Habitat use and preference by guanacos,vicuñas,and livestock in an altitudinal gradient in northwest Argentina

Distribution of ungulates is regulated by natural factors, such as presence of other species, climate, and habitat variability, as well as human activities, including livestock grazing and mining. Understanding the spatial dynamics of landscape use can help solve problems of access to key resources by native herbivores. Laguna Brava Reserve is a protected area of Argentina where the two wild South American camelids, guanacos and vicuñas, coexist with cattle, mules, horses, and goats. Information about the effects of livestock and mining activities, which are widespread throughout the region, on wild camelids is scarce. We used variable-width transects to determine the distribution of guanacos, vicuñas, equids, and cattle in relation to five habitat types defined according to vegetation and topography in the reserve. We carried out a correspondence analysis between the proportion of groups observed in and the proportion of area occupied by each habitat, and a χ ² goodness of fit test to establish if camelids and livestock selected a particular habitat type. Vicuñas were associated with grasslands at high altitudes while guanacos and livestock were associated with shrublands at lower elevations. This coevolutionary segregation between guanacos and vicuñas possibly reduced competition between the two species. Competition between camelids and livestock is probably low because of the low density of livestock. Vegas were preferred by all species year round but used more intensively in summer. In order to conserve the wild camelids of the region, governments must implement measures to conserve the critical habitat provided by the vegas.     

Y‐chromosome and mtDNA variation confirms independent domestications and directional hybridization in South American camelids

Investigations of genetic diversity and domestication in South American camelids (SAC) have relied on autosomal microsatellite and maternally‐inherited mitochondrial data. We present the first integrated analysis of domestic and wild SAC combining male and female sex‐specific markers (male specific Y‐chromosome and female‐specific mtDNA sequence variation) to assess: (i) hypotheses about the origin of domestic camelids, (ii) directionality of introgression among domestic and/or wild taxa as evidence of hybridization and (iii) currently recognized subspecies patterns. Three male‐specific Y‐chromosome markers and control region sequences of mitochondrial DNA are studied here. Although no sequence variation was found in SRY and ZFY, there were seven variable sites in DBY generating five haplotypes on the Y‐chromosome. The haplotype network showed clear separation between haplogroups of guanaco–llama and vicuña–alpaca, indicating two genetically distinct patrilineages with near absence of shared haplotypes between guanacos and vicuñas. Although we document some examples of directional hybridization, the patterns strongly support the hypothesis that llama (Lama glama) is derived from guanaco (Lama guanicoe) and the alpaca (Vicugna pacos) from vicuña (Vicugna vicugna). Within male guanacos we identified a haplogroup formed by three haplotypes with different geographical distributions, the northernmost of which (Peru and northern Chile) was also observed in llamas, supporting the commonly held hypothesis that llamas were domesticated from the northernmost populations of guanacos (L. g. cacilensis). Southern guanacos shared the other two haplotypes. A second haplogroup, consisting of two haplotypes, was mostly present in vicuñas and alpacas. However, Y‐chromosome variation did not distinguish the two subspecies of vicuñas.     

The complete mitochondrial DNA sequence of the guanaco (<Emphasis Type="Italic">Lama guanicoe)</Emphasis>: comparative analysis with the vicuña (<Emphasis Type="Italic">Vicugna vicugna)</Emphasis> genome

South American camelids comprise the guanaco (Lama guanicoe) and the vicuña (Vicugna vicugna), which are wild species, and the domestic llama (Lama glama) and alpaca (Lama pacos). This paper presents the first complete mitochondrial (mt) genome of the guanaco and the mt coding sequence of the vicuña. The guanaco mtDNA is 16,649 nt long and its composition and organization are similar to the mitochondrial genome of other mammals. Excluding the control region, comparison of the complete guanaco and vicuña mtDNA showed 4.4% sequence divergence. Nucleotide differences in peptide coding genes varied from 1.9% in ATP6 to 6.4% in Cyt b. These values are compatible with the close relatedness of both species identified by other authors. Based on the differences between the control region sequence here reported and that previously described, we also discuss the occurrence of NUMTs in the genome of South American camelids.     

