unclear if these remainsreprescnt independent domestication events or the introduction of domesticanimals from the central Andes.

Further south still, the range of L.g. guanicoe, the large austral subspecies,extends from approximately 32 s latitude on the eastern slope of the Chileanand Argentine Andes, throughout Patagonia to 55 s in Tierra del Fuego (Puig,1992; Cunazza, 1992). This subspecies is the best known of all the guanacos(Franklin, 1982, 1983; Raedeke, 1979 among others). There is no evidence thatthe Patagonian guanaco was ever domcsticated, although tamed animals weresometimes kept (Benavente, 1985).A different interpretation of guanaco subspeciation has been proposed byFranklin (1982), with one population restricted to the western slopc of theAndes between 8 and 41 s latitude, and a second isolated population on theeastern side between 18 and 55 s latitude. If such a geographic separation,presumably produced by the salt pans of southern Bolivia and the crests ofthe Andean chain, is real, then a stronger case might be made for subspecificstatus with L.g. cacsilensir being the north-western form and L.g. guanicoe thesouth-eastern.Guanaco numbers have continued to decline since European contact. In


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CAhlEI,II) EVOLU 1 I O N 2771954, Dennler de la Tour (1954a) called attention to the impending disappearanceof the Patagonian guanaco if the hunting of yearling chulengos was notcontrolled and protected reserves established. In 1969, Grimwood found thatthe Peruvian guanaco population was on the edge of extinction, and in 1971the government responded by declaring i t an endangered species. In 1974, theIUCN (International Union for the Conservation of Nature and NaturalResources) declared the guanaco a vulnerable species (Thornback Jenkins,1982). At present the Lguanaco receives protection in 14 reserves in Argentina,four in Chile and three in Peru. None the less, these measures are insufficientand in Bolivia and Paraguay guanaco populations are not protected. Recentlythe South American Camelid Specialist Group of the IUCN has urgentlyrecommended increasing protection for the guanaco in general, and th? relictnorthern L.g. racrilensas in specific ('Torres, 1985). This animal is not only highlyendangered but also virtually unknown to science, and the recommendationshould be taken seriously since archaeozoological and ethiiohistorical datademonstrate based on geographic distribution that L.g. carsdensir is the anceytralform of the domestic llama (Wheeler, 1984, 1986, 1991).

The vicuiia T4rugna uicugna (Molina, 1782)At present vicuiia distribution is limited to areas of extreme elevation between

9 30' and 29 s latitude in the Andes (Fig. 1B). Palaeontological remainssuggest, however, that the genus IScugna may have originated further east onthe Argentine plains as early as two million years ago (Lopez Aranguren, 1930;Cabrera, 1932; Webb, 1974; Harrison, 1985), although a recent revision ofsome of these materials has led Mcnegaz et al. (1989) to conclude that thevicuiia evolved from the guanaco at the beginning of the Holocene. None theless, mtDNA sequence data support a divergence of at least two million yearsbetween vicufia and guanaco (Stanley et a/. 1994), and fossils from Tarija,Bolivia include vicufia remains (Hoffstetter, 1986) dated to between 97 and73000 years ago (MacFadden et al., 1983) indicating that their range hadexpanded westward to the Andes by that date. However, it was only with thePleistocene glacial retreat and establishment of the present Holocene climaticregime between 12-9000 BP that T4cugna moved into their present high elevationpuna habitat (HoEstetter, 1986; Wheeler, Pires-Ferreira Kaulickc, 1976).Vicuiia remains have not been found in either palaeontological deposits(Hoffstetter, 1986) or archaeological sites (Miller Gill, 1990) in Ecuador andColombia. No estimate of preconquest vicufia numbers is available. In 1957,however, Koford calculated the total Andean vicufia population to he at most400000, including 250000 in Peru. By 1969, Grimwood reported only 10000in Peru, and two years later Jungius (1971) estimated a total of between 5000and 10000 in Peru with another 2000 living in Bolivia, Argentina and Chile.The present Andean population probably exceeds 93 000 thanks to rigorousprotection programmes in the area (Wheeler, 199 1).

Two subspecies of vicuiia have been described. The first, J'icugna vicugnauzcugnn (Molina, 1782) is said to occur between 18 and 29 s latitude, whilethe second, V.V. nzensalis (Thomas, 1917) is reported between 9 30' and 18 slatitude. Separation of V.V. inenralis was based on its smaller size relative toV.V. vicugnn, 45 versus 57 mm for length of the molars and 70 versus 90 cm


