Evolution and Origin of the Domestic Camelids

Located 170 km (1,054 miles) northeast of Lima, Peru (11_ 11’S latitude and 75_ 52’W longitude), at 4,420 m (14,498 feet) above sea level, Telarmachay is situated near the absolute upper limits of crop growth potential. Mean annual temperature is 4.8_C (40.6_F), with an average daily variation of greater than 20_C (68_F) and frost occurring 330 nights of the year. Annual precipitation averages from 500 to 1,000 mm (20 to 40 inches) and is normally restricted to the months from November to March, although the timing is irregular and unpredictable, and extended periods of drought occur. No agriculture is practiced in the area today, and grazing ungulates represent the most reliable food resource. This is due to their mobility during periods of drought and their ability to convert the dry ligneous puna grasses into a source of stored protein that can be utilized for human consumption. Palaeoclimatological data indicate that no significant climatic changes have taken place in this area over the last 10,000 years (Van der Hammen and Noldus 1986).

Five seasons of excavation at Telarmachay Rockshelter (Lavallee et al. 1986) revealed a 8,200-year-long occupational sequence and recovered more than 1 metric ton (1.1 ton) of animal bones from the preceramic levels. Archaeozoological analysis of these materials produced evidence of a shift from generalized hunting of guanaco, vicuña, and huemul deer 9,000 to 7,200 years ago, to specialized hunting of guanaco and vicuña approximately 7,200 to 6,000 years ago, then to control of early domestic alpacas and llamas by 6,000 to 5,500 years ago, and, finally, to the establishment of a predominately herding economy beginning 5,500 years ago (Wheeler 1986, unpublished data). It has not been possible to determine if these shifts were associated with body size reduction, as has been documented for other domestic ungulates, because species-specific characters for separating postcranial bones are lacking. Instead, determination of early camelid domestication at Telarmachay is based upon an increase in the frequency of both camelid and neonatal camelid remains, together with changes in dental morphology. During the preceramic period, 9,000 to 1,800 years ago, camelid remains gradually increased from 64.7 percent to 88.6 percent of the faunal assemblage, while deer remains diminished from 34.2 percent to 9.2 percent 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 generalized to specialized hunting and eventual domestication of the camelids.

Between 9,000 and 6,000 years ago, camelid remains increased from 64.7 percent to 81.7 percent of the total faunal sample, with just over one-third (35.3 percent to 37.1 percent) of the bones coming from fetal or neonatal animals (Wheeler 1986). These figures are consistent with a hunting economy because between 35 percent and 40 percent of animals in contemporary guanaco and vicuña populations fall within this category (Franklin 1978, personal communication). Thus, the ever-greater dependence upon camelids in the diet during this period suggests increasing specialization in guanaco and vicuña hunting.

Around 6,000 years ago, however, the frequency of fetal and neonatal camelids increased markedly to 56.8 percent and continued to rise until it reached 73.0 percent of all camelid remains in the deposits dated to 3,800 years ago (Wheeler 1986). These figures suggest either the development of specialized hunting of neonates, an economically unviable strategy, or the appearance of other mortality-inducing factors in the environment. They far exceed expected frequencies for both the fetal/neonatal age group and the natural (that is, no human hunting) mortality rates of 4.5 (Raedeke 1979, 199) to 30 percent (Franklin, 1978, 42) that have been recorded for the guanaco and vicuña, but closely correspond with mortality rates experienced by llama and alpaca breeders today.

At present, up to 70 percent of each year’s young may be lost before two months of age owing, in part, to failure of passive immune transfer (Garmendia et al. 1987) with resulting mortality from Clostridium perfringens Type A enterotoxemia and other pathogens (Leguía 1991; Ramírez 1991). The epizootic nature of enterotoxemia is to some extent controlled by climatic conditions that permit sporulation of the bacteria, as well as by the presence of a critical number of captive or domestic animals. In the Andes, outbreaks of enterotoxemia are associated with unsanitary corraling practices during the wet- season birth period. Similar epidemics are not known to occur in the wild camelids.

Although it is not always possible to distinguish between the bones of a terminal eleven-and-one-half-month-old fetal camelid and those of a neonate, tooth wear studies indicate that the majority of Telarmachay specimens dating from 6,000 to 3,800 years ago were neonatal, whereas those from the earlier levels were primarily fetal, presumably taken in utero through the hunting of pregnant females (Wheeler 1986). This shift from predominantly fetal to neonatal remains coincides with the significant increase in frequency of fetal/neonatal remains described above and permits the hypothesis that mortality induced by disease rather than by intentional butchery was the cause. Additional support for this interpretation comes from the study of bone distribution across the 6,000- to 3,800-year-old living floors, which indicates that newborn camelids were brought into the shelter whole and processed for consumption. The resultant pattern is very similar to that created by contemporary traditional herders who utilize dead llama and alpaca neonates for food. Meat produced by the often massive die-off of camelid neonates does not now, and apparently did not then, go to waste.