Lack of feral livestock interference with native guanaco during the dry season in a South American desert

Analyzing coexistence of exotic and native ungulates in arid areas is important from both a theoretical and a species conservation perspective. We assess the habitat use patterns and possible interference between guanaco (Lama guanicoe) and feral livestock (donkey and cattle) in arid environments of South America. To determine habitat use and niche overlap between exotic and native ungulate species, ten sites with different habitats and six natural waterholes were selected. Plots (20 at each site, ten around each waterhole) were randomly set up and characterized by environmental variables and relative use by cattle, donkey and guanaco through faecal pellet counts. Aggregation, niche breadth and niche overlap of the three herbivores were analyzed at habitat level (mesoscale). A direct redundancy analysis was used to examine the relationships between abundance of herbivore faeces and environmental variables at microscale. Mesoscale analyses showed (i) an extensive use of the area by all three species, with guanaco having the highest niche breadth followed by donkey and cattle and (ii) a large, broad guanaco–donkey and donkey–cattle habitat overlap. However, results at a finer scale showed high spatial aggregation of feral livestock species and an independent use of territory by guanacos. This study is the first to provide information about habitat partitioning between guanacos and feral livestock in the hyper-arid Monte Desert biome and points to an apparent lack of negative effects on the native ungulate.     

History of canids in Chile and impacts on prey adaptations

Artiodactyl prey species of Chile, especially guanacos (Lama guanicoe), are reported to be very susceptible to predation by pack‐hunting feral dogs. It has been previously suggested that guanacos and endemic South American deer may have evolved in the absence of pack‐hunting cursorial predators. However, the paleoecology of canid presence in southern South America and Chile is unclear. Here, we review the literature on South American and Chilean canids, their distributions, ecologies, and hunting behavior. We consider both wild and domestic canids, including Canis familiaris breeds. We establish two known antipredator defense behaviors of guanacos: predator inspection of ambush predators, for example, Puma concolor, and rushing at and kicking smaller cursorial predators, for example, Lycalopex culpaeus. We propose that since the late Pleistocene extinction of hypercarnivorous group‐hunting canids east of the Andes, there were no native species creating group‐hunting predation pressures on guanacos. Endemic deer of Chile may have never experienced group‐hunting selection pressure from native predators. Even hunting dogs (or other canids) used by indigenous groups in the far north and extreme south of Chile (and presumably the center as well) appear to have been used primarily within ambush hunting strategies. This may account for the susceptibility of guanacos and other prey species to feral dog attacks. We detail seven separate hypotheses that require further investigation in order to assess how best to respond to the threat posed by feral dogs to the conservation of native deer and camelids in Chile and other parts of South America.     

Relationships between integumental characteristics and thermoregulation in South American camelids

Hair fibre is regarded as a unique mammalian feature with an important role for endothermy. Artificial selection for hair characteristics resulted in marked changes with regard to follicle number, type, distribution, growth and natural shedding. This review focuses on the fine fibre-producing South American camelids (SACs) and the relationship between their hair coat and thermoregulation. SACs have developed several special integumental characteristics. While the hair coat of the wild lamoids vicuña (Vicugna vicugna) and guanaco (Lama guanicoe) is formed by two types of hair (the coarse outer guard hairs and a finer undercoat), the domesticated llamas (Lama glama) and alpaca (Lama pacos) exhibit variably double coat and predominantly single coat, respectively. The distribution of the hair coat across the body is not homogenous. Thermal windows with shorter hair or thinner skin can be identified at the ventral abdomen, axillary space and inside of the thighs (about 20% of the skin), thus allowing to modulate heat dissipation. In contrast to sheep wool, lamoid fibres are mainly medullated. The thermal conductance of summer pelage was higher than that of the winter fleece and highest for the axillar and lower flanks. Lamoids have developed behavioural strategies to modify heat loss by adopting specific postures according to ambient conditions by closing or opening the thermal windows. Energy savings of 67% attributed to posture were calculated. SACs have shown to be able to adapt to a broad range of different climatic conditions. The specific integumental characteristics of SACs indicate that they have developed adaptation mechanisms particularly suited for cooler climates. Accordingly, hyperthermia might become a problem in hot, humid areas outside of their original habitat. Several studies showed the beneficial effect of shearing against heat stress. In particular, fertility in males exposed to heat stress may be improved by shearing. Infrared thermography reveals that in shorn animals the heat is radiated across the entire body surface and is not restricted to the thermal windows. However, shearing also changes the conditions of the protective layer, resulting in a loss of thermal conductance that may result in adverse effects when animals are kept under cold temperatures. The length of residual fibre appears to be crucial in avoiding excessive heat loss in a cold environment, as demonstrated by shearing experiments with different shearing machines. There is, therefore, potential for welfare considerations to conflict with industrial demands for fibre length or homogenous quality.     

