Evolution and present situation of the South American camelidae

Evolution American JANE Farultad .Qzrtado

and present Camelidae

situation

of the South

C. \VHEELER de Aledicina I ‘eterinaria, Uniz~ersidad .Nacional i\fg>lor de San A farces, 4 I-0068, Lima II, Peru

C:ON’I‘EN’I‘S Imroductior~ ‘I‘hr wild Sot~th Amcriran Camclidac Thr pm”‘0 haa ,&ymime (RIdtrr, 1776) The viruiia licks &pa (hlotina, 1782) O+jn ofrhr domcsuc tixms ‘l’hc domestic South Amcriran Cam&lx ‘I‘hc llama lornngla~r~ (I.innacus, 17.58) The alpaca Lnmo pncrrr (Linnacus, 1758)

271 271 273 277 279 284 285 287 289 290 290

Hybridixation The t-ururc Rrlrrcnces

INTRODUC’I’ION

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, Luma 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 distinguished by: absence of horns or antlers, presence of true canines separated from the premolars by a diastema in both the upper and lower jaws, position 271 0024-4066/95/00327

I +25

$08.00/0

0

t 995 The

Linncan

Society

of London

of the vertebral artery confluent to the neural canal in the cervical vcrtcbrac, anatomy of the rear limbs which permits the animal to bend its legs hcncath the body and rest on its stomach, and the prcscncc of a nail covcrcd dQgital pad rather than a hoof. In 1758, Linnaeus described the two domestic New \Vorld camclids as Gzmelus glamn “Camelus peruvianus Glama dictus” (llama) and Chmelus /xzro.r “Camelus peruvianus laniger Paces dictus” (alpaca), placing them together in a single genus with the Old \Vorld dromedary and hactrian camels, Ci2melu.s dromedatius and Camelus barhianus. The two remaining New \Vorld spccics, the wild
togcthcr with the Olcl \Vorld domestic dromedary Gmzelus dromedrrrius Linnaeus, 1758 and domestic and wilcl Bactrian camels C: bactlia~r Linnaeus, 1758. The trihc Lamini, reprcsentcd by fossils of the genus Pliaucheniu, originated in the Great Plains of wcstcrn North America between 9 and 11 million years ago (Myr) (Harrison, 1985) (Tahlc 2). Two gcncra, illfo@ (10-4.5 Myr) and Heminuchenio (10-O. 1 Myr) (Webb, 1965, 1974) evolved arrison, 1979) W from Pliauchenia approximately 10 Myr. The first of these, illjoljns, and its dcsccndant C’anzelops (4.5-0.1 Myr) remained in North America (Harrison, 1979; Webb, 1965, 1974), while some species of Hemiauchenia migrated to South America during the Pliocene/Pleistocene transition approximately 3 Myr. It is from the latter genus that Lama and Vicugna evolved in South America approximately 2 Myr (L6pez Aranguren, 1930; Cabrcra, 1932; Webb, 1972; Harrison, 1985). Palaeolamn (1.5-o. 1 Myr), the other descendant of Hemiauchena, is no longer considered to be an ancestor of Luma and Vicugna. Only Luma and Vicugna survived the end of the Pleistocene period some 10 000 years ago. THE

\VII>D

The guanaco

SOU’I’H

AAIEKICAN

C.ALlEI.IDAE

Lu ma guuanicoe(Mtiller,

1776)

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

274

date to some 2 Myr (Webb, 1974) (Fig. 1A). They arc also present at Tarija, Bolivia (Hoffstetter, 1986) in strata recently dated 97773 000 years BP (before present) (MacFadden et al., 1983), but did not spread into the high Andean puna ecosystem before the establishment of modern climatic conditions between 12-9000 years ago (Hoffstetter, 1986). Prior to European contact,
I :30 I.50 348 8 16