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278 J. C. WHEELERfor withers height (Thomas, 1917). However, a recent study of 50 male and50 female V.V. mensalis (1.5 to 6.5 years old) from Pampa Galeras, Peru, reportsaverage withers height measurements of 86.5 cm for females and 90.4 cm forthe males (Paucar et al., 1984). N o clearly defined geographic separation existsbetween the two proposed vicuiia subspecies and many authors ignore V.V.mensalis (Osgood, 1943; Gilmore, 1950; Dennler de la Tour, 1954b; Grimwood,1969; Koford, 1957). Osteological remains from Andean archaeological sitespoint to the origin of alpaca domestication from vicufia at high elevationlocalities within the range of V.V. mensalis approximately 6000 years ago(Wheeler, 1984, 1986, 1991; Moore, 1988, 1989). A third purported subspecies,V.V. e h d n e Krumbiegel, 1944 has been described based on specimens foundin German zoos.Recent survey data compiled by Torres (1992) shows vicufia populationsspread throughout the high Andean puna ecosystem (at 3800 m above sealevel and higher) from 9' 30' to 29 s latitude, without any obvious geographicclustering which might suggest two distinct subspecies (Cajal, 1992; Glade, 1992;Hoces, 1992; Villalba, 1992). None the less, sufficient phenotypic differencesappear to exist between the northern (Peruvian, Chilean, Bolivian) and southern(Argentine) vicufia populations to justify the existence of at least two geographicraces.The best studied vicuiia, the northern V.V. mensalis, is distinguished primarilyby the long growth of hair on the chest. The head, neck, back, sides anddorsal surface of the tail are a dark cinnamon colour, with white covering thelower portion of the face, the chest, belly, interior surface of the legs andventral surface of the tail. The eyes and edges of the ears are outlined inwhite. Average coat length is 3.28 cm in adult animals and the long chesthairs reach 18 to 20 cm (Hofmann et al., 1983). Fleece fibre diameter is12.52+ 1.52 pm (Carpio Solari, 1982a) and the average fleece fibre lengthis 3.2 cm in adult males (Carpio Santana, 1982). Follicle density averages78.65 per mm2 (Carpio Solari, 1982b) and the frequency of primary hairin the fleece is 2 /0 (Carpio Solari, 1982a). In contrast, V.V. vicugna lacks thelong chest hairs, and has a lighter beige pelage coloration with white coveringa greater portion of the body, rising halfway up the sides to mid-rib heightand all the way to the ilium crest, as well as covering the anterior portion ofthe rear legs.Total length measurements from the Paucar study of V.V. mensalis fromPampa Galeras (Paucar et al., 1984) are 96.3 cm for females and 110.7 cmfor males, with average weights of 33.2 kg and 36.2 kg respectively. Thesefigures contrast with the total length measurements of 137-181 cm reported byHofmann et al. (1983) for 19 adult vicufias from the same local. Gilmore (1950)and Pearson (1951) also report greater total lengths, 144 to 175 cm, andheavier live weights, 45-55 kg for V.V. mensalis. We have not encounteredsimilar comparative statistics for V.V. vicugna.In contrast with the continuing decline of guanaco numbers, the vicufia hasmade a remarkable recovery over the last 20 years, passing from endangeredstatus in 1969, to vulnerable in 1972 (Thornback &Jenkins, 1982). This changeis a direct result of conservation programmes. In 1957, Koford estimated thetotal Andean vicufia population to be 400000 at the most, including 250000in Peru. In 1969, Grimwood reported only 10000 in Peru, and two years later


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CAMELID E V O L U I I O N 279Jungius (1971) calculated a total of between 5000 and 10000 in Peru withanother 2000 living in Bolivia, Argentina and Chile. In 1982, 15 years afterthe establishment of a protection programme, the Peruvian population hadrisen to 62000 (Franklin, 1982), and in 1991, the total Andean populationreached 92 882 (Wheeler, 199 1).In Peru, conservation efforts at the Pampa Galeras reserve began in 1967with 1753 vicufias. During the first years, an unexpected annual populationincrease of 21% was registered at Pampa Galeras (Sanchez, 1984). This highgrowth rate has subsequently been repeated when Chilean and Bolivian reserveswere established (Rodriguez Torres, 1981; Rabinovitch, Hernandez Cajal,1985). In 1978-79 however, the Pampa Galeras population reached a crisisbrought on by a prolonged drought, overgrazing and over-population, and anegative 11.28% growth rate was registered. During this period pregnancy ratesdecreased from 85% to 58% and recruitment dropped from 76% to 27% (Otte

Hofmann, 1981). Abortion rates rose dramatically and pregnancy was delayedfrom once a year to once every two years (Mtnard , 1982) while adult mortalityincreased from 5.6% to 27.6% (Sanchez, 1984). The present situation appearsto have improved remarkably, but exact census data are lacking due to recentsecurity problems in the area.

ORIGIN OF THE DOMESTIC FORMSTo date the earliest evidence of camelid domestication comes from

archaeological sites located between 4000 and 4900 m elevation, in the punaecosystem of the Peruvian Andes. Both guanaco (L.g. cacsilensis) and vicufia(V.V. mensalis) have inhabited this tundra environment for approximately 12 000years and, together with the huemul deer Hippocamelus antisensis (d'orbigny,1834), were the primary prey of early human hunters. Faunal materials fromarchaeological sites (Wing, 1986; Wheeler, 1984, 1986; Wheeler et al., 1976;Moore, 1988, 1989) indicate that during the earliest occupation of this zone12 000 to 7500 years ago, approximately equal numbers of camelids and deerwere hunted, while during later periods the frequency of camelid remainsincreased dramatically suggesting a shift to the utilization of domestic animals.Archaeozoological data from one of these sites, Telarmachay Rockshelter, haveproduced the most extensive evidence concerning this shift to date (Wheeler,1984, 1986).Located 170 km north-east of Lima, Peru (1 1 11's latitude and 75 52'Wlongitude), at 4420 m above sea level, Telarmachay is situated near the absoluteupper limits of crop growth potential. Mean annual temperature is 4.8 C, withan average daily variation of greater than 20 and frost occurs 330 nights ofthe year. Annual precipitation averages 500 to 1000 mm and is normallyrestricted to the months from November to March, although the timing isirregular and unpredictable, and extended periods of drought occur. Noagriculture is practised in the area today, and grazing ungulates represent themost reliable food resource. This is due to their mobility during periods ofdrought and their ability to convert the dry ligneous puna grasses into asource of stored protein which can be utilized for human consumption.Palaeoclimatological data indicate that no significant climatic changes have


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280 J C \YHEEI,ERtaken place in this area over the last 10000 years (Van der HammenNoldus, 1986).

