Sweating in the guanaco (Lama guanicoe)

Sweat glands are present all over the skin, where sweat production varies from 4.98 to 73.36gm(-2)h(-1) of skin. Ambient temperatures between 20 and 33 degrees C are the main stimuli for activation of sweat glands, generating a heat loss ranging from 11.9 to 37% of standing basal metabolic rate. Respiratory water loss is not an important mechanism for heat dissipation. Water loss is controlled by postural changes in the guanaco.     

Non-genetic factors affecting morphometric and fleece traits in guanaco (Lama guanicoe guanicoe) populations from Argentinean Patagonia

Lama guanicoe guanicoe is the guanaco subspecies located in the Argentinean Patagonia. It is of particular interest to characterize these populations as it is a native species and a high quality fibre production genetical resource. With the aim of describing phenotipically Lama guanicoe guanicoe populations, 262 specimens were sampled. The animals came from four different ecological regions, both wild and bred in captivity, from 5 ranches. Three age classes were established: calves (0–1 year), yearlings (1–2 years) and adults (more than 2 years). Morphometric variables measured were: body length (BL), thoracic depth (TD), half thoracic perimeter (HTP), head length (HL), shin length (ShL) and metacarpal circumference (MC). Fleece samples were evaluated on two ranches (n = 115) for: fleece total weight (FTW) and mean diameter of down (MDFD). Relationships between variables were studied and differences between morphostructures considering sex, age and ranch were analyzed. Age was the most important differencing factor (p < 0.05) for all morphometric traits (except for the ShL between yearlings and adults). Calves had finer MDFD (14.2 ± 0.6 μm) and less FTW (159 ± 47 g) than yearlings (15 ± 5.8 μm and 254 ± 57 g). Sexual dimorphism was not found in calves nor in yearlings for any of the morphometric characters considered. Adult females presented greater significant values (p < 0.05) for TD (42.7 ± 2.1 cm) and HTP (55.4 ± 3.2 cm) than males (41.6 ± 2.3 cm and 53 ± 2.9 cm, respectively). Only calf males presented higher significant values than females for the FTW (169 ± 42 g and 147 ± 51 g, respectively). FTW and MDFD were positively correlated between them (0.42) and with the thoracic measures (TD and HTP), these data imply an inverse relationship between quality and quantity of the fleece and suggest that the thoracic variables are good indicators of the fleece animal production in these Patagonian populations.     

Habitat use and social structure of an isolated population of guanacos (Lama guanicoe) in the Monte Desert, Argentina

We studied abundance, habitat use and social structure of an isolated population of guanacos in the Lihue Calel National Park in central Argentina, based on data from foot surveys during 1998–2000. Based on topography and vegetation characteristics we distinguished three types of habitat: hills, valleys and piedmonts, and lowlands. Three types of social groups were observed: solitary males, family groups and male groups. The guanacos used hills preferentially, followed by valley and piedmont. The guanacos avoided the lowlands where the vegetation is dense. Habitat selection, both on a coarse and a fine scale, could be explained by habitat quality and possibly also predator avoidance behaviour. Abundance of guanacos increased by 21% over the 3-year-study period. However, the specificity of habitat requirements of guanacos and the susceptibility of the study area to wildfires could cause guanacos to move into sub-optimal areas were they are more vulnerable to predation and human persecution.     

Guanaco (Lama guanicoe) mortality by entanglement in wire fences

Wire fences are widely used in rangelands around the world and may have a negative impact on wildlife that varies among species and habitats. The guanaco (Lama guanicoe) is the largest Patagonian ungulate and though entanglement in wire fences is frequently reported, its impact on guanaco populations has not been previously evaluated. We estimated annual mortality rate of wild guanacos due to entanglement in wire fences and evaluated whether the frequency of entanglement was age-dependent in the two wire-fence designs traditionally used in Patagonian sheep ranches. We found that annual yearling mortality on fences (5.53%) was higher than adult mortality (0.84%) and was more frequent in ovine (93 cm high) than bovine (113 cm) fences. Most guanacos died entangled by their legs in the highest wire when trying to jump over the fence. Our results suggest that guanacos are more likely to die due to fence entanglement than ungulates studied in other regions. Indirect effects of wire fences should also be considered as they may act as semi-permeable barriers for guanaco populations. We suggest removal of unnecessary wire fences and replacement by guanaco-friendly fences, like high-tensile electric fences that may reduce mortality and barrier-effect on guanaco populations.     

