Lama guanicoe

Geographic Range

Guanacos are found in South America from northern Peru southward. Their range includes Peru, western Bolivia, Argentina, Chile, Tierra del Fuego, and Navarino Island (Fowler, 2010). Populations in the northern part of the distribution, between northern Peru and northern Chile, are assigned to the subspecies Lama guanicoe cacsilensis . The remaining populations are assigned to the subspecies Lama guanicoe guanicoe (Franklin, 2011). Guanacos were introduced from Argentina to Staats Island, one of the Falkland Islands, in the 1930's, and a thriving population remains there (Franklin and Grigione, 2005).

• Biogeographic Regions
• neotropical neotropical
  • introduced introduced
  • native native

Habitat

Guanacos are the mostly widely distributed of the four species of South American camelids and they occupy the most diverse range of habitats (Franklin, 1982). Adapted to harsh, highly seasonal environments, guanacos are able to cope with such starkly contrasting climates as those of the hyper-arid Atacama Desert in northern Chile and the perpetually wet Tierra del Fuego at the southern tip of South America (Franklin, 1982). Guanacos prefer open, dry habitats, and they avoid steep slopes, cliffs, and rock (Franklin, 1982). They are found in 4 of the 10 major South American habitats: desert and xeric shrublands, montane and lowland grasslands, savannas and shrublands, and wet temperate forests (Franklin, 2011). In general, guanaco habitats are characterized by high winds, low precipitation, and low primary productivity (de Lamo et al., 1998; González et al., 2006). Guanacos are distributed over an elevation range from sea level to 4,500 m (Franklin, 2011).

• Habitat Regions
• temperate temperate
• tropical tropical
• terrestrial terrestrial

• Terrestrial Biomes
• desert or dune desert or dune
• savanna or grassland savanna or grassland
• forest forest
• scrub forest scrub forest
• mountains mountains

Physical Description

Guanacos have long, slender necks and long legs typical of camelids (Franklin, 2011). Adults are 90 to 130 cm high at the shoulders, and adult body mass is between 90 and 140 kg, with the smallest individuals found in northern Peru and the largest in southern Chile (Fowler, 2010; Franklin, 2011). All guanacos have pelage that is light to dark reddish brown, with white countershading on the chest, belly, and legs, and gray or black coloration of the head (Wheeler, 1995). Though the general appearance is similar in all populations, overall coloration can vary somewhat by region, with northern populations tending to be relatively light (Franklin, 2011). There is no sexual dimorphism in body size or coloration, although males have significantly enlarged canines (Franklin, 2011; Sarno and Franklin, 1999).

All camelids have relatively small heads, no horns or antlers, and a split upper lip. South American camelids ( Lama glama , Lama guanicoe , Vicugna pacos , and Vicugna vicugna ) lack humps, and can be distinguished from Old World camelids ( Camelus bactrianus and Camelus dromedarius ) by their smaller size and slender, rather than broad, feet (Franklin, 2011). Guanacos are slightly larger than alpacas and significantly larger than vicuñas , though smaller and less heavily built than llamas (Rosemann and Morrison, 1963). Within the South American camelids, tooth characteristics can be used to distinguish guanacos and llamas (genus Lama ) from vicuñas and alpacas (genus Vicugna ). Guanacos and llamas have spatulate, closed-root lower incisors, and both the labial and lingual surfaces of each crown are enameled. Vicuñas and alpacas have elongate, ever-growing incisors with enamel only on the labial sides (Fowler, 2010). The general dental formula of South American camelids is 1/3, 1/1, 1–2/1–2, 3/3 = 28–32 (Franklin, 2011).

To cope with the harsh and variable climates they encounter throughout their broad distribution, guanacos have developed physiological adaptations that allow them to respond flexibly to changes in environmental conditions (de Lamo, 1998). By adjusting their body position, for example, individuals can “open” or “close” thermal windows—areas of very thin wool located in their front and rear flanks—in order to vary the amount of exposed skin available for heat exchange with the environment (de Lamo et al., 1998; Franklin, 2011). This allows them to reduce heat loss quickly when the ambient temperature drops (de Lamo et al., 1998).