Identification of the species that was brought under domestication at Telarmachay is based upon incisor morphology. Prior to domestication (9000 to 6000 b.p. [before the present]) it is estimated that nine vicuña were hunted for every guanaco based on incisor type and frequency. Vicuñas have rootless hypselodont parallel-sided permanent incisors with enamel covering the entire labial surface, and root-forming deciduous incisors with enamel covering the upper labial surface only (Miller 1924; Wheeler 1982, 1991). Guanacos have rooted deciduous and permanent spatulate incisors with an enamel-covered crown (Miller 1924). By 6000 b.p., however, the remains of permanent incisors with the same morphology as deciduous vicuña incisors appear in the Telarmachay deposits (Wheeler 1982, 1991, unpublished data). These permanent teeth match the dentition of many extant Peruvian alpacas in which both the deciduous and permanent incisors are root forming and parallel sided, with enamel covering only the upper labial surface (Wheeler 1991, unpublished data). Although contemporary alpacas with spatulate llama incisors have been reported by Kent (1982), it is unclear if these are hybrids. The evidence from Telarmachay suggests an ancestral relationship, which may explain the apparent retention of juvenile vicuña dental traits in the adult alpaca. It cannot be determined if animals with llama-type incisors also appeared in the 6000 b.p. deposits, since these are indistinguishable from guanaco incisors, but the presence of both large and small neonates suggests that this may have been the case.

In contrast to the data from Telarmachay and other Andean archaeological sites that indicate that the llama is descended from the guanaco and the alpaca from the vicuña (Figure 1A), other researchers have come to different conclusions 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 the vicuña, an opinion subsequently supported by Ledger (1860), Darwin (1868), Antonius (1922), Faige (1929), Krumbiegel (1944, 1952), Steinbacher (1953), Frechkop (1955), Capurro and Silva (1960), Akimushkin (1971), and Semorile, Crisci, and Vidal-Rioja (in press). Other authors have concluded that both domestic camelids descend from the guanaco, and the vicuña was never domesticated (Figure 1B) (Thomas 1891; Peterson 1904; Hilzheimer 1913; Lönnberg 1913; Brehm 1916; Cook 1925; Weber 1928; Herre 1952, 1953, 1976, 1982; Röhrs 1957; Fallet 1961; Zeuner 1963; Herre and Thiede 1965; Herre and Röhrs 1973; Bates 1975; Pires-Ferreira 1981/82; Kleinschmidt et al. 1986; Kruska 1982; Jürgens et al., 1988; and Piccinini et al. 1990). In the 1930s, López Aranguren (1930) and Cabrera (1932) suggested that llama and alpaca evolved from presently extinct wild precursors, based on the discovery of 2 Myr [million years ago] Plio-Pleistocene L. glama, L. pacos, L. guanicoe, and V. vicugna fossils in Argentina, and that the guanaco and vicuña 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 behavioral traits that the alpaca originated from hybridization between the llama and vicuña (Figure 1C).

Conclusions about llama and alpaca ancestry have, in large part, been based upon morphological changes produced by the domestication process. During the 1950s, Herre and Röhrs (Herre 1952, 1953, 1976; Herre and Röhrs 1973; Röhrs 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.

In contrast, they found the vicuña cranium to be the smallest of all living Lamini and, based on the premise that domestic animals are smaller than their ancestors, concluded that this species was never brought under human control. Herre and Röhrs consider the llama and alpaca to be "races of the same domestic species bred for different purposes" (Herre 1976, 26). Research on the relationship of brain size relative to body size by Kruska (1982) also found the vicuña to be smaller than the alpaca and llama, which in turn were smaller than the guanaco, suggesting that the latter is the only ancestral form. Nevertheless, papers by Jerison (1971) and Hemmer (1990) report the ratio of alpaca brain size to body size to be smaller than in the vicuña, permitting a different conclusion about origins of the domestic forms. These contradictory data on size reduction are almost certainly a product of sampling, as neither subspecific variation in the wild forms nor the possibility of hybridization between the domestic animals were considered in any of the studies.

Based on the study of pelage characteristics (skin thickness, follicle structure, secondary/primary ratio, fiber length and diameter, coloration) in living camelids, Fallet (1961) found the llama to be an intermediate evolutionary stage between the wild guanaco and the specialized fiber-producing alpaca, and concluded that the absence of transitional characteristics between vicuña and alpaca fleeces eliminates the former from consideration as an ancestral form. This deduction is, in part, based on the assumption that llamas have been selected exclusively for use as pack animals while alpacas have been bred for fiber production. Nevertheless, new data on preconquest llama and alpaca breeds in Peru have revealed the prior existence of a fine-fiber-producing llama, as well as an extra-fine-fiber alpaca that is transitional between the vicuña and a second pre-Hispanic fine-fiber alpaca breed (Wheeler, Russel, and Stanley 1992; Wheeler, Russel, and Redden, submitted).