Cooperative vigilance: the guanaco's (Lama guanicoe) key antipredator mechanism

The concept of sociality has been associated with the effectiveness of antipredator mechanisms, like cooperative vigilance and the dilution effect. Lama guanicoe (guanaco) is a social native herbivore in South America and a social species. The objectives of this study were to evaluate the antipredator responses of different-sized groups of guanacos in areas with varying predation risks and to determine antipredator mechanisms in guanacos. For this, we measured different antipredator responses to a potential predator (human subjects). Detection of predator and flight distances from predator both increased with a greater number of guanacos per group and with greater distances among guanacos within the social group. Both buffer distance and flight time decreased with a greater number of guanacos per group, but increased with greater distances among guanacos inside the social group. Solitary adult males moved shorter distance and mixed groups moved greater distances. Flight distances were greater in areas with tall and dense vegetation than in areas with low vegetation. Buffer distance and flight time were shorter in undulating land than on flat lands, and groups were usually observed on hill slopes. Our results suggest that the benefit of social grouping in guanacos is the increased probability of avoiding predator with cooperative vigilance and not with the dilution effect. This means that a predator could be detected earlier when approaching a guanaco group than when approaching a solitary individuals and could thus be avoided.     

Predator occurrence and perceived predation risk determine grouping behavior in guanaco (Lama guanicoe)

Grouping behavior of social ungulates may depend on both predator occurrence and perceived predation risk associated with habitat structure, reproductive state, and density of conspecifics. Over 3 years, we studied grouping behavior of guanaco (Lama guanicoe) families in Chilean Patagonia during the birthing season and determined their response to variation in predator occurrence and perceived predation risk (habitat structure, calf/adult rate, and density of conspecifics). We considered the effect of two predators, puma (Puma concolor) and culpeo fox (Lycalopex culpaeus). We measured two common (family group size and vigilance) and one novel (family group cohesion) behavioral responses of guanaco. Our results show that guanaco family groups adapted their grouping behavior to both predator occurrence and perceived predation risk. Larger family groups were found in open habitats and areas with high puma occurrence, while guanacos stayed in small family groups in areas with high shrub cover or low visibility. Group cohesion increased in areas with higher occurrence of pumas and culpeo foxes, and also increased in smaller family groups and in areas with low guanaco density. Vigilance (number of vigilant adults) was mainly related to group size and visibility, increasing in areas with low visibility, while residual vigilance (vigilance after removing the group‐size effect) did not vary with the explanatory variables examined. Our results suggest that a mix of predator occurrence and perceived predation risk influences guanaco grouping behavior and highlights the importance of evaluating different antipredator responses together and considering all predator species in studies aimed at understanding ungulate behavior.     

Macroscopic and histological investigation of guanaco footpads (Lama guanicoe,Müller 1776)

