3.14.1 Lama guanicoe (guanaco)
3.14.2 Vicugna vicugna (wild vicuña)
The Camelidae Family is represented in South America by four species. Two are wild: the guanaco (Lama guanicoe) and the vicuña (Vicugna vicugna); and two are domestic. The domestic llama (Lama glama) and alpaca (L. pacos) are found throughout the Andean altiplano and the southern part of the continent. They are extraordinarily important in economic, social and cultural terms. The domestic species are believed to derive from the guanaco - or perhaps the alpaca was a hybrid of the guanaco and the vicuña. They were domesticated some time between 4 000 and 3 500 B.C. (191, 217, 626).
Vernacular names: Guanaco, huanaco, luan (Argentina, Bolivia, Chile, Peru).
Geographical variation and distribution: The wild guanaco originally ranged from the Andean areas of northern Peru and Bolivia and adjacent parts of Paraguay down to Tierra del Fuego, covering most of Argentina and Chile. L. guanicoe survives in some isolated parts of the Peruvian altiplano, in northern Chile, in western Argentina from Catamarca to southern Patagonia and in Chile's Magallanes province. There are two recognized subspecies: the smaller L. guanicoe cacsilensis, in Peru and Bolivia, and L. guanicoe guanicoe in Chile and Argentina (106, 154, 217, 488, 244).
Elevational range: The wild guanaco is found from sea level up to 4 200 - 4 500 m (217, 245).
Size and weight: The guanaco, one of South America's largest terrestrial mammals, reaches its maximum size in Magallanes with a total adult length of 176-202 cm (average 190 cm), an average weight of 120 kg (105-140 kg), and a shoulder height of 110-115 cm (488). The Peruvian population is smaller with an adult weight of 70-120 kg (191, 217). There is no sexual dimorphism by size.
Habitat: Cold or temperate open areas such as shrub-steppe and semi-arid brushlands are preferred, but the habitat may range from deserts of the western Andean slopes to the wet coastal rainforests of Tierra del Fuego. Guanaco are often pushed into marginal habitats by the pressure of livestock activities (93, 193, 217, 245, 488).
Abundance: At the time of the Spanish conquest, the guanaco was the most abundant ungulate in South America, with an estimated population of 10 to 30 million (191, 245, 488). Current numbers are thought to be about 575 000, of which nearly 550 000 live in Argentina, 20 000 in southern Chile and 5 000 in Peru (217). Regional population density estimates in Magallanes vary from 0.02-0.2/km2, but there may be 2.8-40.5/km2 in more densely populated areas (488).
Behaviour: Guanaco live in family groups of one male, several females, and their young. The group size varies from two to 30 and averages eight (488) to 16 (306). The groups tend to be bigger during the mating season: some females may leave in the winter (217, 218). Subadults are forcibly evicted from the group at the age of 13 to 15 months and form male groups. The old males live alone.
These family groups live in permanent territories of 30 to 50 ha that the territorial male defends, but may migrate altitudinally or laterally in many areas, and sometimes form large mixed groups that winter together (93, 95, 217, 488).
Guanacos are active during the day, much of which is spent feeding. On detecting a potential predator they give a cry of alarm and flee while maintaining visual contact with the pursuer (93, 95, 218, 306, 488).
Feeding habits: The guanaco is a non-specialized herbivore and basically a grazer, but may also browse. The diet in Magallanes is made up of 62 percent grasses (mainly Festuca) and 15 percent browse (Nothofagus spp). Dicotyledons form 11 percent of the diet and are particularly important in the spring, i.e. the months of October and November. Guanacos also eat epiphytes, lichens and fungi (488).
The guanaco inhabits a great variety of habitats and so its diet may also vary greatly at different times and places. Guanacos are more efficient than sheep at digesting crude fibre and dry matter (280), and may compete with them for winter fodder. Winter food shortages caused by overgrazing by sheep are probably the main cause of guanaco mortality in Magallanes (488).