Research on camelid bahavior has produced contradictory hypotheses concerning llama and alpaca origins. Krumbiegel (1944, 1952) and Steinbacher (1953) argue that the alpaca is the domestic vicuña based on unique shared behavioral traits that are said to differ from those observed in the guanaco and llama. Hemmer, on the other hand, concludes that while some alpaca behavior patterns match those of the vicuña, others are intermediate between those of vicuña and guanaco, suggesting that "The alpaca is a mixture of both lines, [produced] by crossbreeding of captured vicuñas with the only initially available domestic animal, the llama" (1990, 63). It has also been suggested that the vicuña was never domesticated because it is more territorial than the guanaco (Franklin 1974). Nevertheless, this assumption is open to question because it is based upon study of guanacos located at the southernmost extreme of their range, where seasonal migration in response to severe climatic changes is essential for survival (Franklin 1982, 1983). Farther to the north, where vicuña and guanaco ranges overlap and llama and alpaca domestication occurred (Wheeler 1984), a more benign climate and a constant food supply permit the characteristic sedentary social organization of the vicuña (Franklin 1982, 1983). Although data concerning behavior of the guanaco in this region are lacking, it is possible that the limited sedentary territorial organization observed in some Patagonian groups plays a more important role in these less-extreme climatic conditions.

Analysis of hemoglobin amino acid sequences in vicuña, alpaca, llama, and guanaco from the Hannover Zoo, Germany, led Kleinschmidt et al. (1986), Jürgens et al. (1988), and Piccinini et al. (1990) to the conclusion that the vicuña was never domesticated. However, earlier research on blood and muscle samples from llama, alpaca, vicuña, guanaco, and alpaca x vicuña hybrids at Santiago Zoo (Cappuro and Silva 1960) indicated a llama–guanaco and alpaca–vicuña subdivision, as have more recent data from ribosomal genes (Semorile et al., in press). Other researchers utilizing immunological, electrophoretic analysis, and protein sequencing have found it impossible to draw conclusions about llama and alpaca ancestry (Miller, Hollander, and Franklin 1985; Penedo et al. 1988). Cytogenetic studies (Capanna and Civitelli 1965; Taylor et al. 1968; Larramendy et al. 1984; Gentz and Yates 1986) indicate that all four species of the South American Camelidae have the same 2n = 74 karyotype, but information on molecular biology is limited. Vidal-Rioja et al. (1987) and Saluda-Gorgul, Jaworski, and Greger (1990) have examined satellite DNA, and research analyzing the full mitochondrial cytochrome b gene sequence in all six Camelidae has documented hybridization among the domestic South American camelids (Stanley et al. 1994). Recent studies of the fiber from mummified ninth- and tenth-century llamas and alpacas suggests that postconquest hybridization has modified the genetic makeup of living populations (Wheeler et al. 1992), a fact that may well explain the diversity of conclusions about their ancestry.

THE DOMESTIC SOUTH AMERICAN CAMELIDAE

The llama Lama glama (Linnaeus 1758)

The llama is the largest of the domestic South American camelids and resembles its ancestor in almost all aspects of morphology and behavior. Like the guanaco, the llama has adapted to a wide range of environments (Figure 2C). 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 inter-Andean 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 and Shimada 1985) and in Ecuador (Wing 1986; Stahl 1988; Miller and 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. glama has been dated to 3400 b.p., with stronger evidence for herding at 1450 b.p. (Yacobaccio and 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 b.p., 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 Incan rule (1470–1532) llama distribution reached its farthest 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, totaling 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- to 1,000-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 from 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 eleven 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 3).

Average fleece diameter of the former was found to be 22.2 with a between-sample standard deviation of 1.8 µm, compared to 32.7 (SD ±4.2) µm 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 multicolored 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 color of flocks were kept by means of 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 percent of the domestic camelids (Flores Ochoa 1982) as well as 80 percent 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 (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 21_S latitude at elevations of 3,800 to 5,000 meters (about 12,460 to 16,400 feet) 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 "nonwoolly" phenotype, characterized by sparse fiber growth on the body and the absence of fiber on the face and gels. 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’aku 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 fiber characteristics of Argentine llamas has identified the existence of seven distinct fiber types in the population (Frank and 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 color, 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, respectively. 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 µm (female) to 35.5 +17.8 µm (male) for coarse "nonwoolly" q’aras, 30.5 +18.5 µm (female) to 30.5 +17.9 µm (male) for intermediates, and 27.0 +15.6 µm (female) to 29.1 +12.7 µm (male) 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 pre-Spanish llama breeds from El Yaral could account for the entire range of fleece variation observed in today’s animals (Figure 3).