Social group dynamics and composition of managed wild vicuñas (<Emphasis Type="Italic">Vicugna vicugna vicugna</Emphasis>) in Jujuy,Argentina

The vicuña is a high Andean wild camelid that lives year-long in groups. We analyzed the social organization of 98 marked vicuñas after capture and live shearing, focusing on group composition, spatial patterns, territoriality and habitat use. The social units analyzed were families, bachelor groups and solo animals. Location fixes of 54 males and 36 females were registered over a 2-year period, and home range was estimated. All females belonged to family groups, and 8–12 month yearlings changed their home ranges prior to first reproduction at 2 years. Female yearlings dispersed to a new family group, while male yearlings dispersed to a bachelor group. Solitary males appeared during the interval between leaving bachelor or family groups, and vice versa. Our study showed that vicuña polygyny was based on female grazing area requirements. Females that stayed with the same males were defined as loyal, and had a mean 18.9 ha home range. Females that changed both to a different family and male; and grazed over the same home range size, were considered disloyal. Family groups selected the habitat that had a low stratum and the highest coverage of palatable plants, while bachelors used their habitat randomly.     

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.     

Poor horse traders: large mammals trade survival for reproduction during the process of feralization

We investigated density dependence on the demographic parameters of a population of Camargue horses (Equus caballus), individually monitored and unmanaged for eight years. We also analysed the contributions of individual demographic parameters to changes in the population growth rates. The decrease in resources caused a loss of body condition. Adult male survival was not affected, but the survival of foals and adult females decreased with increasing density. Prime-aged females maintained high reproductive performance at high density, and their survival decreased. The higher survival of adult males compared with females at high density presumably results from higher investment in reproduction by mares. The high fecundity in prime-aged females, even when at high density, may result from artificial selection for high reproductive performance, which is known to have occurred in all the major domestic ungulates. Other studies suggest that feral ungulates including cattle and sheep, as these horses, respond differently from wild ungulates to increases in density, by trading adult survival for reproduction. As a consequence, populations of feral animals should oscillate more strongly than their wild counterparts, since they should be both more invasive (as they breed faster), and more sensitive to harsh environmental conditions (as the population growth rate of long-lived species is consistently more sensitive to a given proportional change in adult survival than to the same change in any other vital rate). If this principle proves to be general, it has important implications for management of populations of feral ungulates.     

Seed dispersal by ungulates in the point calimere wildlife sanctuary: A scientific and perspective analysis

Exotic woody weed plants are a very serious threat to seed dispersed by ungulate in the tropical forest of Asia. The ungulates in Point Calimere Wildlife Sanctuary (PCWS) are a significant role in native indigenous seed dispersal. The exotic woody weed tree Prosopis juliflora prevalence distributed in the PCWS and they might potentially alter the native medicinal plant species. In the present investigation, we have assessed the seed dispersal by ungulates in PCWS from January to March 2017. Four different ungulate species were selected to understand their seed dispersal rate of different plant species in selected sanctuary. This investigation was planned to confirm the seed dispersal by ungulates of blackbuck, spotted deer, wild boar and feral horse. Among the four different ungulates tested, the maximum numbers of pellets collected from blackbuck and no seed found in their pellets. The low quantities of pellets were collected from wild boar and this study has recorded medium-sized ungulates which dispersed variety of plant. However, the dispersal of the seed of medicinal plants were not considerably high and relatively moderate percentage of seeds dispersal occurred in medium-sized ungulates like wild boar and spotted deer. P. juliflora had 100% seed germination rate were observed from the faecal samples of wild boar and feral horse. The control seed achieved maximum seedling rate than the ungulates seeds.     

Isolation and characterization of 10 polymorphic dinucleotide microsatellite markers for llama and guanaco

We isolated and characterized five polymorphic dinucleotide microsatellite markers from llama (Lama glama) and five from guanaco (Lama guanicoe). All loci were assayed on wild llamas and guanacos from Argentina, as all of the primers were able to amplify in both species.     