Five seasons of excavation at Telarmachay Rockshelter (Lavallee et al., 1986)revealed a 8200 year long occupational sequence and recovered more than onemetric ton of animal bones fiom the preceramic levels. Archaeozoologicalanalysis of these materials produced evidence of a shift from generalized huntingof guanaco, vicufia and huemul deer 9000-7200 years ago, to specializedhunting of guanaco and vicufia approximately 7200-6000 years ago, then tocontrol of early domestic alpacas and llamas by 6000-5500 years ago, andfinally, to the establishment of a predominately herding economy beginning5500 years ago (Wheeler, 1986, unpublished data). It has not been possible todetermine if these shifts were associated with body size reduction as has beendocumented for other domestic ungulates because species specific characters forseparating postcranial hones are lacking. Instead, determination of early cameliddomestication at Telarmachay is based upon an increase in the frequency ofboth camelid and neonatal camelid remains, together with changes in dentalmorphology. During the preceramic period, 9000 to 1800 years ago, camelidremains gradually increased from 64.7 /0 to 88.6% of the faunal assemblage,while deer remains diminished from 34.20/0 to 9.2% of the total (Wheeler,1986). This shift was not caused by decreased availability of deer in the zone,but rather, by a change in animal utilization patterns from generalixd tospecialized hunting and cventual domestication of the camelids.

Between 9000 and 6000 years ago, camelid remains increased from 64.7%to 81.7% of the total faunal sample, with-just over one third (35.3'10 to 37.1°/0)of the bones coming from fetal or neonatal animals (Wheeler, 1986). Thesefigures are consistent with a hunting economy because between 35'/0 and 40'/0of animals in contemporary guanaco and vicufia populations fall within thiscategory (Franklin, 1978, personal communication). Thus, the ever greaterdependence upon camelids in the diet during this period suggests increasingspecialization in guanaco and vicufia hunting.

Around 6000 years ago, however, the frequency of foetal and neonatalcamelids increased markedly to 56.80/0, and continued to rise until it reached73.Oo/o of all camelid remains in the deposits dated to 3800 years ago (Wheeler,1986). These figures suggest either the development of specialized hunting ofneonates, an economically unviable strategy, or the appearance of other mortalityinducing factors in the environment. They far exceed expected frequencies forboth the foetal/neonatal age group and the natural (i.e. no human hunting)mortality rates of 4.5 (Raedeke, 1979: 199) to 30% (Franklin, 1978: 42) whichhave been recorded for the guanaco and vicufia, but closely correspond withmortality rates experienced by llama and alpaca breeders today.At present, up to 7Oo/o of each year's young may be lost before two monthsof age due, in part, to failure of passive immune transfer (Garmendia et al.,1987) with resulting mortality from Clostrr'dium fxrfringenf Type A eiiterotoxaemiaand other pathogens (Leguia, 1991; Ramirez, 1991). The epizootic nature ofenterotoxaemia is to some extent controlled by climatic conditions that permitsporulation of the bacteria, as well as by the presence of a critical number ofcaptive or domestic animals. In the Andes, outbreaks of enterotoxaemia areassociated with unsanitary corralling practices during the wet season birthperiod. Similar epidemics are not known to occur in the wild camelids.


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CAMELID E\OI.UTION 28Although i t is not always possible to distinguish hctween the boncs of a

terminal eleven and one half month foetal camelid and those of a neonate,tooth wear studies indicate that the majority of Telarmachay specimens datingfrom 6000 to 3800 years ago were neonatal, whereas those from the earlierlevels were primarily foetal, presumably taken ill utero through the h~intingofpregnant females (Wiceler, 1986). This shift from predominantly foetal toneonatal remains coincides with the significant increase in frequency offoetal/neonatal remains described above, and permits the hypothesis thatmortality induced by disease rather than by intentional butchery was the cause.Additional support for this interpretation comes from the study of bonedistribution across the GO00 to 3800 year old living floors which indicates thatnewborn camclids were brought into the shelter whole and processed forconsumption. The resultant pattern is very similar to that created by conteniporarytraditional herders who utilize dead llama and alpaca neonates for food. Meatproduced by the often massive die-off of camelid neonates does not now, andapparently did not then, go to waste.

Identification of the species which was brought under domestication atTelarmachay is based upon incisor morphology. Prior to domestication (9000to 6000 BP) it is estimated that nine vicufia were huntcd for every guanacobased on incisor type and frequency. Vicufias have rootless hypselodont parallel-sided permanent incisors with enamcl covering thc entire labial surface, androot-forming deciduous incisors with enamel covering thc upper lahial surfaceonly (Miller, 1924; Wheeler, 1982, 1991 . Guanacos have rooted deciduous andpermanent spatulate incisors with an enamel covered crown (Miller, 1924). By6000 BP, however, the remains of permanent incisors with the same morphologyas deciduous vicufia incisors appear in the Tclarmachay deposits (Wheeler,1982, 1991, unpublished data). These permanent teeth match the dentition ofmany extant Peruvian alpacas in which both the deciduous and permanentincisors are root forming and parallel sided, with enamel covering only theupper labial surface (Wheeler, 1991, unpublished data). Although contemporaryalpacas with spatulate llama incisors have been reported by Kent (1982), it isunclear if these are hybrids. The evidence from Telarmachay suggests anancestral relationship which may explain the apparent retention of juvenilevicuiia dental traits in thc adult alpaca. It cannot be determined if animalswith llama type incisors also appeared in the 6000 BP deposits, since these areindi~tin~guishablerom guanaco incisors, but the presence of both large andsmall neonates suggests that this may have been the case.In contrast to the data from Telarmachay and other Andean archaeologicalsites which indicate that the llama is descended from the guanaco and thealpaca from the vicufia (Fig. 2A), other researchers have come to differentconclusions about their ancestry based on the study of living animals. In 1775,Frisch attributed the origin of the llama to the _guanaco and the alpaca to thevicuiia, an opinion subsequently supported by Ledger (1860), Darwin (1868),Antonius (1922), Faige (1929), Krumbiegel (1941, 1952), Steinbacher (1953),Frechkop (1955), Capurro Silva (1960), Akimushkin (197 1) and Semorile,Crisci Vidal-Rioja (in press). Other authors have concluded that both domesticcamelids descend from the guanaco, and the vicufia was never domesticated(Fig. 2B) (Thomas, 1891; Peterson, 1904; Hilzheirner, 19 13; L61inberg, 1913;Brehm, 1916; Cook, 1925; Weber, 1928; Herre, 1952, 1953, 1976, 1982;