The alpaca Lama pacos (Linnaeus 1758)

The alpaca is smaller than the llama and resembles the vicuña 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 (Browman 1989). Nevertheless, excavations in the central Peruvian puna have placed its origins between 7,000 and 6,000 years ago (Wheeler 1984, 1986), and it was from this region that the alpaca was subsequently moved to lower-elevation inter-Andean valleys 3,800 years ago (Wing 1972; Shimada 1985). Evidence of alpaca rearing at coastal sites in southern Peru dates from 900 to 1,000 years ago (Wheeler et al. 1992; Wheeler in press; Wheeler et al. submitted) (Figure 2D). Pre-Columbian alpaca remains have not been reported in the faunal materials from archaeological sites in Chile (Dransart 1991a, 1991b; Hesse 1982), Argentina (Elkin, personal communication), or Ecuador (Miller and Gill 1990), although they are found in limited numbers in these regions today. It is impossible to estimate the number of preconquest alpacas. Spanish documents record their rapid decimation and displacement to remote, extreme high-elevation regions of the Andes (Flores Ochoa 1977). Recent data indicate that over the last twenty-five years numbers have fallen significantly in Peru, from 3,290,000 in 1967 to 2,510,912 in 1986, and in 1991 the total Andean alpaca population was estimated to be 2,811,612 (Wheeler 1991).

Representatives of two possible preconquest alpaca breeds have been found among the 1,000-year-old El Yaral mummies. Fine-fiber and extra-fine-fiber alpacas were distinguished, based on physical appearance and average fiber diameter (1,600 fibers measured per animal). The former have fleeces averaging 23.6 (SD +1.9 µm), while the latter fleeces average 17.9 (SD +1.0 µm) (Wheeler et al. 1992; Wheeler et al. submitted). Both groups had lustrous fiber ranging from wavy to crimped and dense to very dense. Hairs were visible in three of the four animals, but were not significantly coarser than the undercoat fibers. Indeed, fiber-diameter variation both within and across the fleece was remarkably low, suggesting that rigorous breeding selection for fine-quality fiber was being practiced (Figure 3).

The Spanish conquest had a disastrous effect on both llama and alpaca populations. Massive mortality accompanied the displacement of alpaca herds from the coast, inter-Andean valleys, and most of the puna, as introduced stockrearing practices pushed the survivors into the marginal, extreme high-elevation pastures where they are found today (Flores Ochoa 1982). At present, alpaca distribution extends from approximately 8_S latitude, where they have been recently reintroduced in Cajamarca, to 20_S latitude, in the vicinity of Lake Poopo, Bolivia, with small populations located farther to the south in northern Chile and northwestern Argentina (Figure 2D).

Today, 75 percent of all alpacas, paqocha in Quechua (Flores Ochoa 1988) and allpachu in Aymara (Dransart 1991b), are held by traditional herders (Novoa 1989). Two alpaca phenotypes, known in the literature by their Quechua names as suri and huacaya, or wakaya, are recognized but these do not breed true. The suri has long straight fibers, organized in waves that fall to each side of the body in much the same manner as a Lincoln sheep, while the huacaya has shorter, crimped fibers that give it a spongy appearance similar to that of a Corriedale sheep. Occasionally animals with intermediate wool characteristics are seen, and these have been named chili by Cardozo (1954). Crosses between huacaya and huacaya produce a certain percentage of suri offspring, and crosses of suri with suri produce some huacaya offspring. Although no artificial selection is made, an estimated 90 percent of all alpacas are huacayas (Novoa 1989). The suri is not known among the Aymara herders of Chile, who refer to their huacayas simply as allpachu or alpacas (Dransart 1991b). The fleece of both phenotypes varies from white to black and brown, passing through all intermediate shades, with a greater tendency to uniform coloration than in the llama. Alpacas with wild vicuña coloration occur.

In comparison to the preconquest El Yaral alpacas, contemporary Andean huacaya and suri fleeces average 31.2 +3.8 µm (Carpio 1991) and 26.8 +6.0 µm (Von Bergen 1963) respectively, are coarser, may have a tendency to hairiness, and are of uneven quality. Some coats containing up to 40 percent hair have been reported for both living varieties, and considerable variation is reported in published statistics on fiber diameter. The origin of this degeneration almost certainly lies in the Spanish conquest, but a breakdown in controlled breeding between the fine and extra-fine El Yaral breed would not alone account for the variation observed today.

The most probable cause of coarsening and hairiness in both huacayas and suris would be through hybridization with the coarse-fiber llama breed, a not-improbable 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 crossbreeding between alpacas and llamas may have occurred since the Spanish conquest and has played a much more important role in the formation of today’s livestock than has been realized (Figure 3).

HYBRIDIZATION

The guanaco, vicuña, llama, and alpaca all possess the same 2n = 74 karyotype (Capanna and Civatelli 1965; Taylor et al. 1968) and can, under human influence, produce fertile hybrids (Gray 1954). Preliminary research on the cytochrome b gene sequence has found no evidence of hybridization between guanaco and vicuña (Stanley et al. 1994). Because this study included samples from the northern populations of both genera, the region where they are sympatric in parts of their range, the findings may possibly support Miller’s (1924) creation of the genus Vicugna.