• Other Physical Features
• endothermic endothermic
• homoiothermic
• bilateral symmetry bilateral symmetry

• Sexual Dimorphism
• sexes alike

Reproduction

Guanaco reproductive strategy is a form of resource-defense polygyny: resident males defend feeding territories against the intrusion of other males (Franklin, 1982). These territories, which may offer protection from predators as well as the food resources essential to breeding females, are typically 0.07 to 0.13 km^2 in size and occupied either year-round or seasonally by family groups (Franklin, 1982; Franklin, 2011; Sarno and Franklin, 1999). Despite the name, the members of a particular family group are not necessarily related (Sarno et al., 2003, p. 23). Each family group is composed of one territorial male and a variable number of females and young, with total adult membership usually between 5 and 13 (González et al., 2006). Female membership in family groups is fluid, with the resident male generally allowing females and juveniles to leave freely, but regulating the entry of new females (Franklin, 2011). Guanaco males become territorial at 4 to 6 years old, and thereafter they engage in violent competition to establish residence in feeding territories and control of family groups (Fowler and Bravo, 2010). Although guanacos are not sexually dimorphic in body size, males have significantly enlarged canines, which they use in intense, frequently injurious male–male fights (Franklin, 2011). Aggressive behaviors seen in guanaco males include spitting (up to 2 m); making threatening displays; chasing and fleeing; biting at the legs, hindquarters, and necks of their opponents; ramming chests; and neck wrestling (Franklin, 2011). Fights are frequent and serious enough that neck, leg, and body wounds are commonly observed in territorial males (Franklin, 2011).

• Mating System
• polygynous polygynous

Guanacos reproduce seasonally and there is one breeding season per year. Mating occurs in family groups during the austral summer, typically between early December and early January (Franklin, 2011). The timing of birth is influenced by environmental conditions and thus varies widely by latitude, but offspring are generally born in November and December (González et al., 2006; Sarno et al., 2003). The gestation period is 11.5 months, and a single offspring, with birth weight about 10% of maternal weight, is born to each breeding female every year (Sarno and Franklin, 1999; Sarno et al., 2003). Twins are extremely rare, and only one neonate ever survives (Franklin, 2011).

As expected in light of their long gestation, guanaco young, called chulengos, are precocial, able to stand as early as 5 to 76 minutes postpartum (González et al., 2006; Hofman, 1993). Chulengos exhibit following, rather than hiding, behavior; this may be a strategy to avoid predation in open habitats (González et al., 2006; Ralls, Kranz, and Lundrigan, 1986). Because of the need to grow quickly in advance of harsh winters, chulengos begin grazing within a few weeks of birth, and they forage almost exclusively by the time they are 8 months old (Franklin, 2011; González et al., 2006; Sarno et al., 1999).

Female guanacos reach sexual maturity at 2 years old and breed for the first time at age 3 (Franklin, 2011). Males reach maturity at 2 to 4 years old and begin breeding after obtaining a feeding territory between age 2 and age 6 (Fowler and Bravo, 2010; Franklin, 2011). Each year, 75% of adult females and 15 to 20% of adult males reproduce (Sarno and Franklin, 1999).

Guanacos are induced ovulators, like other South American camelids, and females undergo a follicular wave cycle rather than a regular estrous cycle (Franklin, 2011). Ovulation occurs 24 to 48 hours after copulation and is triggered by a hormonal signal in the semen (Fowler and Bravo, 2010; Franklin, 2011).

• Key Reproductive Features
• iteroparous iteroparous
• seasonal breeding seasonal breeding
• gonochoric/gonochoristic/dioecious (sexes separate)
• sexual sexual
• induced ovulation induced ovulation
• viviparous viviparous

Guanaco mothers play an important role in defending their young against predators (González et al., 2006). Predation accounts for a large proportion of juvenile mortality, and maternal aggressiveness toward potential predators, including threatening, spitting, charging, and kicking, has been shown to increase the survival of guanaco young (González et al., 2006; Sarno et al., 1999). Birth weight, growth rate, and suckling behavior are similar in male and female offspring (Sarno and Franklin, 1999). All chulengos are weaned between 4 and 8 months of age, although they return to nursing as yearlings after the birth of the next season’s young (Fowler and Bravo, 2010). This period of overlap, when neonates and older juveniles coexist and compete for resources in family groups, suggests a high level of parental investment in guanacos compared with other South American camelids (Franklin, 1982). Juvenile vicuñas , for example, are expelled from family groups before they reach one year of age, and thus are absent before the next birthing season begins (Franklin, 1982). It has been hypothesized that, for guanacos, additional time in the natal group is required for further growth or socialization (Franklin, 1982).

