Past and Recent Effects of Livestock Activity on the Genetic Diversity and Population Structure of Native Guanaco Populations of Arid Patagonia.

Extensive livestock production and urbanization entail modifications of natural landscapes, including installation of fences, development of agriculture, urbanization of natural areas, and construction of roads and infrastructure that, together, impact native fauna. Here, we evaluate the diversity and genetic structure of endemic guanacos (Lama guanicoe) of the Monte and Patagonian Steppe of central Argentina, which have been reduced and displaced by sheep ranching and other impacts of human activities. Analyses of genetic variation of microsatellite loci and d-loop revealed high levels of genetic variation and latitudinal segregation of mitochondrial haplotypes. There were indications of at least two historical populations in the Monte and the Patagonian Steppe based on shared haplotypes and shared demographic history among localities. Currently, guanacos are structured into three groups that were probably reconnected relatively recently, possibly facilitated by a reduction of sheep and livestock in recent decades and a recovery of the guanaco populations. These results provide evidence of the genetic effects of livestock activity and urbanization on wild herbivore populations, which were possibly exacerbated by an arid environment with limited productive areas. The results highlight the importance of enacting conservation management plans to ensure the persistence of ancestral and ecologically functional populations of guanacos.

Choosing what is left: the spatial structure of a wild herbivore population within a livestock-dominated landscape.

Shrublands and grasslands comprise over 30% of the land surface and are among the most exploited ecosystems for livestock production. Across natural landscapes, the distribution and abundance of wild herbivores are affected by interspecific competition for foraging resources, hunting and the development of infrastructure among other factors. In Argentine Patagonia, the abundance of domestic sheep grazing on native vegetation outnumbers the widely distributed guanaco (Lama guanicoe) and sheep ranching monopolizes the most productive lands. In this work, we aimed to assess the spatial variation in the abundance of guanacos in Península Valdés, a representative landscape of Patagonia, investigating the incidence of natural and human-related factors. We conducted ground surveys during the austral autumn in 2017 totaling 383.4 km along areas with and without sheep ranching. We built density surface models to account for the variation in guanaco abundance and obtained a map of guanaco density at a resolution of 4 km2. We estimated an overall density of 11.71 guanacos.km-2 for a prediction area of 3,196 km2, although the density of guanacos tripled in areas where sheep ranching was terminated (in around 20% of the surface of Península Valdés) compared to areas with sheep. Guanacos were more abundant at lower values of primary productivity and sheep stocking rates and further from inhabited ranch buildings, suggesting competition with sheep and conflict with humans. Although guanacos selected open, grass-dominated habitats across sheep-free sites, fences dividing properties and paddocks played a significant role in the spatial structure of their population in Península Valdés affecting negatively the abundance of guanacos. Our results indicate that actions to improve habitat connectivity for guanacos, favor the coexistence among guanacos and sheep ranching, and promote responsible human activities and attitudes towards wildlife are needed.

Modeling the spatial structure of the endemic mara (Dolichotis patagonum) across modified landscapes.

Across modified landscapes, anthropic factors can affect habitat selection by animals and consequently their abundance and distribution patterns. The study of the spatial structure of wild populations is crucial to gain knowledge on species' response to habitat quality, and a key for the design and implementation of conservation actions. This is particularly important for a low-density and widely distributed species such as the mara (Dolichotis patagonum), a large rodent endemic to Argentina across the Monte and Patagonian drylands where extensive sheep ranching predominates. We aimed to assess the spatial variation in the abundance of maras and to identify the natural and anthropic factors influencing the observed patterns in Península Valdés, a representative landscape of Patagonia. We conducted ground surveys during the austral autumn from 2015 to 2017. We built density surface models to account for the variation in mara abundance, and obtained a map of mara density at a resolution of four km2. We estimated an overall density of 0.93 maras.km-2 for the prediction area of 3,476 km2. The location of ranch buildings, indicators of human presence, had a strong positive effect on the abundance of maras, while the significant contribution of the geographic longitude suggested that mara density increases with higher rainfall. Although human presence favored mara abundance, presumably by providing protection against predators, it is likely that the association could bring negative consequences for maras and other species. The use of spatial models allowed us to provide the first estimate of mara abundance at a landscape scale and its spatial variation at a high resolution. Our approach can contribute to the assessment of mara population abundance and the factors shaping its spatial structure elsewhere across the species range, all crucial attributes to identify and prioritize conservation actions.

Ecological drivers of guanaco recruitment: variable carrying capacity and density dependence.