Traditional herders recognize the existence of llama and alpaca crosses. These are referred to by the generic terms wari in Quechua (Flores Ochoa 1988) and wik’ñna in Aymara (Dransart, 1991b). These hybrids are classified as llamawari, or llama-like, and paqowari, or alpaca-like, by Quechua speakers (Flores Ochoa 1977). Aymara speaking herders use waritu and wayki for llama and alpaca phenotype hybrids, as well as the generic term wakayu for any llama ´ alpaca offspring (Dransart 1991b). First-generation crosses are easily recognized, but it is not always possible to identify hybrid animals based upon phenotype alone because it is likely that hybridization has been an ongoing process since the time of the Spanish conquest. The extent to which contemporary llama and alpaca populations have been affected by this process has not been determined, but comparison with preconquest animals suggests that it has been extensive and that breeds of fine-fiber-producing llama and alpaca have likely disappeared in the process (Wheeler et al. 1992). Hybridization has been confirmed through DNA analyses (Stanley et al. 1994) (Figure 3).

Crosses between the wild and domestic South American camelids produce fertile offspring but do not normally occur in nature. The pacovicuña, or alpaca ´ vicuña hybrid, has received considerable attention for its potential as a fine-fiber producer. Carpio et al. (1990) report fiber diameters ranging from 13.3 to 17.3 µm for five first-generation crosses, but this is said to rapidly increase in subsequent generations. The pacovicuña phenotype closely resembles that of the vicuña, although it is slightly larger and less gracile than its wild progenitor. Research on the fixation of phenotypic traits from generation to generation of alpaca ´ vicuña hybrids is lacking, and much remains to be done before its potential as a fine-fiber 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, Xerez observed that domestic llamas were sometimes so numerous some escaped to the wild, and in 1555 Zarate recorded that once each year some llamas were released into the wild as an offering to the gods (Murra 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 group of guanaco and llama hybrids living in a feral state in the province of Cordoba, Argentina. These animals were the 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 were almost entirely white. No observations on changes in body size and fiber quality were recorded. The behavior of these feral hybrids was considered to be virtually identical with that of the guanaco, and they lived and reproduced without problem.

THE FUTURE

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, vicuña, 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 guanaco and vicuña subspecies, as well as llama and alpaca breeds, remain to be answered, although such information is essential for ensuring their future. In the case of the wild camelids, we need to be sure that we are protecting all genetic variants of each species, and not just increasing the numbers of potentially genetically impoverished subgroups. In light of the increased movement of both wild and domestic camelids throughout the Andes and the beginning of exportation in 1983, there is an urgent need to identify relict populations of genetically pure pre-Columbian llama and alpaca breeds in order to ensure both their preservation and the possibility of a return to high-quality fine-fiber production.

About the Author

Jane C. Wheeler holds degrees from American University, Cambridge University, and the University of Michigan and completed postdoctoral studies at the University of Paris. She made her first trip to Peru in 1974 as a Senior Fulbright Fellow, and although she had no intention of continuing to do research there at the time, South American camelids have been the focus of her research ever since. She has published extensively on the origins of alpaca and llama domestication and the evolution of herding in the Andes. Her current research is concentrated on camelid fiber, genetics, and molecular systematics. She is managing director of Camelid Consultants International, based in Ocala, Florida, and a visiting professor at the Facultad de Medicina Veterinaria, Universidad Nacional Mayor de San Marcos, Lima, Peru.

Sources Cited

Akimushkin, J. 1971. Migraciones. Buenos Aires: Editorial Cartaga.

Antonius, O. 1922. Stammesgeschichte der Haustiere. Jena.

Bates, M. 1975. Südamerika-Flora und Fauna. Reinbek: Rowehlt.

Brehm, A. 1916. Die Säugetiere. Leipzig: Bibliographilches Institut.

Brotherston, G. 1989. Andean pastoralism and Incan ideology. In The walking larder patterns of domestication, pastoralism and predation, ed. J. Clutton-Brock, 240–55. London: Unwin Hyman Ltd.

Browman, D. L. 1989. Origins and development of Andean pastoralism: An overview of the past 6000 years. In The walking larder patterns of domestication, pastoralism and predation, ed. J. Clutton-Brock, 256–68. London: Unwin Hyman Ltd.

Cabrera, A. 1932. Sobre los camélidos fosiles y actuales de la América austral. Revista del Museo de la Plata 33:89–117.

Capanna, E., and M. V. Civitelli. 1965. The chromosomes of three species of neotropical camelidae. Mammalian Chromosomes Newsletter 17:75.

Capurro, L. F., and F. Silva. 1960. Estudios cromatagráficos y electroforéticos en camélidos Sudamericanos. Investigaciones Zoológicas Chilenas 6:49–64.

Cardozo, A. 1954. Los auquénidos. La Paz: Editorial Centenario.

Carpio, M. 1991. Aspectos tecnologicos de la fibra de los camélidos Andinos. In Producción de ruminantes menores: Alpacas, ed. C. Novoa and A. Florez, 297–359. Lima: Rerumen.

Carpio, M., C. Leyva, Z. Solari, and J. Sumar. 1990. Diameter and length variations in different types of alpaca fibre and fleeces of llama, vicuña and paco-vicuña. Schriftenreihe des Deutschen Wollforschungsinstituts 106:76–90.

