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.

1. In a recently published paper, Oliva et al. concluded that domestic grazing pressure across Patagonian rangelands approached carrying capacity due to decades of stock adjustment, but that guanaco overpopulation may have altered that balance. The authors argued that unless guanaco numbers are controlled, they will reduce forage available for domestic stock and will negatively affect rangelands. We consider that the herbivore-stock analysis presented is inaccurate and deserves revision, and that the stated conclusions lack empirical support. 2. When the spatial distribution of herbivores is accounted for in the Oliva et al. analysis, domestic stock is far above carrying capacity, indicating that domestic overgrazing continues. 3. Theoretical and empirical evidence on bottom-up regulation and competitive exclusion challenges the supposed guanaco overpopulation and the hypothetical reduction of forage available for livestock. 4. Even if guanaco numbers are reduced, grassland degradation and production losses will continue because their main drivers, domestic overstock and heterogeneous grazing, are still operating. 5. Synthesis and applications. Oversimplified models with poor ecological insight can lead to erroneous conclusions and misguide management decisions. The incorrect inference that Patagonian domestic stock is adjusted to carrying capacity could help to consolidate current domestic overgrazing by reducing incentives to improve livestock management practices. Regarding guanacos, a controversial species in an unfavourable context, control-oriented harvest without a clear justification threatens populations' viability and genuine attempts of productive diversification. Addressing relevant ecological processes, such as niche partitioning, competitive exclusion and population regulation, is essential to correctly assess joint carrying capacity in multi-herbivore systems, as well as to identify the true factors driving degradation processes and to optimize rangeland use on a sustainable basis.

Response of guanacos Lama guanicoe to changes in land management in Península Valdé s, Argentine Patagonia: conservation implications M a r í a V i r g i n i a B u r g i , A n d r e a M a r i n o , M a r í a V i c t o r i a R o d r í g u e z G u s t a v o P a z o s and R i c a r d o B a l d iAbstract The guanaco Lama guanicoe was the only large native herbivore widely distributed across Patagonia until the introduction of domestic sheep Ovis aries. Guanacos have declined because of competition with sheep for forage, high hunting pressure and habitat degradation. Península Valdés is a protected area where sheep ranching is the predominant activity. A ranch formerly dedicated to sheep production was converted into a private wildlife reserve, from which all the sheep were removed in 2005. We studied changes in guanaco abundance inside and outside the reserve after sheep removal, and also plant cover of various vegetation types. We found that guanaco abundance was higher inside than outside the reserve, and increased by three-fold within 3 years. Total plant cover and grass cover were higher inside than outside the reserve. Our results showed that guanacos reacted rapidly to changes in management, and suggest that even at a high density guanacos would not be as damaging to the vegetation as sheep. Although management changes resulted in significant changes in guanaco abundance locally, the size of a protected area influences the persistence of wild populations. A large herbivore such as the guanaco needs to be managed across large areas. We believe it is necessary to implement a management plan for Península Valdés that allows for the coexistence of sustainable livestock production and healthy wildlife populations.

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.

Sheep rearing is the main productive activity in Patagonian rangelands, where guanacos are the only native ungulate. Ranchers perceive a decrease in range carrying capacity as guanaco numbers increase, therefore guanaco conservation within private lands becomes a considerable challenge. This issue is particularly evident in the World Natural Heritage Península Valdés (PV), where there is a need to harmonize livestock production and biodiversity conservation. While sheep rearing prevails as the primary land use in the area, some ecotourism initiatives have been implemented to complement livestock production. In order to study how land use affected guanaco distribution, we characterized PV's ranches in terms of land subdivision, primary productivity, stocking-rate and management type, and assess how these variables affected guanaco encounter rates. Smaller ranches were composed of smaller paddocks (mean size 4.8 km2), which showed higher values of the remote-sensing derived Enhance Vegetation Index (EVI) (mean 0.14) and held higher sheep densities (mean 108.0 sheep/km2), while larger management units (mean size 23.8 km2), showed lower EVI values (mean 0.12) and lower stocking-rates (mean 36.7 sheep/km2). This pattern suggests that primary productivity has been a decisive factor to determine the minimal paddock size set by ranchers in PV, apparently precluding excessive land-subdivision in less productive areas. Guanaco encounter rate, expressed as number of guanacos per travelled kilometre, was inversely related to EVI and stocking-rate. However, land subdivision was the better predictor of guanaco encounter-rate within only sheep ranches, finding more guanacos per kilometre as paddock size increased. In contrast, in ranches where ecotourism was implemented as a complementary activity, guanaco encounter-rates were greater, regardless of paddock size. Our results suggest that the implementation of an additional activity by which landowners derive benefits from wildlife has prompted a beneficial outcome for guanacos, presumably through a decrease in harassment intensity. Finally, we propose possible mechanisms by which land subdivision may affect guanaco distribution and potential alternatives for the inclusion of wildlife conservation in a context of extensive livestock production.