Reproduction: Reproduction takes place in the spring and summer from November to March, with most births falling between mid-December and January. The gestation period is 345-360 days (183, 217), roughly one year, and the female breeds again very soon after giving birth to a litter of one. The 8-12 kg newborn baby guanaco is very precocious and can graze and run about just a few days after birth. The mother may suckle her offspring up to six or eight months (93, 183, 217, 488).
Growth and age: The baby guanaco grows very fast: a female reaches sexual maturity at, perhaps, two, and adult size in three or four years. The age may be estimated by the tooth eruption and replacement pattern and the degree of wear and layers of cementum on the first mandibular incisor (488). Guanacos in captivity may live as long as 23 years (138).
Hunting: The people of South America used to hunt guanaco by driving them into a ravine where another group of hunters waited in ambush to spear them. They were also hunted on horseback with bolas. Today the weapon of choice is a long-range repeater rifle, while baby guanaco are driven to exhaustion by mounted riders and then clubbed to death (93, 193).
Products: Guanaco used to be the major protein source for many indigenous groups in the Andes and in southern South America (93, 144, 193, 217). People prefer to eat guanaco meat in the form of "charki" (jerky), a salted and dried preparation. The fresh meat is not considered very appetizing (93, 191, 217). The dressed weight constitutes 55-57 percent of the total weight and the dried meat 10.2 percent (144).
The soft pelts of baby guanacos two to three weeks old, much persecuted in Patagonia, are the most lucrative product. A total of 443 655 skins (451) were exported from Argentina between 1972 and 1979, but the amount dropped to 13 157 in 1983 and 10 250 in 1984. The short, coarse, scant (250 g/adult) wool is woven in various ways and the cured hides are used for shoes (144, 145, 191, 217).
Management: Argentina's remnant guanaco population is about 8 percent of the original estimated population and Chile's a bare 1 percent (191). Increasing numbers of European immigrants and their livestock, particularly sheep, displaced the guanaco as the prime herbivore on the steppes and mountainsides of southern South America.
Guanaco populations have been slashed by culling to protect pasture for sheep, commercial hunting of baby guanacos, fodder shortages in winter due to overgrazing by sheep, fencing that blocked seasonal migrations and became a fatal trap for guanacos attempting to cross it, and deforestation (93, 145, 191, 193, 488). The species is listed on Appendix II of CITES and is considered endangered in Peru. Guanaco hunting has been forbidden in Peru since 1977 and in Chile since 1972, but it is still authorized in Argentina's southern provinces.
South American camelid research and conservation began to receive considerable attention in the 1970s. The data on guanaco biology and abundance are good, and there are a number of management guidelines. The small, dispersed, remnant populations of Peru and northern Chile still need protection from hunting. In southern Argentina and Chile, there is a need for more explicit management goals and policies and to locate and define areas where guanaco could be given priority. There must be unified laws and management designed to make the rural population the prime beneficiaries and to enlist their support and cooperation.
With proper management, it is believed that guanaco populations could stabilize at about one million in Argentina and 100 000 in Chile, with a respective annual production of 100 000 and 10 000 (145, 191, 217).
Guanaco management options should be designed and analysed as alternative economic uses for marginal land. The social structure of the species would allow a good proportion of the solo males to be harvested without affecting the reproductive capacity of the population as a whole. The key factors in guanaco management are probably primary production and food availability for these affect: land carrying capacity; the effect of density-dependence on fertility; the extension of adequate environments in the management areas; the effect of migration on population; and various socio-economic aspects (486).
Vernacular name: Vicuña.
Geographical variation and distribution: The wild vicuña originally ranged from northern Peru or southernmost Ecuador across the Peruvian and Bolivian altiplano to northern Chile and Argentina. Today's disjunct populations are scattered from the department of Ancash in Peru to the provinces of San Juan in Argentina and Atacama in Chile, including the Bolivian departments of La Paz and Oruro. There are two recognized subspecies: V. v. vicugna in Argentina and V. v. mensalis in Peru and Bolivia (72, 96, 217, 245, 282).
Elevational range: The vicuña is an animal of the Andean altiplano from 3 000-4 800 m, but is mainly found around 4 000 m (71, 96, 217, 245, 282).
Size and weight: The total adult length is 160-180 cm, the shoulder height 80-96 cm, and the weight 35-55 kg. There is no sexual dimorphism by size (93, 217, 282).