Physiological and behavioral indices of short-term stress in wild vicuñas (Vicugna vicugna) in Jujuy Province,Argentina

The management of wild vicuñas can trigger a stress response that may compromise welfare. In Santa Catalina, Jujuy Province, Argentina, indices of short-term stress associated with capture, handling, and shearing were studied in 105 wild vicuñas (Vicugna vicugna). The study included 2 groups (n = 59 and n = 46) of wild vicuñas captured in 2 consecutive days. Independent variables analyzed included sex, restraint time, and groups. Cortisol, creatine kinase, glucose, white blood cells, temperature, heart rate, and respiratory frequency were higher than published values. Respiratory rate increased during handling and correlated with holding time and group size, while heart rate decreased. Packed cell volume was higher in females. Cortisol concentrations differed between restraint groups and sex and inversely correlated with agonistic behavior. The most common behavior was increased vigilance. Sternal recumbency increased over holding time. During handling procedures, frequency of sudden movements like kicking and attempts to stand increased as restraint time increased. Females vocalized more than males. In conclusion, the methods used triggered measurable changes suggestive of short-term stress that appeared to be physiologically tolerated by the vicuñas.     

Bayesian Inference on the Effect of Density Dependence and Weather on a Guanaco Population from Chile

Understanding the mechanisms that drive population dynamics is fundamental for management of wild populations. The guanaco (Lama guanicoe) is one of two wild camelid species in South America. We evaluated the effects of density dependence and weather variables on population regulation based on a time series of 36 years of population sampling of guanacos in Tierra del Fuego, Chile. The population density varied between 2.7 and 30.7 guanaco/km², with an apparent monotonic growth during the first 25 years; however, in the last 10 years the population has shown large fluctuations, suggesting that it might have reached its carrying capacity. We used a Bayesian state-space framework and model selection to determine the effect of density and environmental variables on guanaco population dynamics. Our results show that the population is under density dependent regulation and that it is currently fluctuating around an average carrying capacity of 45,000 guanacos. We also found a significant positive effect of previous winter temperature while sheep density has a strong negative effect on the guanaco population growth. We conclude that there are significant density dependent processes and that climate as well as competition with domestic species have important effects determining the population size of guanacos, with important implications for management and conservation.     

Effects of capture,shearing, and release on the ecology and behavior of wild vicuñas

The goals of sustainable use of wildlife include minimizing long term deleterious impacts of management. We evaluated the ethoecological and demographic responses of wild vicuñas (Vicugna vicugna) to capture, shearing, and release in northern Argentina, as well as effects on behavior, by comparing captured and shorn versus non-captured animals for 2 yr after capture events. We observed subtle and short-term changes in the movement behavior of individuals, likely due to thermal and behavioral stress following capture. We did not observe changes in survival and birth rates, social organization, or distribution of animals following capture. Therefore, we concluded that the capture techniques used resulted in low impacts on the wild vicuña population. © 2011 The Wildlife Society.     

Habitat choice of Grevy's zebras (Equus grevyi) in Laikipia, Kenya

Characterizing habitat choice is essential for endangered species conservation. For the endangered Grevy's zebra ( Equus grevyi ), as with many widely ranging vertebrates, human activities may be an important factor affecting space use. Grevy's zebras are grazing ungulates inhabiting the savannahs of central-northern Kenya and Ethiopia. Past research on their social organization indicates that reproductive status shapes associations and movements. Here, we examine how habitat use varies across four reproductive classes: lactating and nonlactating females, bachelors and territorial males. We also test whether Grevy's zebra avoid locations close to active livestock corrals, or bomas . We find that forage quality, forage quantity and habitat openness of locations used by Grevy's zebra vary significantly depending on individual reproductive state. Lactating females and bachelors use areas with green, short grass and medium-dense bush more frequently than nonlactating females or territorial males. We hypothesize that lactating females trade off forage quantity and safety to access nutrients in growing grass. Across reproductive classes, Grevy's zebra choose locations further from active bomas than if they used the area randomly. Our results suggest that Grevy's zebra may require a range of vegetation characteristics for different reproductive classes. Further, they may need areas free from competition or disturbance by livestock.     

In vitro production of embryos in South American camelids

Studies in reproductive biotechnology techniques have been minimal in South American camelids (SAC). Complex reproductive characteristics of these species contribute to slow progress. Nevertheless, some techniques, such as in vitro fertilization, intracytoplasmic sperm injection and nuclear transfer have been applied and have produced advances in knowledge on embryo environment and in vitro conditions necessary for development. Embryo production may have a high impact in both domestic and wild camelids population. Studies addressed to improve in vitro embryo production and oocyte collection could be a potential key to develop IVF and embryo production as a routine procedure in camelids.     