ml

:330 350 Ii 7

the European conquest. During the nineteenth century the impact of indiscriminate hunting and commercial sheep rearing reduced the guanaco population to 7 million, and at present an estimated 602 907 survive (Wheeler, 1991). All guanacos cxhihit similar pelage coloration varying from a dark reddish hro\vn in the southern populations (L.g. guanicoe) to a lighter brown with ochre yellow tones in the northern variety (L.g. rac.&~zsi.r). The chest, belly and internal portion of the legs arc more or less pure white, the head grey to black with white around the lips, eyes and borders of the ears. Fibre diameter varies from 16.5 pm to 24 pm and contains from 5 to 20% hair (Verscheure is absent except & Garcia, 1980; Carpio & Solar-i, 1982a). Sexual dimorphism for the presence of large canines in the male. Withers height of adult animals varies from 110 to 120 cm for L.g. guaniroe from Patagonia and Tierra de1 Fuego (Cabrera & Yepes, 1960; Franklin, 1982; Herre, 1952; Raedeke, 1979; MacDonagh, 1949), compared to 100 cm for the small northern guanaco L.g. cacsifensis(Herre, 1952). Reported body length, from the tip of the nose to the base of the tail, varies from 167 (MacDonagh, 1949), 185 (Cabrera & Yepes, 1960), 19 1 (Raedcke, 1979) and 2 10 cm (Dennler de la Tour, 1954a) for L.g. guanicoe and 90-100 cm for L.g. cacsilensti from Calipuy, Peru (Kostritsky & Vilchez, 1974). Live weight for adult L.g. guanil-oe varies from 120 to 130 kg (Raedeke, 1979; M i 11er, Rottman & Taber, 1973), compared to 96 kg for L.g. cacsilensis (Kostritsky & Vilchez, 1974). See Table 3 for comparison with the other South American camelids. Four poorly defined subspecies of guanaco have been described: the first, Lama guanicoe guanicoe (Miiller, 1776), is said to occur in Patagonia, Tierra de1 Fuego and Argentina south of 35”s latitude; the second, L.g. huanacw (Molina,

‘76

J, (:. \\'HI:.l-I.I:li

1782), is rcstrictcd to Chile; the third, L.g. rrrc.si/mi.s Liinnbcrg, 19 13, a local high altitude form from southern Peru; and the li)urth. I,.g. rqqrlii Krumbicgcl, 1944, found on the castcrn slope of the Argcntinc Anclcs bct\\.ccn approximately 21” and 32”s latitude. Thr characteristics \vhich set each sulqccics apart arc on zoogcograpliic not fully dctailcd in thcsc early works, and littlc information The most cstcnsi\.cly studiccl \rariations in guanaco morpholo~gy is a\&iblr. populations arc located at the southernmost limits of guanaco distribution, and it is clear that thcsc larger, darker animals (I>.g. g~nrric-0~)contrast markcclly \vith the smaller, lighter colourcd spccimcns (I,.g. u7tsihr.si.s) founcl at the northern boundary. Rcccnt survey data compiled by Torrcs (1992) sho\~s population clusters which may correspond to the four proposccl
~::\;\11~1.11)

I\‘oI.c”I‘Ios

‘77

1954, Ihnlcr dc la ‘I’our ( 193-1-a) called attention to the impending clisappcarancc of the Patagonian guannco if the hunting of ),carling chulcn,gos \vas not controlled ancl protcctccl I-CSCI~W cstablishrd. In 1969, Grim\t,ood found that the Pcru\iaii guanaco population \vas on the cdgc of cstinction, and in 197 1 the go\‘crnmcnt rcsponclccl by clcclaring it an cndangcrcd spccirs. In 1974, the IUCN (International Union fi)r the Conscl\.ation of Nature and Natural Rcsourrcs) clcclarccl the guanaco a ~~ulncrablc spccics (‘l’hornback & Jenkins, 1982). At prcsrnt the guanaco rccci\rcs protection in I-1 rcscilres in Argentina, four in Chile ancl three in Peru. None the Icss, thcsc mcasurcs arc insullicicnt ancl in Bolivia and Paraguay ~guanaco populations arc not protcctccl. Rcccntly the South American Camclid Specialist Group of the IUCN has urgcntl) rcroninicnclccl increasing protection li)r the guanaco in gcncral, and the relict northern L.g. c-ac~.ri/en.ci.s in spccilic (l’orrcs, 1985). This animal is not only highl) cndangcrcd but also \+rtually unkno\\~i to scicncc, and the rcconimcnclation should bc taken seriously since archacozoolo,~cal and cthnohistorical data dcmonstratc basccl on geographic distribution that L.g. racsifer~sisis the ancestral Torm of the domestic llama (\\‘hcclcr, 1984, 1986, 199 1).