The surface of guanaco footpads is characterized by hairless skin with up to 4‐mm‐thick stratum corneum that protects from abrasion. The horny layer is pliable and elastic, and ensures firm contact with irregular ground. It is padded with a particular structure of the subcutaneous layer, the digital cushion. The flat cushions of each of the two digits are of elongated ovate shape, each about 45‐mm long, up to 20‐mm wide, and 8‐mm thick. The cushions are lined by a 1–2‐mm‐thick capsule that resembles a tunica albuginea. The capsule consists of coarse collagen fibers, with elastic fibers absent. The cushion capsule and dermis approach each other, and fuse along a line that runs parallel to the longitudinal axes of cushion and digit. Loose connective tissue rich in elastic fibers and acidic glycosaminoglycans separates dermis and cushion capsule lateral to the narrow interconnecting zone. The cushion capsule encloses cloudy yellowish, gelatinous material. Microscopy shows bundles of elastic fibers in abundant mucinous matrix. Tightly gathered elastic bundles adjoin the inner surface of the capsule. Rough cords of elastic fibers branch out from there and traverse to the opposite side. The cushion is pressed flat, and elastic fibers are stretched when bearing weight. With relief of load, elastic fibers contract and reset the cushion's shape. Contractile cells are absent. A resistant capsule and easily malleable mucinous contents establish the functioning as a gel pad. Mucinous connective tissue between elastic fiber bundles contains abundant basophilic matrix. Hyaluronan, chondroitin sulfate, and dermatan sulfate are main matrix constituents. Spindle‐shaped or stellate fibroblasts contain vimentin, S100 protein, and neuron specific enolase. Moprhology, staining characteristics and synthesis activities of these cells meet the criteria to be classified as myxoid cells. The connective tissue in guanaco digital cushions represents myxoid tissue. J. Morphol. 276:331–341, 2015. © 2014 Wiley Periodicals, Inc.     

Food selection by the guanaco (Lama guanicoe) along an altitudinal gradient in the Southern Andean Precordillera (Argentina)

Wild ungulates like the guanaco are exposed to important changes in climate and plant diversity along altitudinal gradients in the Andes Mountains, such as in the Southern Andean Precordillera where three phytogeographic provinces are present in altitudinal belts. The guanaco’s diet and food availability were seasonally analyzed using microhistological analysis and point-quadrat transects at six sampling sites, representative of the phytogeographic belts along the altitudinal gradient. Plant cover and diversity decreased with growing altitude. Richness of plant species was poorer at the summit than in the lower altitudes, whereas the proportion of species eaten by guanacos increased with altitude. The diet included 77 species. Grasses were preferred and shrubs were avoided all year round. The grass Poa spp. occupied more than 50 % of the diet at all altitudes. Grasses were the main dietary item even at low altitudes, where shrubs constituted the main food available. Decreasing generalism with descending phytogeographic belts agrees with the prediction for altitudinal gradients. The increase of diversity in the diet during the winter decline of plant cover at high and middle altitudes follows that expected from the optimal foraging theory. The winter decline of vegetation and the dietary shift from grazing to browsing proved to be stronger as altitude increases and the climate become more rigorous. Plant species richness, food scarcity, and climate severity are relevant variables to explain altitudinal and seasonal changes in the diet of adaptive ungulates in mountain environments, such as the guanaco in the Southern Andean Precordillera.     

Vigilance Patterns of Territorial Guanacos (Lama guanicoe): The Role of Reproductive Interests and Predation Risk

We conducted focal observations of territorial guanacos, a highly polygynous and social mammal, to compare time budgets between sexes and test the hypothesis that the differences in reproductive interests are associated with differential group size effects on male and female time allocation patterns. In addition, we used group instantaneous sampling to test the hypothesis that grouping improves detection capacity through increased collective vigilance. We fit GLM to assess how group size and group composition (i.e., presence or absence of calves) affected individual time allocation of males and females, and collective vigilance. As expected from differences in reproductive interests, males in family groups devoted more time to scan the surroundings and less to feeding activities compared to females. Both sexes benefited from grouping by reducing the time invested in vigilance and increased foraging effort, according to predation risk theory, but the factors affecting time allocation differed between males and females. Group size effects were significant when females were at less than five body‐lengths from their nearest neighbour, suggesting that grouping benefits arise when females are close to each other. Female time budgets were also affected by season, topography and vegetation structure. In contrast to our expectation, males reduced the time invested in vigilance as the number of females in the group increased, supporting the predation risk theory rather the intrasexual competition hypothesis. The presence of calves was associated with an increase in male individual vigilance; and vegetation type also affected the intensity of the group size effect over male time allocation. In closed habitats, collective vigilance increased with the number of adults but decreased with the number of calves present. Although male and female guanacos differed in their time allocation patterns, our results support the hypothesis that both sexes perceive significant antipredator benefits of group living.     