In guanacos, juveniles of both sexes are expelled from family groups in late spring and early summer, when they are between 11 and 15 months old (Franklin, 1982; Sarno et al., 2003). As the breeding season progresses, territorial males become increasingly aggressive toward juveniles born the previous year, until all yearlings are finally forced to disperse (Sarno et al., 2003). Since forced dispersal is not sex-biased, and since male aggression toward yearlings peaks when neonates are beginning to forage and feeding competition within the family group is high, the expulsion of juveniles is probably aimed at freeing up resources to reduce competition for food (Sarno et al., 2003). Upon dispersal, yearling females often travel alone or together among solitary territorial males (Sarno et al., 2003). Alternatively, they may join female groups or established family groups (Franklin, 2011). Yearling males usually join male groups, where they remain for 1 to 3 years while honing their fighting skills by means of aggressive play (Franklin, 2011).

• Parental Investment
• precocial precocial
• female parental care female parental care
• pre-fertilization
  • provisioning
  • protecting
    • female
• pre-hatching/birth
  • provisioning
    • female
  • protecting
    • female
• pre-weaning/fledging
  • provisioning
    • female
  • protecting
    • female
• pre-independence
  • provisioning
    • female
  • protecting
    • female

Lifespan/Longevity

Guanacos may live up to 28 years in captivity or in the wild (Franklin, 2011). In captivity, the longest known lifespan is 33.7 years (Tacuta et al., 2013).

Behavior

Guanacos have a fairly flexible social system, and populations may be sedentary or migratory depending on the year-round availability of forage (Franklin, 1982). During the breeding season, guanacos are found in three primary social units: family groups, male groups, and associations of solitary males (Franklin, 2011; González et al., 2006). Each family group is headed by a territorial adult male and contains a variable number of adult females and young (González, 2006). Outside of family groups, non-breeding, non-territorial juvenile and adult males form male groups of 3 to 60 individuals and forage in distinct male-group zones (Franklin, 2011). Mature males with territories but without females are called solitary males, and they may form associations of about 3 (Franklin, 2011).

Environmental conditions determine group composition after the breeding season ends (González et al., 2006). Where winters are mild and forage supply is stable, populations are sedentary and breeding males defend their feeding territories year round, though females may leave to form winter groups of 10 to 90 individuals (Franklin, 1982; Franklin, 2011; González et al., 2006). Where drought or snow cover reduces food availability in the winter, guanacos form mixed-sex herds of up to 500 individuals and travel to more sheltered or food-abundant areas (Franklin, 2011; González et al., 2006). These migrations may be altitudinal or lateral shifts, depending on climate and geography (Franklin, 1982). In migratory populations, increased group size during winter migrations may be a strategy for reducing predation, since snow cover and relative isolation in the environment make guanaco populations especially vulnerable during this time (Ortega and Franklin, 1995).

• Key Behaviors
• cursorial
• terricolous
• diurnal diurnal
• motile motile
• migratory migratory
• sedentary sedentary
• territorial territorial
• social social

Home Range

There is a good deal of variability in home range size. Populations in eastern Patagonia have ranges of 4 to 9 km^2, for example, while those in western Patagonia may have ranges twice as large (Baldi et al., 2010).

Communication and Perception

The open habitats that guanacos inhabit facilitate visual communication by allowing gestures to be seen over long distances (Franklin, 2011). Male guanacos perform a series of displays to assert dominance over potential rivals (Franklin, 2011). Broadside and S-neck displays may be used for advertising a male’s presence, asserting his status, or intimidating other males (Franklin, 2011). Other gestures, such as tail wagging, seem to be important for intra-group communication among both males and females (Franklin, 1975). Juvenile guanacos commonly use submissive displays to appease aggressive males attempting to drive them from family groups, and in doing so they can delay their forced dispersal (Sarno et al., 2003; Sarno et al., 2006). When approached or threatened by an adult male, a juvenile will enter a submissive crouch by lowering its neck, bending its knees, and raising its tail, a position that may strategically mimic the nursing posture (Franklin, 2011; Sarno et al., 2006). Guanacos produce a range of vocalizations for conveying information and negotiating social roles (Franklin, 2011). Notable among these are alarm calls, given to warn other group members of the presence of potential predators, and clicking noises, used in agonistic encounters between males (Franklin, 1982; Franklin, 2011). Odor is an important means of communication for many camelids (Franklin, 2011). Male guanacos use dung piles for territory marking and intra-group orientation, or keeping family groups together within territories (Franklin, 1975).