At prcscnt viculia distribution is limited to arcas of cxtrcmc elevation bctwcen 9” 30’ and 29”s latitude in the Andes (Fig. 1B). Palaeontolodcal remains suggest, howc\rcr, that the genus l~i;ugnn may have ori
278

,J. C:. \VHI
for withers height (Thomas, 19 17). Howcvrr, a recent study of 50 malt and 50 female V.v. mensalis (1 .5 to 6.5 years old) from Pampa Galcras, Peru, reports average withers height measurements of 86.5 cm for fcmalcs and 90.1 cm fol the males (Paucar eI al., 1984). No &arty definccl geographic separation csists between the two proposed ~icuiia subspecies ancl many authors i
<:.lhll~I.II)

~\‘oI.t”I‘IoN

279

SJun,+us ( 197 I ) calculated a total of bctwcrn 5000 and 10 000 in Peru with anothrr 2000 li\,ing in Bolivia, Ar,grntina and Chile. In 1982, 1.5 years after the cstahlishmcnt of a protection programmc, the Peruvian population had risen to 62 000 (Franklin, 1982), and in 199 I, the total Andcan population reached 92 882 (IVhcclrr, 199 1). In Peru, conservation cKorts at the Pampa Galcras rcscrvc began in 1967 with 1753 \+culias. During the first years, an unrspcctcd annual population increase of 2 1”/0 cvas rr~gistrrccl at Pampa Galcras (Sinchez, 1984). This high growth rate has subscqucntly hccn rcpcatcd when Chilean and Bolivian reserves wcrc cstablishcd (Rodriguez & Torrcs, 198 1; Rahinovitch, Hcrnandez & Cajal, 1985). In 1978m 79 howcvcr, the Pampa Galcras population reached a crisis brought on by a prolonged drought, overgrazing and over-population, and a ncLgativc I I .28’%1 growth rate was rc
ORIGIN

01:

THE

IX~hll:S’I‘IC:

I:ORhlS

To date the earliest cvidcncc of camelid domestication comes from archacoloQgical sites located between 4000 and 4900 m elevation, in the puna ecosystem of the Peruvian Andes. Both guanaco (L.g. racsilensir) and vicufia (V.V. mendis) have inhahitcd this tundra environment for approximately 12 000 years and, together with the huemul deer Hi@ocamelus nntisensti (d’OrbiLgny, 1834), were the primary prey of early human hunters. Fauna1 materials from archaeolo
‘XII

J. (:. \\‘HIXI.I~I<

taken place in this arca o\.cr tltc last IO 000 )~ars (\‘an drr Hammcn C% Noldus, 1986). Fi\rc seasons of csca\.ation a( ‘l’clat~maciia~~ Rockshcltc*r !I.a\~allrc PI nl., 1986) rc\~nlcd a 8200 ycnr long occupational sccluctic~ alld rcco\Wccl mot-c than one metric ton of animal l~otics Ii-om the prcccramic lc\.cls. ;\rcliacozoolo~ic~~l analysis of thrsc materials producc’d ~\iclrncc 01‘ ii sliili li-om gcticralizd hunting of ,pa11ac0, \icutia and hucmul deer 9000 7200 years ago. (0 spccializccl hunting of guanaco and \icufia al~l~rosiinatcl~. 7200 ~4X)OOyears a,go. tltcn to control of early domestic alpacas and Ilalllas I,). 6000 5500 )‘CWs a,qo, allcl finally, to tl1r cstal~lislitnctit of a l~rccl~~iiiitiatcl~~ hcrditig ccononi)~ l~cgititiitig 5500 years ago j\\‘licclct-, 1986, util~ul~lislicd data). It has not hccn possiMc to detcrtninc if thcsc shifts wcrc associated \\.itli I~ocl~~size rcductioti as has 11ccti documented for other dotncstic u~i~ulatcs lxrausc slxcics spccilic cliaractcrs hi separating postcranial l~otics arc lacking. Instcad, dctcrniinatiott of car1~. cati~clicl domestication at Trlarmachay is hascd upon an iticrcasc in dir hcc~urtic~~ of‘ both catnrlid and neonatal catnclid rcmaitis, togcthcr \\.itli cliatig~-cs in clcntal morphology. During the preccratnic period, 9000 to 1800 years ago, camclid rctnains ,gradually incrcascd from 64.7’%1 to 88.G”A~ of‘ the litunal assct~~l~lagc, while drcr remains diminished fi-om 34.2’s) to 9.2’%) of thr tool (\\‘hcclcr, 1986). This shift was not caused 1,). dccrcasccl a\.ailabilit). of deer in the zone, hut rather, by a change in animal utilization patterns Ii-om ~gcncralizccl to spccializcd hunting and c1.rritual domestication of the camclids. Bctwrcn 9000 and 6000 years ago, catnclid remains incrcasccl frotn 64.7”% to 8 1.7”/0 of thr total fauna1 sample, \vith- .just o\‘cr 011~ thircl (35.3’%, to 37.1 I%)) of the l~ones coming frotn fetal or neonatal animals (\\‘hcclcr, 1986). These fi
(:.\.\II~;I.II)