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282A

J. C. WHEELERGUANACO

B.

C.

VICUNA

LLAMA ALPACA

I

CUANACO

LLAMA ALPACA

GUANACO

VICUNANOT DOMESTICATED

LLAMA VICUNA

LPACAFigure 2 . Evolutionary models of llama and alpaca domestication. A , Model derived fromarchaeozoological evidence in which the llama is domesticated from the guanaco and the alpacafrom the vicuiia, with subsequent hybridization between the two domestic forms producing thewan. B, Model derivcd from study of extant animals in which both the llama and alpaca descendfrom the guanaco and the vicuiia was never domesticated. C, Model derived from study of extantanimals in which the llama is a domesticated form of the guanaco and the alpaca results fromcrossing the domestic llama with the wild vicufia.

Rohrs, 1957; Fallet, 1961; Zeuner, 1963; Herre Thiede, 1965; HerreRohrs, 1973; Bates, 1975; Pires-Ferreira, 1981/82; Kleinschmidt et al., 1986;Kruska, 1982; Jiirgens et al., 1988; and Piccinini et al., 1990). In the 1930s,Lopez Aranguren (1930) and Cabrera (1932) suggested that llama and alpacaevolved from presently extinct wild precursors, based on the discovery of 2Myr Plio-Pleistocene L. glama, L. pacos, L. guanicoe and V. vicugna fossils inArgentina, and that the guanaco and vicufia were never domesticated. Thisposition is no longer considered a possible alternative. Finally, Hemmer (1975,1983, 1990) attributes llama ancestry to the guanaco, but has deduced on thebasis of shared morphological and behavioural traits that the alpaca originatedfrom hybridization between the llama and vicuiia (Fig. 2C).

Conclusions about llama and alpaca ancestry have, in large part, been basedupon morphological changes produced by the domestication process. Duringthe 1950s, Herre and Rohrs (Herre, 1952, 1953, 1976; Herre Rohrs, 1973;Rohrs, 1957) examined alterations in the mesotympanal area of the skull relatedto a decrease in llama and alpaca hearing acuity, and reported an overallreduction in cranial capacity of both domestic species relative to the guanaco.


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CAMELID EVOLUTION 283In contrast, they found the vicufia cranium to be the smallest of all livingLamini, and, based on the premise that domestic animals are smaller thantheir ancestors, concluded that this species was never brought under humancontrol. Herre and Rohrs consider the llama and alpaca to be “races of thesame domestic species bred for different purposes’’ (Herre, 1976: 26). Researchon the relationship of brain size relative to body size by Kruska (1982) alsofound the vicufia to be smaller than alpaca and llama, which in turn weresmaller than the guanaco suggesting that the latter is the only ancestral form.None the less, papers by Jerison (1971) and Hemmer (1990) report the ratioof alpaca brain size to body size to be smaller than in the vicuiia permittinga different conclusion about origins of the domestic forms. These contradictorydata on size reduction are almost certainly a product of sampling, as neithersubspecific variation in the wild forms nor the possibility of hybridizationbetween the domestic animals were considered in any of the studies.

Based on the study of pelage characteristics (skin thickness, follicle structure,secondary/primary ratio, fibre length and diameter, coloration) in living camelids,Fallet (196 1) found the llama to be an intermediate evolutionary stage betweenthe wild guanaco and the specialized fibre-producing alpaca, and concludedthat the absence of transitional characteristics between vicufia and alpaca fleeceseliminates the former from consideration as an ancestral form. This deductionis, in part, based on the assumption that llamas have been selected exclusivelyfor use as pack animals while alpacas have been bred for fibre production.None the less, new data on preconquest llama and alpaca breeds in Peru haverevealed the prior existence of a fine fibre-producing llama, as well as an extrafine fibre alpaca which is transitional between the vicufia and a secondprehispanic fine fibre alpaca breed (Wheeler, Russel Stanley, 1992; Wheeler,Russel Redden, submitted).Research on camelid behaviour has produced contradictory hypothesesconcerning llama and alpaca origins. Krumbiegel (1944, 1952) and Steinbacher(1953) argue that the alpaca is the domestic vicufia based on unique sharedbehavioural traits which are said to differ from those observed in the guanacoand llama. Hemmer, on the other hand, concludes that while some alpacabehaviour patterns match those of the vicufia, others are intermediate betweenthose of vicufia and guanaco, suggesting that “the alpaca is a mixture of bothlines, [produced] by crossbreeding of captured vicuiias with the only initiallyavailable domestic animal, the llama” (1990: 63). It has also been suggestedthat the vicuiia was never domesticated because it is more territorial than theguanaco (Franklin, 1974). None the less, this assumption is open to questionbecause it is based upon study of guanacos located at the southernmost extremeof their range where seasonal migration in response to severe climatic changesis essential for survival (Franklin, 1982, 1983). Further to the north, wherevicufia and guanaco ranges overlap and llama and alpaca domestication occurred(Wheeler, 1984), a more benign climate and a constant food supply permit thecharacteristic sedentary social organization of the vicufia (Franklin, 1982, 1983).Although data concerning behaviour of the guanaco in this region are lacking,it is possible that the limited sedentary territorial organization observed in somePatagonian groups plays a more important role in these less extreme climaticconditions.

