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Evolution and Origin of the Domestic Camelids
By Jane C. Wheeler, PhD
To date the earliest evidence of camelid domestication comes
from archaeological sites located between 4,000 and 4,900
m (13,120- to 16,072-foot) elevation, in the puna ecosystem
of the Peruvian Andes. Both guanaco (Lama glama cacsilensis)
and vicuña (Vicugna vicugna mensalis) have inhabited
this tundra environment for approximately 12,000 years and,
together with the huemul deer Hippocamelus antisensis (d’Orbigny
1834), were the primary prey of early human hunters. Faunal
materials from archaeological sites (Wing 1986; Wheeler 1984,
1986; Wheeler et al. 1976; Moore 1988, 1989) indicate that
during the earliest occupation of this zone 12,000 to 7,500
years ago, approximately equal numbers of camelids and deer
were hunted, while during later periods the frequency of camelid
remains increased dramatically, suggesting a shift to the
utilization of domestic animals. Archaeozoological data from
one of these sites, Telarmachay Rockshelter, have produced
the most extensive evidence concerning this shift to date
(Wheeler 1984, 1986).
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.
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