I:\‘ol.I”I’los

‘HI

:\ltho~~gh it is not al\\~)~ possible to distinguish bc.t\\.cc.n llir Imncs of a terminal clc\.cbn ancl on(’ half month Ii)&11 camclicl ancl those of a nconatc, tooth \\‘c‘ar studies inclic.atc that lhr majorit). of ‘I’clarmachay spccinicns elating from (iOO0 to 3800 )x21-s ago \rcrc neonatal, \\.hcrcas those horn the carlic1 Ic\.cls \\‘crc primaril], li~clal, presumably taken ir, II/WI throiigh thr hunting of pregnant fcmalcs (\\‘hcrlcr, 1986). ‘I‘his shift fi-om l~r~clominantl~. foctal to nc.oliatal remains coincides \vith the significant incrcasc in frcclucncy of Ib~tal/nc~oli;ital rcmailis drscribccl abo\,r, ancl permits the hypothesis that mortalit). inducccl 1,). discasc rather than 1,). intentional butchcry \vas the cause. :1clclitional support li)r this intrll>rctation c‘omcs l-r0111 the stucly of lmne distribution across thr 6000 to 3800 ?.car olcl li\Gig floors \vhich inclicates that nc\vborn camclicls \vcrc broL+$it into the shcltcr \\holc ancl proccsscd for consuniption. The resultant pattern is \.c~ similar to that crcatecl 1,)~contemporary traditional hrrclcrs \\.lio utilize clcacl llama ancl alpaca nconatcs fill- Ibocl. hleat produced hy tlic olicn massi\*c die-ofl‘ of camclid nconatcs dors not now, and apparcnlly did not then, ,go to ~vaslc. Idrntification of the spccics Lvhich \vas brought unclcr domestication at Trlarniarha)~ is basccl upon incisor morpholo,~. Prior to clomcstication (9000 to 6000 BP) it is cstimatccl that nine \.icluia \\.crc hunted fixc\uy gumaco based on incisor lyJ>c and frcqucncy. Vicufias lia\rc rootless hypscloclont parallclsiclcd pcrmancnt incisors \vith cnamcl co\prring the cntirc labial surface, and root-li)rmin
282

J, C:. \YHIIKI.I-R A.

GUANACO

VICUtiA

LLAMA

ALPACA

WAR1

0.

VICUtiA

GUANACO NOT LLAMA

C.

DOMESTICATED

ALPACA

GUANACO

I

LLAMA

VICUI;IA

Rohrs, 1957; Fallet, 1961; Zeuner, 1963; Herre & Thicde, 1965; Herre & Rohrs, 1973; Bates, 1975; Pires-Ferreira, 198 l/82; Kleinschmidt et al., 1986; Kruska, 1982; Jurgens et al., 1988; and Piccinini et al., 1990). In the 1930s Lopez Aranguren (1930) and Cabrera (1932) suggested that llama and alpaca evolved from presently extinct wild precursors, based on the discovery of 2 Myr Plio-Pleistocene L. glama, L. paces, L. guanicoe and V. vicugna fossils in Argentina, and that the guanaco and vicuria were never domesticated. This position is no longer considered a possible alternative. Finally, Hemmer (1975, 1983, 1990) attributes llama ancestry to the guanaco, but has deduced on the basis of shared morphological and behavioural traits that the alpaca originated from hybridization between the llama and vicuna (Fig. 2C). Conclusions about llama and alpaca ancestry have, in large part, been based upon morphological changes produced by the domestication process. During the 195Os, Herre and Rohrs (Herre, 1952, 1953, 1976; Hcrre & Rohrs, 1973; Rohrs, 1957) examined alterations in the mesotympanal area of the skull related to a decrease in llama and alpaca hearing acuity, and reported an overall reduction in cranial capacity of both domestic species relative to the guanaco.