Analysis of haemoglobin amino acid sequences in vicuiia, alpaca, llama and


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84 J. C . WHEELERguanaco from Hannover Zoo, Germany, led Kleinschmidt et al. (1986), Jurgenset al. (1988), and Piccinini et al. (1990) to the conclusion that the vicufia wasnever domesticated. However, earlier research on blood and muscle samplesfrom llama, alpaca, vicuiia, guanaco and alpaca x vicufia hybrids at SantiagoZoo (Cappuro Silva, 1960) indicated a llama-guanaco and alpaca-vicuiiasubdivision, as have more recent data from ribosomal genes (Semorile et al., inpress). Other researchers utilizing immunological, electrophoretic analysis andprotein sequencing have found it impossible to draw conclusions about llamaand alpaca ancestry (Miller, Hollander Franklin, 1985; Penedo et al., 1988).Cytogenetic studies (Capanna Civitelli, 1965; Taylor et al., 1968; Larramendyet al., 1984; Gentz Yates, 1986) indicate that all four species of the SouthAmerican Camelidae have the same 211 = 74 karyotype, but information onmolecular biology is limited. Vidal Rioja et al. (1987) and Saluda-Gorgul,Jaworski Greger (1990) have examined satellite DNA, and research analysingthe full mitochondria1 cytochrome b gene sequence in all six Camelidae hasdocumented hybridization among the domestic South American camelids (Stanleyet al., 1994). Recent studies of the fibre from mummified ninth and tenthcentury llamas and alpacas suggests that post-conquest hybridization has modifiedthe genetic makeup of living populations (Wheeler et al., 1992), a fact whichmay well explain the diversity of conclusions about their ancestry.

' I 'H E D O M E S T I C S O U T H A M E R I C A N C A M E L I D A EThe llama Lama glama (Linnaeus, 1758)

The llama is the largest of the domestic South American camelids andresembles its ancestor in almost all aspects of morphology and behaviour. Likethe guanaco, the llama has adapted to a wide range of environments (Fig. 1C).After domestication in the Peruvian puna between 7000 and 6000 years ago(Wheeler, 1984, 1991; Wing, 1977, 1986), the llama was moved to the lowerelevation interAndean valleys and into northern Chile where their remains havebeen found in archaeological sites dated to 3800 years ago (Wing, 1986; Hesse,1982; Dransart, 1991a). Some 2400 years later they were being bred on thenorth coast of Peru (Shimada Shimada, 1985) and in Ecuador (Wing, 1986;Stahl, 1988; Miller Gill, 1990). Although it is often assumed that the LakeTiticaca region was also a centre of llama domestication, relevant data arelacking from early archaeological sites in Bolivia (Browman, 1989). In north-western Argentina, a single cranium of L. glama has been dated to 3400 yearsBP with stronger evidence for herding at 1450 BP (Yacobaccio Madero,1992; Reigadas, 1992), and it is thought that domestication may have occurredindependently in both this region (ibid.) and northern Chile (Hesse, 1982).Shortly thereafter, 900-1000 BP, evidence of llama rearing has been recoveredat sites located in the cloud forest on the eastern slope of the central Andes,as well as in the dry Osmore drainage of south coastal Peru (Wheeler, 1991,in press). Under Inca rule (1470-1532) llama distribution reached its furthermostexpansion as pack trains accompanied the royal armies to southern Colombiaand central Chile. It is impossible to estimate the size of this preconquest llamapopulation, but it clearly must have exceeded present numbers for early Spanishadministrative documents record the virtual disappearance of these animals


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CARIEIJD EVOLUTIONLLAMA ALPACA

285

COARSE FIBREFINE FIBRE

P R E C O N Q U E S T B R E E D S

FINE FIBREEXTRA FINE FIBRE

Q RA W A k S UR ICH'KU PACOWARI HUACAYAINTERMEDIATE LLAMAWARI INTERMEDIATEFigure 3 . Pi)st-doniestica1ioii llama and alpaca evolutioii sho\ring the relationship betwccnpreronquest hrccds and cxtant phenotypes produccd b y a hrcalidown in controlled hrceding andhybridization.

within a century of contact (Flores Ochoa, 1977). In recent years the llamapopulation has remained relatively stable, totalling 3 776 793 in 1991 (Wheeler,

Because Andean civilization was nonliterate, knowledge of pre-Spanish llamaand alpaca herding practices must be reconstructed from archaeological remains.The recent discovery of 900-1000 year old naturally desiccated llamas andalpacas at El Yaral, an archaeological site in the Moquegua valley of southernPeru (Rice, 1993), has provided a first view of preconquest breeds (Wheeler etal., 1992; Wheeler et al., submitted). Associated with the pre-Inca Chiribayaculture, these animals had been sacrificed by a blow between the ears andimmediately buried beneath house floors where thcy became naturally mummifieddue to the extreme aridity of the environment.