• Communication Channels
• visual visual
• tactile tactile
• acoustic acoustic
• chemical chemical

• Perception Channels
• visual visual
• tactile tactile
• acoustic acoustic
• chemical chemical

Food Habits

Guanacos are herbivorous. As browser-grazers, they are able to exploit a diverse range of food sources, and they exhibit flexible feeding behavior that varies across space and time (Franklin, 1982; González et al., 2006). In the Andean foothills of Argentina, two shrub species, Mulinum spinosum and Colletia spinosissima , constitute a large proportion of the year-round diet (Bahamonde, Martin, and Sbriller, 1986). When their preferred foods are not available, however, guanacos will eat lichens, fungi, cacti, fruit, and flowers in addition to grasses and shrubs (Franklin, 2011). Their generalist diet and efficient water and energy metabolism have enabled them to survive under harsh circumstances, including in extremely arid climates (Franklin, 1982).

• Primary Diet
• herbivore herbivore
  • folivore folivore

• Plant Foods
• leaves
• fruit
• flowers
• lichens

• Other Foods
• fungus

Predation

The primary predators of guanacos are pumas , which coexist with guanacos throughout their distribution except on Navarino Island and the islands of Tierra del Fuego (González et al., 2006; Novaro et al., 2009). In some populations, predation by pumas accounts for up to 80% of chulengo mortality (Sarno et al., 1999). Although pumas were the only confirmed predators for many years, researchers have recently reported attacks on juvenile guanacos by culpeos , which are present on Tierra del Fuego as well as in other parts of the guanaco range (Novaro et al., 2009).

For guanacos, group living is an important anti-predation strategy (Franklin, 2011; Novaro et al., 2009). Thanks to the protective effects of dilution and early detection, individuals living in groups are able to invest less time in vigilance, and more time in foraging, than are individuals living alone (Franklin, 2011; Novaro et al., 2009). Guanacos generally exhibit a “see and flee” response when they encounter potential predators (Franklin, 1982; Novaro et al., 2009). An individual will maintain visual contact with the predator until it gets too close, then give an alarm call to alert the rest of the group and escape by running away (Franklin, 1982). This strategy tends to be effective against pumas, which do not pursue their prey for long distances. By contrast, a more aggressive approach may be advantageous in dealing with smaller, cursorial predators such as culpeos. In at least one instance, adult guanacos have been observed engaging in cooperative defense against a culpeo: they cornered it, kicked it, and eventually drove it away after it chased and attacked a chulengo (Novaro et al., 2009).

Ecosystem Roles

Throughout their distribution, guanacos play an important role in maintaining ecosystem function. They disperse seeds in their feces, control vegetation growth by grazing and browsing, and serve as food sources for scavengers and predators (González et al., 2006). Guanaco carcasses are commonly eaten by foxes ( Lycalopex griseus , Lycalopex culpaeus ) and birds of prey ( Polyborus plancus , Vultur gryphus ), and guanacos are hunted by pumas and culpeos (González et al., 2006; Novaro et al., 2009).

Throughout the Andes and in parts of Patagonia, guanacos have historically been the primary prey species for pumas (Franklin et al., 1999; González et al., 2006). But as hunting pressures and habitat degradation have caused guanaco population densities to decline, they have been largely replaced in predator diets by introduced and domesticated species, such as red deer , sheep , and European hares . In some areas, interactions with pumas and other species have fallen to negligible levels and guanacos are said to be ecologically extinct (Novaro, Funes, and Walker, 2000). Guanacos compete with introduced domesticated species for forage, including compete for forage with sheep , goats , horses , donkeys , and cattle .