~:r\hll~I.II~

I:VoI.U’I‘IoN

283

In contrast, they fomld the vicufia cranium to he the smallest of all living Lamini, and, based on the premise that domestic animals arc smaller than their ancestors, concluded that this species was never brought under human control. Herrc and Riihrs consider the llama and alpaca to be “races of the same domrstic species hrcd for different purposes” (Herrc, 1976: 26). Research on the relationship of brain size relative to body size by Kruska (1982) also found the vicuiia to he smaller than alpaca and llama, which in turn were smaller than the
,J. C. \\‘HI:RI.I~K

284

r/ nl. (1986), Jurgrns guanaco from Hannovcr Zoo, Germany, led Klcinschmidt et al. (1988), and Piccinini et a/. (1990) to the conclusion that the vicmia was never domesticated. However, rarlicr rcscarch on blood and rn~~sclr samples from llama, alpaca, vicmia, guanaco and alpaca x vicmia hybrids at Santiago Zoo (Cappuro & Silva, 1960) indicated a Ilamaguannco and alpaca-cicutia subdivision, as have more recent data from rihosomal gcncs (Semorilc et al., in press). Other researchers utilizing immunolo~gical, clcctrophorctic analysis and protein sequencing have found it impossihlc to draw conclusions about llama and alpaca ancestry (Miller, Hollander & Franklin, 1985; Pcncdo ef al., 1988). Cytogenetic studies (Capanna & Civitclli, 1965; Taylor et nl., 1968; Larramendy et al., 1984; Gentz & Yates, 1986) indicate that all four species of the South American Camelidae have the same 2n = 74 karyotypc, but information on molecular biology is limited. Vidal Rioja et al. (1987) and Saluda-Gor~gul, Jaworski & Greger (1990) have examined satellite DNA, and research analysing the full mitochondrial cytochrome b gcnc sequence in all six Camelidae has documented hybridization among the domestic South American camelids (Stanlcy et al., 1994). Recent studies of the fibrc from mummified ninth and tenth century llamas and alpacas suggests that post-conquest hybridization has modified the genetic makeup of living populations (Wheeler et nl., 1992), a fact which may well explain the diversity of conclusions about their ancestry.

THE

DORlESI‘IC

SOUTH

/UiIEKICAN

The llama Lama glumu

C:ARIEI,ID,\E

(Linnaeus,

1758)

The llama is the largest of the domestic South American camelids and resembles its ancestor in almost all aspects of morphology and behaviour. Like ‘the guanaco, the llama has adapted to a wide range of environments (Fig. 1C). After domestication in the Peruvian puna between 7000 and GO00 years ago (Wheeler, 1984, 1991; Wing, 1977, 1986), 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, 1986; Hesse, 1982; Dransart, 199 la). Some 2400 years later they were being bred on the north coast of Peru (Shimada & Shimada, 1985) and in Ecuador (Wing, 1986; Stahl, 1988; Miller & Gill, 1990). Although it is often assumed that the Lake Titicaca region was also a centre of llama domestication, relevant data are lacking from early archaeological sites in Bolivia (Browman, 1989). In northwestern Argentina, a single cranium of L. glumu has been dated to 3400 years BP with stronger evidence for herding at 1450 BP (Yacobaccio & Madero, 1992; Reigadas, 1992), and it is thought that domestication may have occurred independently in both this region (ibid.) and northern Chile (Hesse, 1982). Shortly thereafter, 900-1000 BP, evidence of llama rearing has been recovered at 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) 11ama distribution reached its furthermost expansion as pack trains accompanied the royal armies to southern Colombia and central Chile. It is impossible to estimate the size of this preconquest llama population, but it clearly must have exceeded present numbers for early Spanish administrative documents record the virtual disappearance of these animals

