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The Guanaco

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

The guanaco is the largest wild artiodactyl in South America.  Fossil remains of L. guanicoe are found in Argentine Pleistocene deposits (López Aranguren, 1930; Cabrera, 1932; Menegaz, Goin & Ortiz Jaureguizar, 1989) which probably date to some 2 Myr (Webb, 1974)(Figure 3A).  They are also present at Tarija, Bolivia (Hoffstetter, 1986) in strata recently dated 97-73,000 years BP (before present)(MacFadden et al., 1983), but did not spread into the high Andean puna ecosystem before the establishment of modern climatic conditions between 12­9,000 years ago (Hoffstetter, 1986).  Prior to European contact, guanacos were found along the Pacific shore and into the high Andes from approximately 8' South latitude to Tierra del Fuego, as well as east into the Paraguayan Chaco and across the pampas to the Province of Buenos Aires (Torres, 1985; Tonni & Politis, 1980).  Neither fossil nor recent guanaco remains have been found in Ecuador or Colombia.  Raedeke (1979) has estimated the prehispanic guanaco population to have been 30 to 50 million based on carrying capacity of the territory which they occupied.  These numbers rapidly declined during the European conquest.  During the nineteenth century the impact of indiscriminate hunting and commercial sheep rearing reduced the guanaco population to 7 million, and at present an estimated 602,907 survive (Wheeler, 1991).

All guanacos exhibit similar pelage coloration varying from a dark reddish brown in the southern populations (L.g. guanicoe to a lighter brown with ocher yellow tones in the northern variety (L.g. cacsilensis).  The chest, belly and internal portion of the legs are more or less pure white, the head grey to black with white around the lips, eyes and borders of the ears.  Fiber diameter varies from 16.5 Jim to 24 itm and contains from 5 to 20% hair (Verscheure & Garcia, 1980; Carpio & Solari, 1982a).  Sexual dimorphism is absent except for the presence of large canines in the male.  Withers height of adult animals varies from II 0 to 120 cm for L.g. Lyuanicoe from Patagonia and Tierra del Fuego (Cabrera & Yepes, 1960; Franklin, 1982; Herre, 1952; Raedeke, 1979; MacDonagh, 1949), compared to 100 cm for the small northern guanaco L.g. cacsilensis (Herre, 1952).  Reported body length, from the tip of the nose to the base of the tail, varies from 167 (MacDonagh, 1949), 185 (Cabrera & Yepes, 1960), 191 (Raedeke, 1979) and 210 cm (Dennler de la Tour, 1954a) for L.g. gmanicoe and 90-100 cm for L.g. cacsilensis from Calipuy, Peru (Kostritsky & Vilchez, 1974).  Live weight for adult L.g. guanicoe varies from 120 to 130 kg (Raedeke, 1979; Miller, Rottman & Taber, 1973), compared to 96 kg for L.g. cacsilensis (Kostritsky & Vilchez, 1974).  See Figure 4 for comparison with the other South American camelids.

Four poorly defined subspecies of guanaco have been described: the first, Lama guaunicoe guanicoe (Miiller, 1776), is said to occur in Patagonia, Tierra del Fuego and Argentina south of 35' S latitude; the second, L.g. huanacus (Molina, 1782), is restricted to Chile; the third, L.g. cacsilensis L6nnberg, 1913, a local high altitude form from southern Peru; and the fourth, L.g. Krumbiegel, 1944, found on the eastern slope of the Argentine Andes between approximately 21' and 32' S latitude.  The characteristics which set each subspecies apart are not fully detailed in these early works, and little information on zoogeographic variations in guanaco morphology is available.  The most extensively studied populations are located at the southernmost limits of guanaco distribution, and it is clear that these larger, darker animals (L.g. icoe) contrast markedly with the smaller, lighter coloured specimens (L.g. cacsilensis) found at the northern boundary.

Recent survey data compiled by Torres (1992) shows population clusters which may correspond to the four proposed guanaco subspecies.  A predominately high altitude, small northern subspecies, L.g. cacsilensis, is found between 8' and 20' S latitude in Peru and northern Chile (Hoces, 1992; Cunazza, 1992).  At Pampa Galeras (Ayacucho, Peru) these animals descend to the coast suggesting that human disturbance has displaced them from this part of their range in other areas.  None the less, some authors (Ponce del Prado & Otte, 1984) have postulated the possible existence of an undescribed coastal subspecies of guanaco.  Osteological remains from Andean archaeological sites document the origin of llama domestication from the guanaco at high elevation localities within the range of L.g. cacsilensis starting some 6,000 years ago (Wing, 1986; Wheeler, 1984, 1991; Moore, 1988, 1989).

To the south, a second population cluster [L.g. huanacus ?] is found on the western slope of the Andes between 22' and 28' S latitude (Cunazza 1992), with a third [L.g. ?] in south-eastern Bolivia (Villalba, 1992), northwestern Paraguay (Torres, 1985) and on the eastern slope of the Andes from 19' to 30' or 3 1 ' S latitude (Puig, 1992).  Within each of these areas, domestic llama remains appear in archaeological sites located between 22' and 24' S latitude.  In the Salar de Atacama, Chile (Hesse, 1982; Dransart, 1991a & b) they are dated to 4,500 BP, while in the Province of Jujuy, Argentina llamas may be present by 3,400 BP (Yacobaccio & Madero, 1992).  It is unclear if these remains represent independent domestication events or the introduction of domestic animals from the central Andes.

Further south still, the range of L.g. guanicoe, the large austral subspecies, extends from approximately 32' S latitude on the eastern slope of the Chilean and Argentine Andes, throughout Patagonia to 55' S in Tierra del Fuego (Puig, 1992; Cunazza, 1992).  This subspecies is the best known of all the guanacos (Franklin, 1982, 1983; Raedeke, 1979 among others).  There is no evidence that the Patagonian guanaco was ever domesticated, although tamed animals were sometimes kept (Benavente, 1985).

A different interpretation of guanaco subspeciation has been proposed by Franklin (1982), with one population restricted to the western slope of the Andes between 8' and 41' S latitude, and a second isolated population on the eastern side between 18' and 55' S latitude.  If such a geographic separation, presumably produced by the salt pans of southern Bolivia and the crests of the Andean chain, is real then a stronger case might be made for subspecific status with L.g. cacsilensis being the northwestern form and L.g. guanicoe the southeastern.

Guanaco numbers have continued to decline since European contact.  In 1954, Dennler de la Tour (1954a) called attention to the impending disappearance of the Patagonian guanaco if the hunting of yearling chulengos was not controlled and protected reserves established.  In 1969, Grimwood found that the Peruvian guanaco population was on the edge of extinction, and in 1971 the goverment responded by declaring it an endangered species.  In 1974, the IUCN (International Union for the Conservation of Nature and Natural Resources) declared the guanaco a vulnerable species (Thornback & Jenkins, 1982).  At present the guanaco receives protection in 14 reserves in Argentina, 4 in Chile and 3 in Peru.  None the less these measures are insufficient and in Bolivia and Paraguay guanaco populations are not protected.  Recently the South American Camelid Specialist Group of the IUCN has urgently recommended increasing protection for the guanaco in general, and the relict northern L.g. cacsilensis in specific (Torres, 1985).  This animal is not only highly endangered but also virtually unknown to science, and the recommendation should be taken seriously since archaeozoological and ethnohistorical data demonstrate based on geographic distribution that L.g. cacsilensis is the ancestral form of the domestic llama (Wheeler, 1984, 1986, 1991).

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