Habitat: Vicuñas live in the cold, semi-arid puna with its plains, hills and valleys covered with herb or shrub vegetation, and prefer the relatively flat areas where grasses predominate. In the Pampa Galeras National Vicuña Reserve this constitutes 33 percent of the total area (217). The vicuña's habitat may border and occasionally overlap with that of the guanaco of the foothills of the cordillera at lower altitudes (71, 218, 282, 322). The vicuña's habitat may be shared by llamas, alpacas and sheep, and this means health problems and competition for fodder (282, 322).
Abundance: The vicuña was the dominant ungulate in the puna of the high Andes under natural conditions: the Peruvian population in Inca times is estimated at 1.5 million (71). In 1957 the population was probably still about 400 000 (322) but by the end of the 1960s the figure had dropped to 7 000-12 000 (313). By 1985 the total population was back up to 129 000, of which 100 000 in Peru (191). The IUCN lists the vicuña as vulnerable (576) and it is on Appendix I of CITES (221).
Population densities in the Pampa Galeras National Vicuña Reserve in the province of Ayacucho, Peru, can be as high as 87/km2 (217) whereas the figures are 0.9-1.8/km2 for Lauca National Park in northern Chile (506). The optimum estimated density for vicuña production in Pampa Galeras is 40-43/km2 (72, 486).
Behaviour: The vicuña is a diurnal, gregarious and territorial animal. Most of the population (60 percent) lives in permanent, territorial family groups, usually made up of a male, three females and two sets of offspring. They occupy the best habitats. Group size varies in accordance with the size and productivity of the territory from 2 to 19. There are also territorial family groups in marginal habitats, mobile family groups, bachelor groups of non-breeding males (5-50 individuals representing 22-24 percent of the population) and solitary males (218, 282, 306, 322, 506).
The sedentary groups have a sleeping territory where they spend the night, usually in high areas, and a feeding territory in the valleys and plains. The average size of the feeding territories in Pampa Galeras is 18.4 ha but the range is 2 to 56 ha. The young males are expelled from the group by the territorial male when they are 4-9 months old and the females at 10-11 months, before the next litter is born. The evicted males form bachelor groups whereas females generally attach themselves to marginal or mobile family groups (218, 282, 322). They normally have fixed scat stations and drink once a day. Their main sense is sight and their flight distance with respect to people varies from 50 m in protected reserves to 500 m (242).
Feeding habits: The vicuña is classified as a selective and efficient grazer, preferring bunch grasses such as Calamagostris vicunarum, and spends most of the day grazing (217, 218, 282, 322). There are apparently no specific studies on the vicuña's diet.
Reproduction: Vicuña give birth in the summer from February to March, the most favourable season in the puna with mild temperatures, frequent rains and very good grazing (71, 217, 282). After a gestation period of 330-350 days, the female gives birth to one offspring. The newborn 4-6 kg lamb is extremely precocious (183, 282) and can follow its mother and keep up with the family group only half an hour after birth. The lamb can eat grass at the age of one month but is suckled up to the age of 6-8 months (282).
The female mates again about two weeks after the birth. Copulation lasts 30 minutes and ovulation is copulation-induced. The female may be sexually mature at the age of one year (282) but most females experience first gestation after the age of two (217). In Pampa Galeras the total number of offspring for 100 females varied from 22 to 75, the low figures corresponding to very dry years (282). The population growth rate can be as great as 22 percent a year in the accelerated phase (191).
Mortality: An estimated 10-30 percent of newborn vicuñas die from hypothermia due to cold storms, or from pneumonia, diarrhoea caused by Escherichia coli and depredation (217). Stray and/or sheepdogs are cited as top predators. The fox Dusicyon culpaeus and the puma Felis concolor may occasionally attack vicuñas. The incidence of mange is linked to malnutrition (282, 322). The record age in captivity is nearly 24 years (138).