• Ecosystem Impact
• disperses seeds

Economic Importance for Humans: Positive

Guanaco wool is highly valued on the international market and prized for its softness and warmth (Baldi et al., 2010; Franklin, 2011). Although some guanacos are kept in captivity for the purpose of producing wool, efforts to capture, shear, and release wild guanacos have increased rapidly since these initiatives began in the late 1990's (Baldi et al., 2010). The demand for guanaco shearing is growing and Argentina now produces 1500 kg of guanaco wool per year (Baldi et al., 2010; Franklin, 2011). In addition, specialists called chulengueros hunt chulengos in Argentina for their pelts (Franklin, 1982).

• Positive Impacts
• body parts are source of valuable material

Economic Importance for Humans: Negative

Guanacos compete for forage with domesticated sheep, goats, horses, donkeys, and cattle and are considered by many ranchers to be pests (Baldi et al., 2010; González et al., 2006).

Conservation Status

Because guanacos remain widespread in South America, they are classified as Least Concern on the IUCN Red List. Still, careful management of local populations is necessary to prevent population declines (Baldi et al., 2008). This is true especially in light of the increasing demand for live shearing, which has resulted in capture myopathy in some wild guanacos and which may have further negative consequences for the growing number of populations involved (Baldi et al., 2008; Baldi et al., 2010). Guanaco populations are also threatened by disease transmission from domestic livestock; by overhunting, especially of chulengos for pelts; and by land degradation due to intensification of agriculture and overgrazing by sheep (Baldi et al., 2004; Baldi et al., 2008; Baldi et al., 2010; Franklin, 1982; González et al., 2006). Fences erected by sheep ranchers can interfere with guanaco migratory routes and can kill chulengos that become entangled in the wires (Baldi et al., 2010; Franklin, 1982).

As a result of human pressures, guanacos now occupy less than 40% of their original distribution in South America and existing populations are frequently small and fragmented (Baldi et al., 2010; Franklin, 2011). The governments of Argentina, Bolivia, Chile, and Peru regulate the use of wild guanacos within their borders, but enforcement of conservation laws tends to be weak, and most guanaco habitats are not under effective protection (Baldi et al., 2008).

Other Comments

Guanacos are the ancestors of domesticated llamas (Franklin, 2011).

Additional Links

Encyclopedia of Life

Contributors

Eva Hoffman (author), Yale University, Eric Sargis (editor), Yale University, Tanya Dewey (editor), University of Michigan-Ann Arbor.

References

Bahamonde, N., S. Martin, A. Sbriller. 1986. Diet of guanaco and red deer in Neuquen Province, Argentina. Journal of Range Management , 39/1: 22-24.

Baldi, B., G. Lichtenstein, B. González, M. Funes, E. Cuéller, L. Villalba, D. Hoces, S. Puig. 2008. " Lama guanicoe " (On-line). IUCN 2013. IUCN Red List of Threatened Species. Version 2013.2. Accessed December 17, 2013 at http://www.iucnredlist.org/details/full/11186/0 .

Baldi, R., A. Novaro, M. Funes, S. Walker, P. Ferrando, M. Failla, P. Carmanchahi. 2010. Guanaco management in Patagonian rangelands: A conservation opportunity on the brink of collapse. Pp. 266-290 in Wild rangelands: Conserving wildlife while maintaining livestock in semi-arid ecosystems . Hoboken, NJ: Wiley-Blackwell.

Baldi, R., A. Pelliza-Sbriller, D. Elston, S. Albon. 2004. High potential for competition between guanacos and sheep in Patagonia. Journal of Wildlife Management , 68/4: 924-938.

Convention on International Trade in Endangered Species of Wild Fauna and Flora, 2013. "Appendices I, II and III" (On-line pdf). Accessed November 15, 2013 at http://www.cites.org/eng/app/2013/E-Appendices-2013-06-12.pdf .

Eisenberg, J. 1989. Mammals of the Neotropics . Chicago: University of Chicago Press.

Fowler, M., W. Bravo. 2010. Reproduction. Pp. 429-478 in Medicine and surgery of camelids . Hoboken, NJ: Wiley-Blackwell.

Fowler, M. 2010. Medicine and surgery of camelids . Hoboken, NJ: Wiley-Blackwell.

Franklin, W., M. Grigione. 2005. The enigma of guanacos in the Falkland Islands: the legacy of John Hamilton. Journal of Biogeography , 32: 661-675.

Franklin, W., W. Johnson, R. Sarno, J. Iriarte. 1999. Ecology of the Patagonia puma Felis concolor patagonica in southern Chile. Biological Conservation , 90: 33-40.