Research on the physical appearance of the El Yaral llamas, as well asanalysis of skin and fibre samples taken at 11 different locations across thebody, revealed the possible existence of both a fine fibre and a coarse fibrebreed (Wheeler e f nl., 1992; Wheeler et nl., submitted) (Fig. 3). Average fleecediameter of the former was found to be 22.2 with a between sample standarddeviation of 1.8 pm, compared to 32.7 (SD k4 .2 ) pin for the latter, based onthe measurement of up to 1600 fibres per animal. Th e reduction of both fibrediameter and variation in the fine fibre llama fleece was certainly produced byselective breeding for a single-coat through modification of the primary hair toresemble secondary undercoat fibre. The uniform coloration and fineness, aswell as the absence of visible hairs in the El Yaral fine llama fleeces are ideallysuited for textile production, and contrast markedly with the multicoloureddouble-coat of the coarse fibre breed. An additional evidence of specializedbreeding is the accelerated fibre growth rate recorded for El Yaral fine llamas

1991).


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286 J. C . WHEELERrelative to contemporary animals (Wheeler et al., submitted). The growth curveand live weights of the llamas from El Yaral, and other Chiribaya culture sitesof the same region, are very similar to those of contemporary llamas raised inthe puna, and their age at death reflects controlled stockrearing with eliminationof undesirable animals from the herd (Wheeler, in press).Prior to discovery of the El Yaral mummies, our most detailed data onpreconquest camelid breeding practices came from written documents of thecolonial period. These records describe the use of llamas as pack animals forthe Inca army, but make no mention of fine fibre producing llamas. This maybe due to the general failure of the early Spanish writers to distinguish betweenllamas and alpacas, as well as their special interest in pack animals for use intransporting ore. Despite their European perspective, these documents do providedetails about Inca husbandry. Expansive state and shrine herds were managedby the llama camayoc, members of a hereditary caste of herding specialists,and emphasis was placed on breeding pure brown, black and white animalsfor sacrifice to specific deities, as well as on quality fibre production for thestate controlled textile industry (Murra, 1965, 1975, 1978; Brotherston, 1989).Detailed data on size and colour of flocks were kept utilizing the quipu, amemory assistance device made of knotted camelid fibre cords. Communallyand individually owned herds also existed.

Native Andean stockrearing was largely destroyed by the arrival of theSpanish. Within little more than a century of the conquest in 1532, administrativedocuments record the disappearance of approximately 90% of the domesticcamelids (Flores Ochoa, 1982), as well as 80% of the human population(Wachtel, 1977). Coastal and highland valley herds were the first to disappear,as their grazing lands were usurped for the production of sheep, goats, cattleand pigs. In the puna this process was somewhat slower because both theSpanish and their livestock found the harsh climate and extreme elevationinhospitable. This region became a refuge for native livestock and herders, andtheir descendants continue to inhabit the same marginal lands today. Theprolonged Spanish civil wars and heavy tribute levies, paid either in domesticcamelids or in money obtained from their sale, resulted in depletion of theherds. Introduced livestock diseases may also have played an important role inthis process. By 1651, llamas and alpacas had practically disappeared even inthe Lake Titicaca basin (Flora Ochoa, 1982), the former heartland of theirdistribution (Murra, 1975). The impact of such catastrophic mortality uponcamelid genetic diversity and breeding practices has yet to be fully explored.Today, the total llama population is estimated to be 3 776 793 (Wheeler, 1991).Small groups are found near Pasto, Colombia (1 N latitude) and Riobamba,Ecuador (2 s latitude). To the south they extend to 27 in central Chile, butthe most important production zone is located between 11 and 21 s latitudeat elevations of 3800-5000 meters above sea level.

The name llama comes from Quechua (Flores Ochoa, 1988), and it is knownas qawra by Aymara speakers (Dransart, 1991b). Although specific llama breedsdo not exist, at least three varieties of llama are recognized. Most llamas inPeru, Bolivia and northern Chile are of the 'nonwoolly' phenotype characterizedby sparse fibre growth on the body and the absence of fibre on the face andlegs. To the south, especially in Argentina, the 'woolly' llama is more commonand has a greater density of fibre on the body which extends forward between


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CA ME L ID E V O L U T IO N 287the ears and grows from inside the ears but is absent on the legs. The woollytype is known as ch’aku in Quechua (Flores Ochoa, 1988) and t’awrani inAymara (Dransart, 1991b), while the nonwoolly type is called q’ara in bothlanguages (ibid.). In both areas llamas with intermediate phenotypes are alsorecognized. Recent research on the fibre characteristics of Argentine llamas hasidentified the existence of seven distinct fibre types in the population (FrankWehbe, 1994), raising the possibility that more than three varieties of llamaexist. A different classification has been proposed by Cardozo (1954: 61) whodivides llamas into brachymorphic (round, short profile, abundant fibre) anddolichomorphic (narrow, elongated profile, sparse fibre).

The vast majority of llamas are held by traditional Andean pastoralists whoutilize elaborate classification hierarchies based on colour, fibre and conformationcharacteristics to describe their animals. The existence of these systems amongboth Quechua (Flores Ochoa, 1988) and Aymara (Dransart, 1991b) speakingherders suggests that earlier management strategies may have been directed atproducing animals with specific fibre types, but it is not clear to what extentselection is made for these characteristics today. Contemporary llamas lack thephenotypic uniformity associated with true breeds, and Flores Ochoa (1988)indicates that the primary breeding criteria used by Quechua speaking herdersin southern Peru is to divide llamas into ‘allin millmayuq’ and ‘mana allinmillmayuq’ or fine and coarse fibre animals. Pelage coloration varies from whiteto black and brown passing through all intermediate shades with a tendencyto spots and irregular colour patterns, and llamas with wild guanaco colorationoccur. Fleece quality is uneven, with wide variation in fibre diameter and astrong tendency to hairiness, ranging from 32.5 Ifr 17.9 pm (9) to 35.5 Ifr 17.8pm 6)or coarse ‘nonwoolly’ q’aras, 30.5) 18.5 pm (0) to 30.5f 17.9 pm 6)for intermediates, and 27.0+ 15.6 pm (0) to 29.1 12.7 pm 6)or ‘woolly’cha’kus (Vidal, 1967). For this reason, the primary value of the llama presentlylies in its use as a pack animal rather than as a fibre producer. The variabilityof present day llama fibre is related to an increase in hairs and generalcoarsening of the fleece, which probably began at the time of the Spanishconquest. Increased hairiness is produced by lack of controlled breeding, andcrossing between the two prespanish llama breeds from El Yaral could accountfor the entire range of fleece variation observed in todays animals (Fig. 3).

