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The Llama

Jane C. Wheeler
Facultad de Medicina Veterinaria
Universidad Nacional Mayor de San Marcos
Apartado 41-0068
Lima 41, PERU

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 (Figure 3C).  After domestication in the Peruvian puna between 7,000 and 6,000 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 3,800 years ago (Wing, 1986; Hesse, 1982; Dransart, 1991a).  Some 2,400 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 center of llama domestication, relevant data are lacking from early archaeological sites in Bolivia (Browman, 1989).  In northwestern Argentina, a single cranium of L. 21ama has been dated to 3,400 years BP with stronger evidence for herding at 1,450 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, 900-1,000 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) llama 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 within a century of contact (Flores Ochoa, 1977).  In recent years the llama population has remained relatively stable, totalling 3,776,793 in 1991 (Wheeler, 1991).

Because Andean civilization was nonliterate, knowledge of pre-Spanish llama and alpaca herding practices must be reconstructed from archaeological remains.  The recent discovery of 900-1000 year old naturally desiccated llamas and alpacas at El Yaral, an archaeological site in the Moquegua valley of southern Peru (Rice, 1993), has provided a first view of preconquest breeds (Wheeler et aL, 1992; Wheeler et al., submitted).  Associated with the pre-Inca Chiribaya culture, these animals had been sacrificed by a blow between the ears and immediately buried beneath house floors where they became naturally mummified due to the extreme aridity of the environment.

Research on the physical appearance of the El Yaral llamas, as well as analysis of skin and fiber samples taken at I I different locations across the body, revealed the possible existence of both a fine fiber and a coarse fiber breed (Wheeler et aL, 1992; Wheeler et al., submitted)(Figure 6). Average fleece diameter of the former was found to be 22.2 with a between sample standard deviation of 1.8 Jim, compared to 32.7 (sd ± 4.2) iLm for the latter, based on the measurement of up to 1,600 fibers per animal. 

The reduction of both fiber diameter and variation in the fine fiber llama fleece was certainly produced by selective breeding for a single-coat through modification of the primary hair to resemble secondary undercoat fiber.  The uniform coloration and fineness, as well as the absence of visible hairs in the El Yaral fine llama fleeces are ideally suited for textile production, and contrast markedly with the multi-colored double-coat of the coarse fiber breed.

An additional evidence of specialized breeding is the accelerated fiber growth rate recorded for El Yaral fine llamas 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 stockrearing with elimination of undesirable animals from the herd (Wheeler, 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 fiber 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 fiber 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 fiber 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 1651, llamas and alpacas had practically disappeared even in the Lake Titicaca basin (Flores 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 (I' 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 I I' and 2 1 ' S latitude at elevations of 3,800-5,000 meters above sea level.
The name llama comes from Quechua (Flores Ochoa, 1988), and it is known as qawra by Aymara speakers (Dransart, 1991b).  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 linonwoolly" phenotype characterized by sparse fiber growth on the body and the absence of fiber on the face and legs.  To the south, especially in Argentina, the "woolly" llama is more common and has a greater density of fiber on the body which extends forward between the ears and grows from inside the ears but is absent on the legs. 

The woolly type is known as ch'alcu in Quechua (Flores Ochoa, 1988) and t'awrani in Aymara (Dransart, 1991b), while the nonwoolly type is called q'ara in both languages (ibid.). In both areas 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 & Wehbe, 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 fiber) and dolichomorphic (narrow, elongated profile, sparse fiber).

The vast majority of llamas are held by traditional Andean pastoralists who utilize elaborate classification hierarchies based on colour, fiber 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 fiber 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 fiber animals. 

Pelage coloration varies from white to black and brown passing through all intermediate shades with a tendency to spots and irregular color patterns, and llamas with wild guanaco coloration occur.  Fleece quality is uneven, with wide variation in fiber diameter and a strong tendency to hairiness, ranging from 32.5 ± 17.9 lkm (@) to 35.5 ± 17.8 14m (@ for coarse "nonwoolly" q'aras, 30.5 ± 18.5 Jim (@) to 30.5 ± 17.9 itm (d) for intermediates, and 27.0 ± 15.6 ikm (@) to 29.1 ± 12.7 itm (CO for"woolly" cha'kus (Vidal, 1967).  For this reason, the primary value of the llama presently lies in its use as a pack animal rather than as a fiber producer. 

The variability of present day llama fiber 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 (Figure 6).

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