Cook, O. F. 1925. Peru as a center of domestication. Journal of Heredity 16.

Darwin, C. 1868. The variation of animals and plants under domestication. London: Murray.

Dransart, P. Z. 1991a. Llamas, herders and the exploitation of raw materials in the Atacama desert. World Archaeology 22(3):304–19.

_____. 1991b. Fibre to fabric: the role of fibre in camelid economies in prehispanic and contemporary Chile. Unpublished D.Phil. thesis, Linacre College, Oxford University.

Faige, F. 1929. Haustierkunde U Haustiereucht. Leipzig.

Fallet, M. 1961. Vergleichende Untersuchungen zur Wollbildung südamerikanischer Tylopoden. Zeitschrift für Tierzüchtung und Züchtungsbiologie 75:34–56.

Flores Ochoa, J. A. 1977. Pastores de alpacas de los Andes. In Pastores de Puna, ed. J. A. Flores Ochoa, 15–52. Lima: Instituto de Estudios Peruanos.

_____. 1982. Causas que originaron la actual distribución espacial de las alpacas y llamas. In El hombre y su ambiente en los Andes Centrales, Senri Ethnological Studies, vol. 10, ed. L. Milliones and H. Tomoeda, 63–92. Osaka: National Museum of Ethnology.

_____. 1988. Clasificación y nominación de camélidos sudamericanos. In Llamichos y paqocheros, ed. J. A. Flores Ochoa, 121–37. Cuzco: Editorial Universitaria, UNSAAC.

Frank, E. N., and V. E. Wehbe. 1994. Primer informe de avance del componente "camélidos domesticos." Acuerdo Republica Argentina-Unión Europea para el Programa de Apoyo para la Mejora en la Producción de Pelos Finos de Camélidos Argentinos.

Franklin, W. L. 1974. The social behaviour of the vicuña. In The behaviour of ungulates and its relation to management, vol. 24, no. 1, ed. V. Geist and F. Walther, 477–87. IUCN.

_____. 1978. Socioecology of the vicuña. Ann Arbor: University Microfilms International.

_____. 1982. Biology, ecology, and relationship to man of the South American camelids. Mammalian biology in South America, Pymatuning Laboratory of Ecology Special Publication 6, ed. M. A. Mares and H. H. Genoways, 457–89. Linesville: University of Pittsburgh.

_____. 1983. Contrasting socioecologies of South America’s wild camelids: the vicuña and the guanaco. Advances in the study of mammalian behavior, Special Publication of the American Society of Mammalogists, vol. 7, ed. J. F. Eisenberg and D. K. Kleinman, 573–629.

Garamendia, A., G. Palmer, J. C. DeMartini, and T. McGuire. 1987. Failure of passive immunoglobulin transfer: A major determinant of mortality in newborn alpacas (Lama pacos). American Journal of Veterinary Research 49(10):1472–76.

Gentz, E. J., and T. L. Yates. 1986. Genetic identification of camelids. Zoo Biology 5:349–54.

Gray, A. P. 1954. Mammalian hybrids. Slough: Commonwealth Animal Bureaux.

Hemmer, H. 1975. Zur Herkunft des Alpakas. Zeitschrift des Kölner Zoo 18(2):59-66.

_____. 1983. Domestikation Verarmung der Merkwelt. Braunschweig/Wiesbaden: Friedr, Vieweg und Sohn.

_____. 1990. Domestication: The decline of environmental appreciation. Cambridge: Cambridge University Press.

Herre, W. 1952. Studien über die wilden und domestizierten Tylopoden Südamerikas. Der Zoologische Garten (Leipzig, N.F.) 19(2–4):70–98.

_____. 1953. Studien am Skelet des Mittelhores Wilder und Domestizierter Formen der Gattung Lama Frisch. Basel Acta Anatomica 19:271–89.

_____. 1976. Die Herkunft des Alpaka. Zeitschrift des Kölner Zoo 19(1):22–26.

_____. 1982. Zur Stammesgeschichte der Tylopoden. Verhandlugen der Deutschen Zoologischen Gesellschaft 75:159–71.

Herre, W., and M. Röhrs. 1973. Haustiere—zoologisch gesehen. Stuttgart: Gustav Fischer Verlag.

Herre, W., and U. Thiede. 1965. Studien an Ghiren Südamerikanischer Tylopoden. Zoologisches Jahrbuch Anatomie 81:155–76.

Hesse, B. 1982. Archaeological evidence for camelid exploitation in the Chilean Andes. Säugetierkundliche Mitteilungen 30(3):201–11.

Hilzheimer, M. 1913. Überblick über die Geschichte der Haustierforschung, besonders der letzen 30 Jahre. Zoologischer Anzeiger 5:233–54.

Jerison, H. J. 1971. Quantitative analysis of the evolution of the camelid brain. American Naturalist 105:227–39.

Jürgens, K. D., M. Pietschmann, K. Yamaguchi, and T. Keinschmidt. 1988. Oxygen binding properties, capillary densities and heart weights in high altitude camelids. Journal of Comparative Physiology 158:469–77.