The alpaca Lama pacos (Linnaeus, 1758)The alpaca is smaller than the llama and resembles the vicufia in manyaspects of morphology and social organization. Although the Lake Titicacabasin of southern Peru and Bolivia has long been considered the focus ofalpaca domestication, archaeological evidence is not presently available toevaluate this hypothesis (Browman, 1989). Nevertheless, excavations in thecentral Peruvian puna have placed its origins between 7000 and 6000 yearsago (Wheeler, 1984, 1986), and it was from this region that the alpaca wassubsequently moved to lower elevation interAndean valleys 3800 years ago(Wing, 1972; Shimada, 1985). Evidence of alpaca rearing at coastal sites insouthern Peru dates from 900 to 1000 years ago (Wheeler et al., 1992; Wheeler,in press; Wheeler et al., submitted) (Fig. 1D). Precolumbian alpaca remains havenot been reported in the faunal materials from archaeological sites in Chile


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288 J. C. WHEELER(Dransart, 1991a b; Hesse, 1982), Argentina (Elkin, personal communication)or Ecuador (Miller Gill, 1990), although they are found in limited numbersin these regions today. It is impossible to estimate the number of preconquestalpacas. Spanish documents record their rapid decimation and displacement toremote, extreme high elevation regions of the Andes (Flores Ochoa, 1977).Recent data indicates that over the last 25 years alpaca numbers have fallensignificantly in Peru, from 3 290 000 in 1967 to 2 510 912 in 1986, and in 1991the total Andean alpaca population was estimated to be 2 811 612 (Wheeler,

Representatives of two possible preconquest alpaca breeds have been foundamong the 1000 year old El Yaral mummies. Fine fibre and extra fine fibrealpacas were distinguished, based on physical appearance and average fibrediameter (1600 fibres measured per animal). The former have fleeces averaging23.6 (SD f 1.9 pm), while the latter fleeces average 17.9 (SD f 1.0 pm) (Wheeleret al., 1992; Wheeler et nl., submitted). Both groups had lustrous fibre rangingfrom wavy to crimped and dense to very dense. Hairs were visible in three ofthe four animals, but were not significantly coarser than the undercoat fibres.Indced, fibre diameter variation both within and across the fleece was remarkablylow, suggesting that rigorous breeding selection for fine quality fibre was beingpracticed (Fig. 3).The Spanish conquest had a disastrous effect on both llama and alpacapopulations. Massive mortality accompanied the displacement of alpaca herdsfrom the coast, interAndean valleys and most of the puna, as introducedstockrearing practices pushed the survivors into the marginal, extreme highelevation pastures where they are found today (Flores Ochoa, 1982). At prescnt,alpaca distribution extends from approximately 8 s latitude, where they havebeen recently reintroduced in Cajamarca, to 20 s latitude, in the vicinity ofLake Poopo, Bolivia, with small populations located further to the south innorthern Chile and north-western Argentina (Fig. 1D).

Today, 75 of all alpacas, paqocha in Quechua (Flores Ochoa, 1988) andallpachu in Aymara (Dransart, 1991b), are held by traditional herders (Novoa,1989). Two alpaca phenotypes, known in the literature by their Quechua namesas suri and huacaya or wakaya, are recognized but these do not breed true.The suri has long straight fibres, organized in waves which fall to each sideof the body in much the same manner as a Lincoln sheep, while the huacayahas shorter, crimped fibres which give it a spongy appearance similar to thatof a Corriedale sheep. Occasionally animals with intermediate wood characteristicsare seen, and these have been named chili by Cardozo (1954). Crosses betweenhuacaya and huacaya produce a certain percentage of suri offspring, and crossesof suri with suri produce some huacaya offspring. Although no artificial selectionis made, an estimated 90% of all alpacas are huacayas (Novoa, 1989). Thesuri is not known among the Aymara herders of Chile who refer to theirhuacayas simply as allpachu or alpacas (Dransart, 1991b). The fleece of bothphenotypes varies from white to black and brown, passing through allintermediate shades, with a greater tendency to uniform coloration than in thellama. Alpacas with wild vicuiia coloration occur.

In comparison to the preconquest El Yaral alpacas, contemporary Andeanhuacaya and suri fleeces average 31.2k3.8 pm (Carpio, 1991) and 26.81f16.0pm (Von Bergen, 1963) respectively, are coarser, may have a tendency to

199 1).


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C A M E L I D E V O L U T I O N 289hairiness, and are of uneven quality. Some coats containing up to 40% hairhave been reported for both living varieties, and considerable variation isreported in published statistics on fibre diameter. The origin of this degenerationalmost certainly lies in the Spanish conquest, but a breakdown in controlledbreeding between the fine and extra fine El Yard breeds would not aloneaccount for the variation observed today.