(:.\.\11~;1.11)

I~\‘oI.t”I‘Ios

LLAMA

ALPACA

PRECONQUEST

BREEDS

COARSE FIBRE FINE FIBRE

/

Q’ AitA

EXTRA

POSTCONQUEST

FINE FINE

FIBRE FIBRE

PHENOTYPES

WAR1

SURI

within a century of contact (Florcs Ochoa, 1977). In recent years the llama population has remained relatively stable, totalling 3 776 793 in 1991 (JVheeler, 199 1). Bccausc Andcan civilization was nonliterate, knowledge of pre-Spanish llama and alpaca herding practices must be reconstructed from archaeological remains. The recent discovery of 900-l 000 year old naturally desiccated llamas and alpacas at El Yaral, an archaeoloLgical site in the Moque
286

.J. C. \YHEEI,ER

relative to contemporary animals (Wheeler et al., submitted). The growth curve and live weights of the llamas from El Yaral, and other Chiribaya culture sites of the same region, are very similar to those of contemporary llamas raised in the puna, and their age at death reflects controlled stockrcaring with elimination of undesirable animals from the herd (Wheclcr, in press). Prior to discovery of the El Yaral mummies, our most detailed data on preconquest camelid breeding practices came from written documents of the colonial period. These records describe the use of llamas as pack animals for the Inca army, but make no mention of fine fibre producing llamas. This may be due to the general failure of the early Spanish writers to distinguish between llamas and alpacas, as well as their special interest in pack animals for use in transporting ore. Despite their European perspective, these documents do provide details about Inca husbandry. Expansive state and shrine herds were managed by the llama camayoc, members of a hereditary caste of herding specialists, and emphasis was placed on breeding pure brown, black and white animals for sacrifice to specific deities, as well as on quality fibre production for the state controlled textile industry (Murra, 1965, 1975, 1978; Brotherston, 1989). Detailed data on size and colour of flocks were kept utilizing the quipu, a memory assistance device made of knotted camelid fibre cords. Communally and individually owned herds also existed. Native Andean stockrearing was largely destroyed by the arrival of the Spanish. Within little more than a century of the conquest in 1532, administrative documents record the disappearance of approximately 90% of the domestic camelids (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, cattle and pigs. In the puna this process was somewhat slower because both the Spanish and their livestock found the harsh climate and extreme elevation inhospitable. This region became a refuge for native livestock and herders, and their descendants continue to inhabit the same marginal lands today. The prolonged Spanish civil wars and heavy tribute levies, paid either in domestic camelids or in money obtained from their sale, resulted in depletion of the herds. Introduced livestock diseases may also have played an important role in this process. By 165 1, llamas and alpacas had practically disappeared even in the Lake Titicaca basin (Flora Ochoa, 1982), the former heartland of their distribution (Murra, 1975). The impact of such catastrophic mortality upon camelid 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, but the most important production zone is located between 11” and 2 1”s latitude at elevations of 3800-5000 meters above sea level. The name llama comes from Quechua (F’lores Ochoa, 1988), and it is known as qawra by Aymara speakers (Dransart, 1991 b). Although specific llama breeds do not exist, at least three varieties of llama are recognized. Most llamas in Peru, 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. To the south, especially in Argentina, the ‘woolly’ llama is more common and has a greater density of fibre on the body which extends forward between