Kent, J. D. 1982. The domestication and exploitation of the South American camelids: Methods of analysis and their application to circum-lacustrine archaeological sites in Bolivia and Peru. Ann Arbor: University Microflims International.

Kleinschmidt, T., J. Marz, K. D. Jürgens, and G. Braunitzer. 1986. The primary structure of two Tylopoda hemoglobins with high oxygen affinity: vicuña (Lama vicugna) and alpaca (Lama pacos). Biological Chemistry Hoppe-Seyler 367:153–60.

Krumbiegel, I. 1944. Die neuveltlichen Tylopoden. Zoo-logischer Anzeiger 145:45–70.

_____. 1952. Lamas. Leipzig: Neue Brehmbücherei.

Kruska, D. 1982. Hirngrößenänderungen bei Tylopoden während der Stammesgeschichte und in der Domestikation. Verhandlugen der Deutschen Zoologischen Gesellschaft 75:173–83.

Larramendy, M., L. Vidal-Rioja, L. Bianchi, and M. Bianchi. 1984. Camélidos sudamericanos: estudios genéticos. Boletín de Lima 6(35):92–96.

Lavallee, D., M. Julien, J. C. Wheeler, and C. Karlin. 1986. Telarmachay chasseurs et pasteurs préhistoriques des Andes I. Paris: Editions Recherches sur les Civilisations, ADPF.

Ledger, C. 1860. Sur un tropeau d’alpacas introduit en Australie. Bulletin Société Impérial Zoologique d’Acclimatation (Paris) 7.

Leguía, 0. 1991. Enfermedades parasitarias. In Avances y perspectivas del conocimiento de los camélidos Sudamericanos, ed. S. Fernandez-Baca, 325–62. Santiago: FAO.

Lönnherg, E. 1913. Notes on guanacos. Ackiv für Zoologi (Stockholm) 8(19):1–8.

López Aranguren, D. J. 1930. Camélidos fosiles argentinos. Anales de la Sociedad Cientifica Argentina (Buenos Aires) 109:15–35, 97–126.

Miller, G. R., and A. R. Gill. 1990. Zooarchaeology at Pirincay, a formative period site in highland Ecuador. Journal of Field Archaeology 17:49–68.

Miller, G. S., Jr. 1924. A second instance of the development of rodent-like incisors in an artiodactyl. Proceedings of the United States Museum 66(8), no. 2545:1–4.

Miller, W. J., J. P. Hollander, and W. L. Franklin. 1985. Blood typing South American camelids. The Journal of Heredity 76:369–71.

Moore, K. M. 1988. Hunting and herding economies on the Junin Puna. In Economic prehistory of the Central Andes, ed. E. S. Wing and J. C. Wheeler, 154–66. Oxford: BAR International Series 427.

_____. 1989. Hunting and the origins of herding in Peru. Ann Arbor: University Microfilms International.

Murra, J. V. 1965. Herds and herders in the Inca state. In Culture and animals, ed. A. Leeds and A. P. Vayda, 185–216. Washington: American Association for the Advancement of Science Publication 78.

_____. 1975. Formaciones economicas y politicas del mundo andino. Lima: Instituto de Estudios Peruanos.

_____. 1978. La organizacion economica del estado Inca. Mexico: Siglo Ventiuno.

Novoa, C. 1989. Genetic improvement of South American camelids. Revista Brasileira de Genetica 12(3):123–35.

Penedo, M. C. T., M. E. Fowler, A. T. Bowling, D. L. Anderson, and L. Gordon. 1988. Genetic variation in the blood of llamas, Lama glama, and alpacas, Lama pacos. Animal Genetics 19:267–76.

Peterson, O. A. 1904. Osteology of oxydactylus. Annals of the Carnegie Museum 2(3), article 8:434–76.

Piccinini, M., T. Kleinschmidt, K. D. Jürgens, and G. Baunitzer. 1990. Primary structure and oxygen-binding properies of the hemoglobin from guanaco (Lama Guanacoë [sic]), Tylopoda. Biological Chemistry Hoppe-Seyler 371:641–48.

Pires-Ferreira, E. 1981/1982. Nomenclatura y nueva classificacion de los camélidos Sudamericanos. Revista do Museu Paulista 28:203–19.

Raedeke, K. J. 1979. Population dynamics and socioecology of the guanaco (Lama guanicoe) of Magallanes, Chile. Ann Arbor: University Microfilms International.

Ramírez, A. 1991. Enfermedades infecciosas. In Avances y perspectivas del conocimiento de los camélidos Sudamericanos, ed. S. Fernandez-Baca, 263–324. Santiago: FAO.

Reigadas, M. C. 1992. La punta del ovillo: Determinación de domesticación y pastoreo a partir del análisis microscopio de fibras y folículos pilosos de camélidos. Arqueologia 2:9–52. Rice, D. S. 1993. Late intermediate period domestic architecture and residential organization at La Yaral, Peru. In Andean domestic architecture, ed. M. S. Aldenderfer, 66–82. Iowa City: University of Iowa Press.

