Perception of optical illusions in ungulates: insights from goats, sheep, guanacos and llamas.

Optical illusions have long been used in behavioural studies to investigate the perceptual mechanisms underlying vision in animals. So far, three studies have focused on ungulates, providing evidence that they may be susceptible to some optical illusions, in a way similar to humans. Here, we used two food-choice tasks to study susceptibility to the Müller-Lyer and Delboeuf illusions in 17 captive individuals belonging to four ungulate species (Lama guanicoe, Lama glama, Ovis aries, Capra hircus). At the group level, there was a significant preference for the longer/larger food over the shorter/smaller one in control trials. Additionally, the whole group significantly preferred the food stick between two inward arrowheads over an identical one between two outward arrowheads in experimental trials of the Müller-Lyer task, and also preferred the food on the smaller circle over an identical one on the larger circle in the experimental trials of the Delboeuf task. Group-level analyses further showed no significant differences across species, although at the individual level we found significant variation in performance. Our findings suggest that, in line with our predictions, ungulates are overall susceptible to the Müller-Lyer and the Delboeuf illusions, and indicate that the perceptual mechanisms underlying size estimation in artiodactyls might be similar to those of other species, including humans.

Spatial perseveration error by alpacas (Vicugna pacos) in an A-not-B detour task.

Spatial perseveration has been documented for domestic animals such as mules, donkeys, horses and dogs. However, evidence for this spatial cognition behavior among other domestic species is scarce. Alpacas have been domesticated for at least 7000 years yet their cognitive ability has not been officially reported. The present article used an A-not-B detour task to study the spatial problem-solving abilities of alpacas (Vicugna pacos) and to identify the perseveration errors, which refers to a tendency to maintain a learned route, despite having another available path. The study tested 51 alpacas, which had to pass through a gap at one end of a barrier in order to reach a reward. After one, two, three or four repeats (A trials), the gap was moved to the opposite end of the barrier (B trials). In contrast to what has been found in other domestic animals tested with the same task, the present study did not find clear evidence of spatial perseveration. Individuals' performance in the subsequent B trials, following the change of gap location, suggests no error persistence in alpacas. Results suggest that alpacas are more flexible than other domestic animals tested with this same task, which has important implications in planning proper training for experimental designs or productive purposes. These results could contribute toward enhancing alpacas' welfare and our understanding of their cognitive abilities.

The effects of poaching and habitat structure on anti-predator behavioral strategies: A guanaco population in a high cold desert as case study.

The effects of poaching on wildlife have been widely studied in conservation biology and can be heterogeneous, particularly on ungulates. These effects can be estimated through different methodologies whose use depends on several conditions such as Flight-initiation distance (FID). Our objectives were: 1- to evaluate whether poaching affects the FID and group structure of a guanaco (Lama guanicoe) population in a high cold desert in San Juan (Argentina); 2- to assess whether habitat structure (slope and vegetation cover) influences FID and group structure in this population. The study area included a site with poaching (unprotected area), and a site without poaching (protected area). We recorded 100 groups of guanacos: 70 in the protected and 30 in the unprotected area. FID and group size were greater in the unprotected than in the protected area, whereas proportions of group categories (with offspring, without offspring and solitary) were similar between areas. Besides, in relation to habitat structure, FID increased when vegetation cover decreased. On the other hand, FID and group size were not affected by slope. Our study shows that guanacos respond to poaching pressure as do other ungulate species, and that other factors such as vegetation cover also affect this behavior. Managers should be aware when interpreting FID due to its relation to habitat structure; the guanaco appears to assume greater risk (lower FID) in areas with high vegetation cover.

Hearing in alpacas (Vicugna pacos): audiogram, localization acuity, and use of binaural locus cues.