Dominance and anti-predator behaviour in blue tits (Parus caeruleus)

Evidence from captive flocks of blue tits (Parus caeruleus) suggests that following the appearance of a predator, socially dominant individuals are likely to wait until subordinate members of their flock resume feeding before doing so themselves. After a model sparrowhawk (Accipiter nisus) was flown over captive flocks of blue tits at a feeder, there was a significant negative correlation between sequence of return to the feeder and status within a linear dominance hierarchy. During this period, birds increased the proportion of their time budgets devoted to scanning for predators. These results suggest that during periods of danger, high-ranking individuals are able to be more cautious than are low-ranking individuals, possibly because their ability to control food resources reduces the energetic costs of their extra caution.     

Die Halsäste des Nervus vagus beim Alpaka,Lama guanicoe pacos (Linné 1758)

Demmel, U. 1980. Die Halsäste des Nervus vagus beim Alpaka, Lama guanicoe pacos (Linné 1758). [The branches of the vagus nerve in the neck of Alpaka, Lama guanicoe pacos (Linné 1758).] (Anatomisches Institut der Universität Köln, Bundesrepublik Deutschland.) — Acta zool. (Stockh.) 61(3): 141–146. Dissections of two newborn and two adult alpacas have shown: that the trapezius musculature of either side is supplied by branches of seven cervical spinal nerves and not by the external ramus of truncus nervi accessorii as in mammals in general; that the nerves which innervate the inner laryngic muscles as well as the mucous membrane of the lower part of the larynx are given off from the main trunk of the vagus nerve just outside the jugular foramen; that a nerve corresponding to the n. laryngeus recurrens of other mammals is absent. It is concluded that the lack of recurvation of the inferior laryngic nerve and the absence of the spinal part of the accessory nerve are character states of tylopod artiodactyls.     

A case of allosuckling in wild guanacos (<Emphasis Type="Italic">Lama guanicoe</Emphasis>)

Although allosuckling, the lactation of non-filial offspring, can be a costly behavior, it has been reported in several species across a wide range of mammalian orders. Monotocous species such as ungulates exhibit this behavior less frequently than polytocous species, and most cases have been reported among captive specimens. A case of allosuckling of a free-ranging guanaco (Lama guanicoe) calf is reported. Allosuckling observed in this species may represent opportunistic behavior by the calf, supporting the “milk theft” hypothesis.     

The influence of the arid Andean high plateau on the phylogeography and population genetics of guanaco (Lama guanicoe) in South America

A comprehensive study of the phylogeography and population genetics of the largest wild artiodactyl in the arid and cold‐temperate South American environments, the guanaco (Lama guanicoe) was conducted. Patterns of molecular genetic structure were described using 514 bp of mtDNA sequence and 14 biparentally inherited microsatellite markers from 314 samples. These individuals originated from 17 localities throughout the current distribution across Peru, Bolivia, Argentina and Chile. This confirmed well‐defined genetic differentiation and subspecies designation of populations geographically separated to the northwest (L. g. cacsilensis) and southeast (L. g. guanicoe) of the central Andes plateau. However, these populations are not completely isolated, as shown by admixture prevalent throughout a limited contact zone, and a strong signal of expansion from north to south in the beginning of the Holocene. Microsatellite analyses differentiated three northwestern and 4–5 southeastern populations, suggesting patterns of genetic contact among these populations. Possible genetic refuges were identified, as were source‐sink patterns of gene flow at historical and recent time scales. Conservation and management of guanaco should be implemented with an understanding of these local population dynamics while also considering the preservation of broader adaptive variation and evolutionary processes.