Home Range Use and Movement Patterns of Non-Native Feral Goats in a Tropical Island Montane Dry Landscape

Advances in wildlife telemetry and remote sensing technology facilitate studies of broad-scale movements of ungulates in relation to phenological shifts in vegetation. In tropical island dry landscapes, home range use and movements of non-native feral goats (Capra hircus) are largely unknown, yet this information is important to help guide the conservation and restoration of some of the world’s most critically endangered ecosystems. We hypothesized that feral goats would respond to resource pulses in vegetation by traveling to areas of recent green-up. To address this hypothesis, we fitted six male and seven female feral goats with Global Positioning System (GPS) collars equipped with an Argos satellite upload link to examine goat movements in relation to the plant phenology using the Normalized Difference Vegetation Index (NDVI). Movement patterns of 50% of males and 40% of females suggested conditional movement between non-overlapping home ranges throughout the year. A shift in NDVI values corresponded with movement between primary and secondary ranges of goats that exhibited long-distance movement, suggesting that vegetation phenology as captured by NDVI is a good indicator of the habitat and movement patterns of feral goats in tropical island dry landscapes. In the context of conservation and restoration of tropical island landscapes, the results of our study identify how non-native feral goats use resources across a broad landscape to sustain their populations and facilitate invasion of native plant communities.     

Fragmentation of the Habitat of Wild Ungulates by Anthropogenic Barriers in Mongolia

Habitat loss and habitat fragmentation caused by anthropogenic activities are the main factors that constrain long-distance movement of ungulates. Mongolian gazelles (Procapra gutturosa) and Asiatic wild asses (Equus hemionus) in Mongolia are facing habitat fragmentation and loss. To better understand how their movements respond to potential anthropogenic and natural barriers, we tracked 24 Mongolian gazelles and 12 wild asses near the Ulaanbaatar–Beijing Railroad and the fenced international border between Mongolia and China between 2002 and 2012. None of the tracked gazelles crossed the railroad, even though gazelles were captured on both sides of the tracks at the start of the study. Similarly, we did not observe cross-border movements between Mongolia and China for either species, even though some animals used areas adjacent to the border. The both species used close areas to the anthropogenic barriers more frequently during winter than summer. These results suggest strong impacts by the artificial barriers. The construction of new railroads and roads to permit mining and other resource development therefore creates the threat of further habitat fragmentation, because the planned routes will divide the remaining non-fragmented habitats of the ungulates into smaller pieces. To conserve long-distance movement of the ungulates in this area, it will be necessary to remove or mitigate the barrier effects of the existing and planned roads and railroads and to adopt a landscape-level approach to allow access by ungulates to wide ranges throughout their distribution.     

Trophic niche overlap and wild ungulate consumption by red fox and wolf in a mountain area in Italy

Red fox (Vulpes vulpes) and wolf (Canis lupus) are two widespread opportunistic predators living in simpatry in many areas. Nonetheless, scarce information are available on their trophic interactions. We investigated food habits of these two carnivores in a mountain area in Italy and assessed the extent of their trophic niche overlap, focusing on the consumption of wild ungulates. Thereby we analyzed the content of 669 red fox scats and 253 wolf scats collected between May 2008 and April 2009. Red foxes resulted to have a more than three times higher niche breadth than wolves. Vegetables, small mammals, wild ungulates, and invertebrates were major items (altogether 92% of volume) of the red fox annual diet. On the contrary wolf annual diet relied on wild ungulates (94% of volume) with wild boar (Sus scrofa) being the main food item. The degree of trophic niche overlap between the two species was found to be low (Pianka's O = 0.356). Diet variation between the warm and the cold seasons was limited in both species, and higher in red fox than in wolf. The two canids appeared to use wild ungulates unevenly being the former more selective for younger preys, smaller in size (newborn piglets and roe deer Capreolus capreolus fawns), whereas the latter exhibited a preference for medium-sized and large ungulates (10–35 kg wild boar and adult roe deer). Even if wild ungulates represent the main shared food category, the different use of age/weight classes by the two predators, together with their possible consumption as carrions by red fox, suggests a very limited trophic competition between wolf and red fox.This study represents a contribution to the knowledge of trophic interaction in predator–prey systems where sympatric carnivores are present.