Hunting: In pre-Colombian times vicuña were often driven into stone-walled corrals or ditches, or hunted with bolas (93, 322). They are currently hunted with medium-calibre repeating rifles, preferably with telescopic sights, on foot or from vehicles, the hunters getting as close as possible for a sure shot (roughly 100 m). Or the vicuña may be hounded to someplace where the hunters can reach them on horseback or with dogs.
When family groups are hunted the male is killed first, and then the rest of the group, disoriented by the absence of their leader. An alternative is to kill several animals in a male group before the rest can flee out of gunshot range (282). In the Pampa Galeras reserve, when vicuñas need to be captured alive for relocation or shearing, a system has been worked out to drive the animals into a funnel arrangement of wood and nets with sides 400 m long, converging into a holding pen built at a 45° angle (282).
Products: The products of the vicuña are its meat, skin or hide and wool. The dark red, lean meat is highly esteemed by indigenous people in the high pampa (322, 634), and there are good prospects for marketing meat fresh, frozen, or as jerky (dried salted meat of the whole carcass with bone in). The animals are in peak condition for slaughter at the end of the rainy season, i.e. May-June (282).
The skins are stretched, preferably salted, and spread to dry in the shade, pegged to the dry ground with the inner side up. Vicuna products made from cured skins and wool, particularly the wool of young vicuña, are attractive and in great demand by tourists (96, 217, 322), but the very slender fibre limits the vicuña's value as a pelt. It is recommended that the wool be shorn or separated by sweating and the hides cured (72, 282). There is little demand for the hides, which are not appropriate for gloves or shoes, although they can be used for purses, bags and other upmarket craft goods.
The fine wool of the vicuña, with a diameter of 10-16 micras and length of 3-6 cm, is the most valuable product. The animals are shorn alive every two years (each vicuña produces about 250 g) or the wool may be taken from the hides of slaughtered animals (72, 96, 282). It is very good for certain types of weaving, preferably in the natural tones, and is worth from 500-1 000 US dollars per pound (217).
Management: The vicuña is unquestionably the key wild species of the Andean high country, with great economic and social potential for management (72, 164, 282, 506). The species was driven to the brink of extinction in the 1960s, but Andean countries (Argentina, Bolivia, Chile and Peru) then joined forces in an ambitious vicuña population recovery effort constituting what was apparently the most successful wildlife management plan in Latin America. The project has so far consisted of effective vicuña protection against poachers, reintroduction of the species in specially regulated areas, biological research and a series of experimental management measures. The success of the project can be attributed to the following: 1) the prohibition of hunting and international trade through vicuña conservation agreements (La Paz 1969, Lima 1979); 2) the organization of relatively effective supervision in protected areas; 3) substantial research efforts covering vicuña sociobiology, habitat carrying capacity, estimates of population levels and productivity; 4) fairly substantial backing from the rural population (targeted as the prime economic beneficiary should the project prove successful) and, 5) the existence of large stretches of marginal land unfit for other economic purposes.
A good part of the technical problems of vicuña use at phase three of the project are currently being solved ((282, 486, 506). Population increases have so far been achieved only in strictly protected areas and the Peruvian objective of repopulating 150 000 km2 of puna with 3 million vicuñas (72) still seems very far in the future. Close working relationships between the national programme promoters and the campesinos have yet to be achieved, and local community responsibility for supervision has not been as successful as hoped (282), vicuña recovery has recently suffered serious setbacks as a result of guerrilla activities in the mountains of Peru (P. Vásquez, pers. com., 1987). Educational extension and campesino organizational work must be increased if the rural community is to take greater responsibility during stage three of the project. Moreover, an ecological and economic study of the relationship between vicuñas and domestic herbivores in the habitat is also needed for effective vicuña management within the overall ecosystem.
Captive breeding: Newborn vicuñas are rather easily tamed and the species can be maintained quite well in zoo collections. The valuable wool has spurred a number of attempts at domestication and captive breeding (96, 138, 282, 322). However, the animal's shy nature makes it difficult to manage whereas the rigid social and territorial organization generates constant fighting when vicuña are confined in small areas, particularly between males, and vicuña-proof fencing of vast areas of its natural habitat is counter-productive, economically speaking. Captive breeding does not seem to be a viable alternative to rational vicuña management (86, 282, 322).