Röhrs, M. 1957. Ökologische Beobachtungen an wildebenden Tylopoden Südamerikas. Velhanlung der Deutchen Zoologischen Gesellschaft, 538–54.

Saluda-Gorgul, A., J. Jaworski, and J. Greger. 1990. Nucleotide sequence of satellite I and II DNA from alpaca (Lama pacos) genome. Acta Biochimica Polonica 37(2):283–97.

Semorile, L. C., J. V. Crisci, and L. Vidal-Rioja. In press. Restriction sites variation of the ribosomal DNA in Camelidae. Genetica 90.

Shimada, M. 1985. Continuities and changes in pattern of faunal resource utilization: formative through Cajamarca periods. In Excavations at Huacaloma in the Cajamarca Valley, Peru, 1979, ed. K. Terada and K. Onuki, 303–36. Tokyo: University of Tokyo Press.

Shimada, M., and I. Shimada. 1985. Prehistoric llama breeding and herding on the north coast of Peru. American Antiquity 50(l):3–26.

Stabl, P. W. 1988. Prehistoric camelids in the lowlands of western Ecuador. Journal of Archaeological Science 15:355–65.

Stanley, H. F., M. Kadwell, and J. C. Wheeler. 1994. Molecular evolution of the family Camelidae—A mitochondrial DNA study. Proceedings of the Royal Society London B 256:1–6.

Steinbacher, G. 1953. Zur Abstammung des Alpakka, Lama pacos (Linne, 1758). Säugetierkundliche Mitteilungen 1(2):78–79.

Taylor, K. M., D. A. Hungerford, R. L. Snyder, and F. A. Ulmer Jr. 1968. Uniformity of karotypes in the Camelidae. Cytogenics 7:8—15.

Thomas, 0. 1891. Notes on some ungulate mammals. Proceedings of the Zoological Society of London, 384–89.

Van der Hammen, T., and G. W. Noldus. 1986. Pollen analysis of the Telarmachay Rockshelter. In Telarmachay chasseurs et pasteurs préhistoriques des Andes I, ed. D. Lavallee, M. Julien, J. C. Wheeler, and C. Karlin, 379–87. Paris: Editions Recherches sur les Civilisations, ADPF.

Vidal, 0. 1967. La crianza de la llama y algunas caracteristicas de su fibra. Tesis, Universidad Nacional Agraria La Molina, Lima.

Vidal-Rioja, L., L. Semorile, N. O. Biachini, and J. Padron. 1987. DNA composition in South American camelids I. Characterization and in situ hybridization of satellite DNA fractions. Genetica 72:137–46.

Von Bergen, W., ed. 1963. Wool handbook. New York: Interscience.

Wachtel, N. 1977. The vision of the vanquished. New York: Barnes and Noble.

Weber, M. 1928. Die Säugetiere. Jena.

Wheeler, J. C. 1982. Aging llamas and alpacas by their teeth. Llama World 1(2):12–17.

_____. 1984. On the origin and early development of camelid pastoralism in the Andes. In Animals and archaeology, vol.. 3, Early herders and their flocks, ed. J. Clutton-Brock and C. Grigson, 395–410. Oxford: BAR International Series 202.

_____. 1986. De la chasse a l’elevage. In Telarmachay chasseurs et pasteurs préhistoriques des Andes I, ed. D. Lavallee, M. Julien, J. C. Wheeler, and C. Karlin, 21–59. Paris: Editions Recherches sur les Civilisations, ADPF.

_____. 1991. Origen, evolución y status actual. In Avances y perspectivas del conocimiento de los camélidos Sudamericanos, ed. S. Fernandez-Baca, 11–48. Santiago: FAO.

_____. In press. Faunal remains from the Chiribaya Alta tombs. In Chiribaya: A biocultural approach to the study of a prehistoric Andean polity, ed. S. Williams and J. Buikstra.

Wheeler, J. C., E. Pires-Ferreira, and P. Kaulicke. 1976. Preceramic animal utilization in the Central Peruvian Andes. Science 194(4264):483–90.

Wheeler, J. C., A. J. F. Russel, and H. F. Stanley. 1992. A measure of loss: prehispanic llama and alpaca breeds. Archivos de Zootecnia (Cordoba) 41 (extra):467–75.

Wing, E. S. 1972. Utilization of animal resources in the Peruvian Andes. In Andes 4: Excavations at Kotosh, Peru, 1963 and 1964, ed. I. Seiichiand K. Terada, 327–52. Tokyo: University of Tokyo Press.

_____. 1977. Animal domestication in the Andes. In Origins of agriculture, ed. C. A. Reed, 837–59. The Hague: Mouton.

_____. 1986. Domestication of Andean mammals. In High altitude biogeography, ed. F. Vulleumier and M. Monasterio, 246–64. Oxford: Oxford University Press.

Yacobaccio, H. D., and G. M. Madero. 1992. Zooarqueología de Huachichocana III (Jujuy, Argentina). Arqueología 2:149–88.