Ungulates living in predator-free reserves offer the opportunity to study the influence of food limitation on population dynamics without the potentially confounding effects of top-down regulation or livestock competition. We assessed the influence of relative forage availability and population density on guanaco recruitment in two predator-free reserves in eastern Patagonia, with contrasting scenarios of population density. We also explored the relative contribution of the observed recruitment to population growth using a deterministic linear model to test the assumption that the studied populations were closed units. The observed densities increased twice as fast as our theoretical populations, indicating that marked immigration has taken place during the recovery phase experienced by both populations, thus we rejected the closed-population assumption. Regarding the factors driving variation in recruitment, in the low- to medium-density setting, we found a positive linear relationship between recruitment and surrogates of annual primary production, whereas no density dependence was detected. In contrast, in the high-density scenario, both annual primary production and population density showed marked effects, indicating a positive relationship between recruitment and per capita food availability above a food-limitation threshold. Our results support the idea that environmental carrying capacity fluctuates in response to climatic variation, and that these fluctuations have relevant consequences for herbivore dynamics, such as amplifying density dependence in drier years. We conclude that including the coupling between environmental variability in resources and density dependence is crucial to model ungulate population dynamics; to overlook temporal changes in carrying capacity may even mask density dependence as well as other important processes.

Ecological correlates of group-size variation in a resource-defense ungulate, the sedentary guanaco.

For large herbivores, predation-risk, habitat structure and population density are often reported as major determinants of group size variation within and between species. However, whether the underlying causes of these relationships imply an ecological adaptation or are the result of a purely mechanistic process in which fusion and fragmentation events only depend on the rate of group meeting, is still under debate. The aim of this study was to model guanaco family and bachelor group sizes in contrasting ecological settings in order to test hypotheses regarding the adaptive significance of group-size variation. We surveyed guanaco group sizes within three wildlife reserves located in eastern Patagonia where guanacos occupy a mosaic of grasslands and shrublands. Two of these reserves have been free from predators for decades while in the third, pumas often prey on guanacos. All locations have experienced important changes in guanaco abundance throughout the study offering the opportunity to test for density effects. We found that bachelor group size increased with increasing density, as expected by the mechanistic approach, but was independent of habitat structure or predation risk. In contrast, the smaller and territorial family groups were larger in the predator-exposed than in the predator-free locations, and were larger in open grasslands than in shrublands. However, the influence of population density on these social units was very weak. Therefore, family group data supported the adaptive significance of group-size variation but did not support the mechanistic idea. Yet, the magnitude of the effects was small and between-population variation in family group size after controlling for habitat and predation was negligible, suggesting that plasticity of these social units is considerably low. Our results showed that different social units might respond differentially to local ecological conditions, supporting two contrasting hypotheses in a single species, and highlight the importance of taking into account the proximate interests and constraints to which group members may be exposed to when deriving predictions about group-size variation.

The impact of conservation on the status of the world's vertebrates.

Using data for 25,780 species categorized on the International Union for Conservation of Nature Red List, we present an assessment of the status of the world's vertebrates. One-fifth of species are classified as Threatened, and we show that this figure is increasing: On average, 52 species of mammals, birds, and amphibians move one category closer to extinction each year. However, this overall pattern conceals the impact of conservation successes, and we show that the rate of deterioration would have been at least one-fifth again as much in the absence of these. Nonetheless, current conservation efforts remain insufficient to offset the main drivers of biodiversity loss in these groups: agricultural expansion, logging, overexploitation, and invasive alien species.

The status of the world's land and marine mammals: diversity, threat, and knowledge.

Knowledge of mammalian diversity is still surprisingly disparate, both regionally and taxonomically. Here, we present a comprehensive assessment of the conservation status and distribution of the world's mammals. Data, compiled by 1700+ experts, cover all 5487 species, including marine mammals. Global macroecological patterns are very different for land and marine species but suggest common mechanisms driving diversity and endemism across systems. Compared with land species, threat levels are higher among marine mammals, driven by different processes (accidental mortality and pollution, rather than habitat loss), and are spatially distinct (peaking in northern oceans, rather than in Southeast Asia). Marine mammals are also disproportionately poorly known. These data are made freely available to support further scientific developments and conservation action.

Guanacos and sheep: evidence for continuing competition in arid Patagonia.

Guanacos (Lama guanicoe) are the only wild ungulate species widely distributed across the Patagonian steppe and have undergone a precipitous population decline since the introduction of domestic sheep (Ovis aries) 100 years ago. There has been speculation that sheep ranching may have played a major role in guanaco population decline after monopolising the most productive land because of competition for forage plants. Our aim in this work was to estimate guanaco abundance and account for its variation across nine different sites, two seasons and two years. We conducted over 100 ground surveys of both guanacos and sheep and estimated the availability of the preferred plant species both animal species might select in their diet. We found that (1) sheep densities were up to 23 times higher than guanaco densities in sympatric conditions, (2) at a protected site without sheep, guanaco density was one order of magnitude higher than at the rest of the sites, (3) across nine different sites, sheep densities alone accounted for around 60% of the variation in guanaco abundance, (4) guanaco densities were negatively related to both total plant cover and availability of the preferred plant species in their diet, which were both positively associated with sheep density, and (5) within-site changes in guanaco densities between seasons and years were negatively related to changes in sheep densities. Our results are consistent with predictions on interspecific competition for food resources, although we cannot rule out possible effects of other human-related activities influencing guanaco abundance. We conclude that sheep compete with guanacos for forage in arid Patagonia.