CAhlI:I.ID

WOILJ’I‘ION

287

the ears and grows from inside the ears but is absent on the legs. The woolly type is known as ch’aku in Qucchua (Flores Ochoa, 1988) and t’awrani in Aymara (Dransart, 199 1b), while the nonwoolly type is called q’ara in both languages (ibid.). In both arcas llamas with intermediate phenotypes are also recognized. Recent research on the fibre characteristics of Argentine llamas has identified the existence of seven distinct fibre types in the population (Frank & Wehbc, 1994), raising the possibility that more than three varieties of llama exist. A different classification has been proposed by Cardozo (1954: 61) who divides llamas into brachymorphic (round, short profile, abundant fibre) and dolichomorphic (narrow, elongated profile, sparse fibre). The vast majority of llamas are held by traditional Andean pastoralists who utilize elaborate classification hierarchies based on colour, fibre and conformation characteristics to describe their animals. The existence of these systems among both Quechua (Flores Ochoa, 1988) and Aymara (Dransart, 1991b) speaking herders suggests that earlier management strategies may have been directed at producing animals with specific fibre types, but it is not clear to what extent selection is made for these characteristics today. Contemporary llamas lack the phenotypic uniformity associated with true breeds, 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. Pelage coloration varies from white to black and brown passing through all intermediate shades with a tendency to spots and irregular colour patterns, and llamas with wild guanaco coloration occur. Fleece quality is uneven, with wide variation in fibre diameter and a strong tendency to hairiness, ranging from 32.5f 17.9 /tm (0) to 35.5+ 17.8 pm (3) for coarse ‘nonwoolly’ q’aras, 30.5 + 18.5 pm (9) to 30.5 & 17.9 pm (8) for intermediates, and 27.0f 15.6 pm (0) to 29.1 & 12.7 pm (& for ‘woolly’ cha’kus (Vidal, 1967). F or this reason, the primary value of the llama presently lies in its use as a pack animal rather than as a fibre producer. The variability of present day llama fibre is related to an increase in hairs and general coarsening of the fleece, which probably began at the time of the Spanish conquest. Increased hairiness is produced by lack of controlled breeding, 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. 3). The alpaca Lama paces (Linnaeus,

1758)

The alpaca is smaller than the llama and resembles the vicuiia in many aspects of morphology and social organization. Although the Lake Titicaca basin of southern Peru and Bolivia has long been considered the focus of alpaca domestication, archaeological evidence is not presently available to evaluate this hypothesis (Brown-tan, 1989). Nevertheless, excavations in the central Peruvian puna have placed its origins between 7000 and 6000 years ago (Wheeler, 1984, 1986), and it was from this region that the alpaca was subsequently moved to lower elevation interAndean valleys 3800 years ago (Wing, 1972; Sh imada, 1985). Evidence of alpaca rearing at coastal sites in southern Peru dates from 900 to 1000 years ago (Wheeler et al., 1992; Wheeler, in press; Wheeler et al., submitted) (Fig. 1D). P recolumbian alpaca remains have not been reported in the faunal materials from archaeological sites in Chile

288

J. C. \VHIXI.l.X

(Dransart, 199 1a & b; HCSSC, 1982), Argentina (Elkin, personal communication) or Ecuador (Miller & Gill, 1990), although they arc found in limited numbers in these re
(::\1\11~1.11~~VOI.L”I‘ION

289

hairiness, and arc of UI~C\TII quality. Some coats containing up to 40% hair have lx~n rcportcd for both living varictics, and considerable variation is rcportcd in published statistics on fibre diamctcr. The ori
HI'RRIDI%:l'I‘ION

The guanaco, vicufia, llama and alpaca all possess the same 2n = 74 karyotype (Capanna & Civatelli, 1965; Taylor e/ nl., 1968) and can, under human influcncc, product fertile hybrids (Gray, 1954). Preliminary research on the cytochrome b gent scqucnce has found no evidence of hybridization between
290

J. C.

\\‘HEfI,I:R

generation of alpaca x vicmia hybrids is lacking, and much remains to be done before its potential as a fine fibre producer can be evaluated. The possibility that feral llamas and alpacas exist and might have crossed with wild camelids has not been fully explored. In 1534, Xercz obscrvcd that domestic llamas were sometimes so numerous some escaped to the wild, and in 1555 Zarate recorded that once each year some llamas were rclcased into the wild as an offering to the gods (hilurra, 1978). It is unclear, however, if feral populations existed at that time. The current consensus of opinion in the central Andean region is that no such populations exist today. Even so, MacDonagh (1940) has described a
I;U’I‘URE