Behavioral audiograms and sound localization abilities were determined for three alpacas (Vicugna pacos). Their hearing at a level of 60 dB sound pressure level (SPL) (re 20 µPa) extended from 40 Hz to 32.8 kHz, a range of 9.7 octaves. They were most sensitive at 8 kHz, with an average threshold of -0.5 dB SPL. The minimum audible angle around the midline for 100-ms broadband noise was 23°, indicating relatively poor localization acuity and potentially supporting the finding that animals with broad areas of best vision have poorer sound localization acuity. The alpacas were able to localize low-frequency pure tones, indicating that they can use the binaural phase cue, but they were unable to localize pure tones above the frequency of phase ambiguity, thus indicating complete inability to use the binaural intensity-difference cue. In contrast, the alpacas relied on their high-frequency hearing for pinna cues; they could discriminate front-back sound sources using 3-kHz high-pass noise, but not 3-kHz low-pass noise. These results are compared to those of other hoofed mammals and to mammals more generally.

Social structure in a family group of Guanaco (Lama guanicoe, Ungulate): is female hierarchy based on 'prior attributes' or 'social dynamics'?

Social life involves costs and benefits mostly associated with how individuals interact with each other. The formation of hierarchies inside social groups has evolved as a common strategy to avoid high costs stemming from social interactions. Hierarchical relationships seem to be associated with different features such as body size, body condition and/or age, which determine dominance ability ('prior attributes' hypothesis). In contrast, the 'social dynamic' hypothesis suggests that an initial social context is a determinant in the formation of the hierarchy, more so than specific individual attributes. Hierarchical rank places individuals in higher positions, which presumably increases resource accessibility to their benefit, including opportunities for reproduction. We evaluate the maintenance of hierarchy in a family group of guanacos (Lama guanicoe) and evaluate the possible mechanisms involved in the stability of these interactions and their consequences. We estimate the linearity of social hierarchy and their dynamics. We find evidence of the formation of a highly linear hierarchy among females with males positioned at the bottom of the hierarchy. This hierarchy is not affected by physical characteristics or age, suggesting that it is established only through intra-group interactions. Rank is not related with calves' weight gain either; however, subordinated females, with lower rank, exhibit higher rates of allosuckling. We found no evidence of hierarchical structure in calves suggesting that hierarchical relationship in guanacos could be established during the formation of the family group. Hence, our results suggest that hierarchical dynamics could be related more to social dynamics than to prior attributes. We finally discuss the importance of hierarchies established by dominance and their role in minimizing social costs of interactions.

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.

Gradual training of alpacas to the confinement of metabolism pens reduces stress when normal excretion behavior is accommodated.

Confinement in metabolism pens may provoke a stress response in alpacas that will reduce the welfare of the animal and jeopardize the validity of scientific results obtained in such pens. In this study, we tested a protocol designed to successfully train alpacas to be held in a specially designed metabolism pen so that the animals' confinement would not jeopardize their welfare. We hypothesized that the alpacas would show fewer behaviors associated with a response to stress as training gradually progressed, and that they would adapt to being in the confinement of the metabolism pen. The training protocol was successful at introducing alpacas to the metabolism pens, and it did reduce the incidence of behavioral responses to stress as the training progressed. The success of the training protocol may be attributed to the progressive nature of the training, the tailoring of the protocol to suit alpacas, and the use of positive reinforcement. This study demonstrated that both animal welfare and the validity of the scientific outcomes could be maximized by the gradual training of experimental animals, thereby minimizing the stress imposed on the animals during experimental procedures.

Llama handling and training.

This article offers insights into the relationship of llama owners to their animals and the role of veterinarians as part of the animal care team. The effect of human behavior and handling techniques on llama behavior and marketability are discussed. Progressive ideas for nonforceful llama handling equipment, procedures, and training ideas are outlined in detail. Included are specific training plans for routine herd management chores such as injections and toenail trimming. This article is useful for both veterinarians and llama owners.

A hierarchical model of the evolution of cooperation in cultural systems.

In this paper the following problem is addressed: "Under what conditions can a collection of individual organisms learn to cooperate when cooperation appears to outwardly degrade individual performance at the outset. In order to attempt a theoretical solution to this problem, data from a real world problem in anthropology is used. A distributed simulation model of this system was developed to assess its long term behavior using using an approach suggested by Zeigler (Zeigler, B.P., 1984, Multifaceted Modelling and Discrete Event Simulation (Academic Press, London)). The results of the simulation are used to show that although cooperation degrades the performance potential of each individual, it enhances the persistence of the individual's partial solution to the problem in certain situations."