Franklin, W. 1982. Biology, ecology, and relationship to man of the South American camelids. Pp. 457-490 in Mammalian biology in South America , Vol. 6. Pittsburgh: University of Pittsburgh Press.

Franklin, W. 2011. Family Camelidae (camels). Pp. 206-247 in Handbook of the mammals of the world , Vol. 2. Barcelona: Lynx Edicions.

Franklin, W. 1975. Guanacos in Peru. Oryx , 13/2: 191-202.

González, B., R. Palma, B. Zapata, J. Marín. 2006. Taxonomic and biogeographical status of guanaco Lama guanicoe (Artiodactyla, Camelidae). Mammal Review , 36/2: 157-178.

Hofman, M. 1993. Encephalization and the evolution of longevity in mammals. Journal of Evolutionary Biology , 6: 209-227.

Novaro, A., M. Funes, R. Walker. 2000. Ecological extinction of native prey of a carnivore assemblage in Argentine Patagonia. Biological Conservation , 92/1: 25-33.

Novaro, A., C. Moraga, C. Briceño, M. Funes, A. Marino. 2009. First records of culpeo ( Lycalopex culpaeus ) attacks and cooperative defense by guanacos ( Lama guanicoe ). Mammalia , 73: 148-150.

Ortega, I., W. Franklin. 1995. Social organization, distribution and movements of a migratory guanaco population in the Chilean Patagonia. Revista Chilena de Historia Natural , 68: 489-500.

Ralls, K., K. Kranz, B. Lundrigan. 1986. Mother–young relationships in captive ungulates: variability and clustering. Animal Behavior , 34: 134-145.

Rosenmann, M., P. Morrison. 1963. Physiological response to heat and dehydration in the guanaco. Physiological Zoology , 36/1: 45-51.

Sarno, R., M. Bank, H. Stern, W. Franklin. 2006. Effects of age, sex, season, and social dynamics on juvenile guanaco subordinate behavior. Journal of Mammalogy , 87/1: 41-47.

Sarno, R., M. Bank, H. Stern, W. Franklin. 2003. Forced dispersal of juvenile guanacos ( Lama guanicoe ): causes, variation, and fates of individuals dispersing at different times. Behavioral Ecology and Sociobiology , 54: 22-29.

Sarno, R., W. Clark, M. Bank, W. Prexl, M. Behl, W. Johnson, W. Franklin. 1999. Juvenile guanaco survival: management and conservation implications. Journal of Applied Ecology , 36: 937-945.

Sarno, R., W. Franklin. 1999. Maternal expenditure in the polygynous and monomorphic guanaco: suckling behavior, reproductive effort, yearly variation, and influence on juvenile survival. Behavioral Ecology , 10/1: 41-47.

Sosa, R., J. Sarasola. 2005. Habitat use and social structure of an isolated population of guanacos ( Lama guanicoe ) in the Monte Desert, Argentina. European Journal of Wildlife Research , 51/3: 207-209.

South American Camelid Specialist Group 1996, 2013. " Lama guanicoe ssp. cacsilensis " (On-line). IUCN Red List of Threatened Species. Version 2013.1. Accessed November 15, 2013 at http://www.iucnredlist.org/details/11187/0 .

South American Camelid Specialist Group 1996, 2013. " Lama guanicoe ssp. guanicoe " (On-line). IUCN Red List of Threatened Species. Version 2013.1. Accessed November 15, 2013 at http://www.iucnredlist.org/details/11190/0 .

Tacutu, R., T. Craig, A. Budovsky, D. Wuttke, G. Lehmann, D. Taranukha, J. Costa, V. Fraifeld, J. de Magalhaes. 2013. "AnAge: The animal ageing and longevity database" (On-line). Human Ageing Genomic Resources. Accessed November 15, 2013 at http://genomics.senescence.info/species/entry.php?species=Lama_guanicoe .

Wheeler, J. 1995. Evolution and present situation of the South American Camelidae. Biological Journal of the Linnean Society , 54: 271-295.

de Lamo, D., A. Sanborn, C. Carrasco, D. Scott. 1998. Daily activity and thermoregulation of the guanaco ( Lama guanicoe ) in winter. Canadian Journal of Zoology , 76: 1388-1393.