The present status of the South American camelids is the product of a largely unknown past. To name but two historic factors, the potential influence of genetic bottlenecks and/or hybridization in the formation of contemporary guanaco, vicuiia, llama and alpaca populations have never been fully investigated, although there is evidence to suggest that they may have played an important role. The most basic questions concerning genetic variation and the systematic classification of presumed
Akimushkin J. 1971. ~&mciones. Buenos Aires: Editorial Cartaga. Auto&us 0. 1922. .Sknnmeqschichfe der Hau&x Jcna. Bates M. 1975. Siidameriko-Flora und Fauna. Reinbck: Rowchlt. Benavente MA. 1985. ReHcxioncs en torno al procrso de domcsticacihn

de 10s camelidos rn 10s vattcs drl de Chile. Bolt!in de1 Mueo Regimml Amucania 2: 37 52. Bohlken H. 1960. Remark on d~e stomach and the systematic posilion of die Tytopoda. Proneding UJ‘ /hi .+ologicol Sorie~v OJ- London 134: 207-2 15. Brehm A. 1916. Dh Sbge&re. Leipzig, Bihiographilches Institut. Brotherston G. 1989. Andean Pasloralism and Inca ideology. In: Glutton-Brork J, rd. 77~ walkkg /n&r pok-nu oJ domes&bon, parb~ro~bn and predokm. London: Unwin Hyman Ltd, 240 255. Browman DL. 1989. Origins and devclopmcn~ or Andcan pa5roratism: an overview of the pas1 6000 yrars. In: Clwon-Brock J, cd. 77~ &king larder polknu of dome&o&m, pnrb~r&m and pred&x London: Unwin Hyman Ltd, 256-268. centro

y sur

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Rodriguez R, Torres H. 1981. .\le~o~olo,~~~z rr.w~rlo pnn d&wrinm ICI poblnri~irr de t~rru~ia~ ( IYrrr~,~rrn z*rrr!er,ni cn t-1 I’arquc .4brionnl Inurn. A&a: Corporarifm National Forestal. RShrs M. 1957. <1kologisrhc Brobachrungcn an wildlcbrndcn ‘l‘vlotx&n , Sildanicrikas. I~~ll~nrt~lrrr~~ &r Ih~khen ~oohpkhtn (h~rlhd~~~ji .538 5.54. Saluda-Gorgul A, Jaworski J, Greger J. 1990. Nurtrotidr scqurnw of sa~cllilc I and II DNA liom Alpaca (.!mo pnros) gr”““le. ilrla Biorhimif-0 f’olo~~ica 37(2): 283 297. Stichez E. 1984. Sobrc poblaciim y nrwsidad clr estraccihn dc vicmias VII Pampa Galwas. In: \‘ittigrrI:, cd. La Gtin. I.ima: Editorial 1.0s Pines. I2 18. Semorile LC, Crisci JV, Vidal-Bioja L. in press. Rcstrirtion sites variation 01‘ rhc 1-ibosomat DNA in Camelidar. &n&n 90. Shimada M. 1985. Cominui&s and rhangcs in patrcrn ot ti~unal rcsourw utilizarion: lixmarivc d~rough Cajamarca pcriocls. In: Trrada K. Onuki K, rds. EwototiurtJ a/ Ifulmtalwm in t/w G~~~morco I id/T. I’rru. 1979. Tokyo: Uni\wsity of ‘I‘okyo Prrss. 303 336. Shimada M, Shimada I. 1985. Prchisroric llama brcrding and herding on lhc north roasl of Prru. Anr&an .4n/iqrri!r 50( 1): 3 26. Stahl PW. 1988. Prehistoric camclids in the lowlands of wcswrn I:cuador. ,7ow71dcf .4rr/meo/u,~irn/ .S&wr 15: 3.5.5 365. Stanley HF, Kadwell M, Wheeler JC. 1994. hlotcwtar cvolu~ion or lhc Igmity Camclidar A mimchnndrial DNA study. I’rorerrfu~.r of /ire Rovnl Socit-!y London U 256: I Ii. Steinbacher G. 1953. %ur Abstammung drs Alpakka, Intttn pnrw (I.imw, I 7.58). .S~il~~~ci~rkrrrrd~i~/~[ir/lP ;Ili/Mrt~,~en l(2): 78 -79. Taylor KM, Hungerford DA, Snyder BL, Ulmer FA Jr. 1968. Uniformity of karyoL)lxs in lh( Camelidae. Qlogetrb 7: 8- 15. Thomas 0. 1891. Notes 011 some un,qutaw mammals. I’ruc~~~~rqr OJ l/w +xhgiml .So:orir!v CI/ Lortdoon. 384 389. Thomas 0. 1917. Prciiminary dia