The most probable cause of coarsening and hairiness in both huacayas andsuris would be through hybridization with the coarse fibre llama breed, a notimprobable scenario amid the chaos and destruction of the conquest. Clearly,however, such a process would not have affected only the alpaca gene pool.The El Yaral mummies indicate the possibility that extensive crossbreedingbetween alpacas and llamas may have occurred since the Spanish conquest andhas played a much more important role in the formation of today’s livestockthan has been realized (Fig. 3).

HYBRIDIZATIONThe guanaco, vicufia, llama and alpaca all possess the same 2n = 74 karyotype

(Capanna Civatelli, 1965; Taylor et nl., 1968) and can, under humaninfluence, produce fertile hybrids (Gray, 1954). Preliminary research on thecytochrome b gene sequence has found no evidence of hybridization betweenguanaco and vicufia (Stanley et a/., 1994). Since this study included samplesfrom the northern populations of both genera, the region where they aresympatric in parts of their range, the findings may possibly support Miller’s(1924) creation of the genus Kcugna.Traditional herders recognize the existence of llama and alpaca crosses. Theseare referred to by the generic terms wari in Quechua (Flores Ochoa, 1988)and wik‘ufia in Aymara (Dransart, 1991b). These hybrids are classified asllamawari or llama-like and paqowari or alpaca-like by Quechua speakers (FloresOchoa, 1977). Aymara speaking herders use waritu and wayki for llama andalpaca phenotype hybrids, as well as the generic term wakayu for anyllama x alpaca offspring (Dransart, 1991b). First generation crosses are easilyrecognized, but it is not always possible to identify hybrid animals based uponphenotype alone because it is likely that hybridization has been an ongoingprocess since the time of the Spanish conquest. The extent to which contemporaryllama and alpaca populations have been affected by this process has not beendetermined, but comparison with preconquest animals suggests that it has beenextensive and that breeds of fine fibre-producing llama and alpaca have likelydisappeared in the process (Wheeler et al., 1992). Hybridization has beenconfirmed through DNA analyses (Stanley et al., 1994) (Fig. 3).Crosses between the wild and domestic South American camelids producefertile offspring, but do not normally occur in nature. The pacovicufia, oralpaca x vicufia hybrid, has received considerable attention for its potential asa fine fibre producer. Carpio et al. (1990) report fibre diameters ranging from13.3 to 17.3 pm for five first generation crosses, but this is said to rapidlyincrease in subsequent generations. The pacovicufia phenotype closely resemblesthat of the vicuiia, although it is slightly larger and less gracile than its wildprogenitor. Research on the fixation of phenotypic traits from generation to


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290 J. C. WHEELERgeneration of alpaca x vicuiia hybrids is lacking, and much remains to be donebefore its potential as a fine fibre producer can be evaluated.

The possibility that feral llamas and alpacas exist and might have crossedwith wild camelids has not been fully explored. In 1534, Xerez observed thatdomestic llamas were sometimes so numerous some escaped to the wild, andin 1555 Zarate recorded that once each year some llamas were released intothe wild as an offering to the gods (Murra, 1978). It is unclear, however, ifferal populations existed at that time. The current consensus of opinion in thecentral Andean region is that no such populations exist today. Even so,MacDonagh (1940) has described a group of guanaco and llama hybrids livingin a feral state in the Province of Cordoba, Argentina. These animals werethe product of natural crosses, and generally exhibited the guanaco phenotype,although some had white blotches on the head and upper part of the neck,and others are almost entirely white. No observations on changes in body sizeand fibre quality were recorded. The behaviour of these feral hybrids wasconsidered to be virtually identical with that of the guanaco, and they livedand reproduced without problem.

THE FUTUREThe present status of the South American camelids is the product of a

largely unknown past. To name but two historic factors, the potential influenceof genetic bottlenecks and/or hybridization in the formation of contemporaryguanaco, vicuiia, llama and alpaca populations have never been fully investigated,although there is evidence to suggest that they may have played an importantrole. The most basic questions concerning genetic variation and the systematicclassification of presumed guanaco and vicuiia subspecies, as well as llama andalpaca breeds, remain to be answered, although such information is essentialfor ensuring their future. In the case of the wild camelids we need to be surethat we are protecting all genetic variants of each species, and not justincreasing the numbers of potentially genetically impoverished subgroup/s. Inlight of the increased movement of both wild and domestic camelids throughoutthe Andes and the beginning of exportation in 1983, there is an urgent needto identify relict populations of genetically pure precolumbian llama and alpacabreeds in order to insure both their preservation and the possibility of a returnto high quality fine fibre production.

REFERENCESAkimushkin J. 1971. M@aciones. Buenos Aires: Editorial Cartaga.Antonius 0 1922. Stummesge.schichte der Haustiere. Jena.Bates M . 1975. ,Sudanierika-Flora und Fauna. Reinbek Rowehlt.Benavente MA. 1985. Reflexiones en torno a1 proceso de domesticacion de 10s camelidos en 10s valles delBohlken H. 1960. Remarks on the stomach and the systematic position of the Tylopoda. Proceedings o theBrehm A. 1916. Die Suugetiere. Lcipzig, Bihiographilches Institut.Brotherston G. 1989. Andean Pastoralism and Inca ideology. In: Clutton-Brock J, ed. ’The uialking larderp a t t m s of domestication, pastoralih and predation. London: Unwin Hyman Ltd, 240-255.Browman DL. 1989. Origins and development of Andean pastoralism: an overview of the past 6000 years.

In: Clutton-Rrock J, ed. The uialking larder patterns of dumesticatiun, pastoralisnr and predation. London: UnwinHyman Ltd, 256-268.

centro y sur de Chile. Boletin del h h e o Regional Araucania 2: 37- 52 .


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