Abstract: Most horses are trained by negative reinforcement. Currently, however, no standardised test for evaluating horses' negative reinforcement learning ability is available. The aim of this study was to develop an objective test to investigate negative reinforcement learning in horses. Twenty-four Icelandic horses (3 years old) were included in this study. The horses were tested in a pressure-release task on three separate days with 10, 7 and 5 trials on each side, respectively. Each trial consisted of pressure being applied on the hindquarter with an algometer. The force of the pressure was increased until the horse moved laterally away from the point of pressure. There was a significant decrease in required force over trials on the first test day (P<0.001), but not the second and third day. The intercepts on days 2 and 3 differed significantly from day 1 (P<0.001), but not each other. Significantly stronger force was required on the right side compared to the left (P<0.001), but there was no difference between first and second side tested (P=0.56). Individual performance was evaluated by median-force and the change in force over trials on the first test day. These two measures may explain different characteristics of negative reinforcement learning. In conclusion, this study presents a novel, standardised test for evaluating negative reinforcement learning ability in horses.

Abstract: Summary Reliable assessment of animal populations is a long-standing challenge in wildlife ecology. Technological advances have led to widespread adoption of camera traps (CTs) to survey wildlife distribution, abundance and behaviour. As for any wildlife survey method, camera trapping must contend with sources of sampling error such as imperfect detection. Early applications focused on density estimation of naturally marked species, but there is growing interest in broad-scale CT surveys of unmarked populations and communities. Nevertheless, inferences based on detection indices are controversial, and the suitability of alternatives such as occupancy estimation is debatable. We reviewed 266 CT studies published between 2008 and 2013. We recorded study objectives and methodologies, evaluating the consistency of CT protocols and sampling designs, the extent to which CT surveys considered sampling error, and the linkages between analytical assumptions and species ecology. Nearly two-thirds of studies surveyed more than one species, and a majority used response variables that ignored imperfect detection (e.g. presence?absence, relative abundance). Many studies used opportunistic sampling and did not explicitly report details of sampling design and camera deployment that could affect conclusions. Most studies estimating density used capture?recapture methods on marked species, with spatially explicit methods becoming more prominent. Few studies estimated density for unmarked species, focusing instead on occupancy modelling or measures of relative abundance. While occupancy studies estimated detectability, most did not explicitly define key components of the modelling framework (e.g. a site) or discuss potential violations of model assumptions (e.g. site closure). Studies using relative abundance relied on assumptions of equal detectability, and most did not explicitly define expected relationships between measured responses and underlying ecological processes (e.g. animal abundance and movement). Synthesis and applications. The rapid adoption of camera traps represents an exciting transition in wildlife survey methodology. We remain optimistic about the technology's promise, but call for more explicit consideration of underlying processes of animal abundance, movement and detection by cameras, including more thorough reporting of methodological details and assumptions. Such transparency will facilitate efforts to evaluate and improve the reliability of camera trap surveys, ultimately leading to stronger inferences and helping to meet modern needs for effective ecological inquiry and biodiversity monitoring.

Abstract: ABSTRACT Reciprocity is probably the most debated of the evolutionary explanations for cooperation. Part of the confusion surrounding this debate stems from a failure to note that two different processes can result in reciprocity: partner control and partner choice. We suggest that the common observation that group-living animals direct their cooperative behaviours preferentially to those individuals from which they receive most cooperation is to be interpreted as the result of the sum of the two separate processes of partner control and partner choice. We review evidence that partner choice is the prevalent process in primates and propose explanations for this pattern. We make predictions that highlight the need for studies that separate the effects of partner control and partner choice in a broader variety of group-living taxa.

Abstract: We analysed the willingness of blue foxes (Vulpes lagopus) to work for and utilise five resources: a platform, wooden block, sand floor, nest box and empty space. Ten juvenile blue fox males were housed singly in apparatus consisting of three cages connected with one-way doors through the walls in between the cages and subjected to work for each of the five resources, one at a time. The resource was placed in one of the outermost cages of the apparatus. Force needed to open the door leading to the resource cage was increased daily by 0.25 or 0.5kg. The number of daily entries, visit durations and interaction with the resource were recorded on workloads of 0, 0.5, 1.5, 2.5, 3.5, 5, 6.5, and 8kg of extra weight. The latency to start interacting with the resource after entering the resource cage was measured on a workload of 3.5kg. The mean number of daily entries in the resource and the other outermost, i.e. control cage varied from 7 to 28 and from 17 to 44, respectively. The increasing workload decreased the number of entries in the resource cage, increased those in the control cage (Linear Mixed Model: F1,638=79.5, P<0.001) and lengthened the visit durations in both cages (F1,642=7.2, P<0.01). The foxes made most (F4,643=9.0, P<0.001) and shortest (F4,641=2.8, P<0.05) visits to the outermost cages when the available resource was either a platform or empty space. The visit durations were longest when the available resource was a nest box. The foxes interacted regularly with the wooden block, but five foxes were not observed interacting with the platform. The nest box was utilised approximately 50% of the time spent in the resource cage, while the platform was utilised only 1-6% and wooden block 2-17% of the time. The mean latency to start interacting with the resource after entering the resource cage was shortest for the sand floor (8s) and longest for the platform (113s, F3,335=26.3, P<0.001). The results show that the foxes re-scheduled their activities on increasing workloads in the apparatus. Based on the number of entries and visit durations, blue foxes valued the wooden block, nest box and sand floor more than the platform or an empty cage. After entering the resource cage, the foxes started interacting fastest with the sand floor, showing high motivation to interact. After entering the resource cage, the foxes make use of the roof of the nest box more urgently than the interior of the nest box. Long bouts in the cage with nest box indicate resting behaviour.

Abstract: To understand the relative benefits of social and personal information use in foraging decisions, we developed an agent-based model of social learning that predicts social information should be more adaptive where resources are highly variable and personal information where resources vary little. We tested our predictions with bumblebees and found that foragers relied more on social information when resources were variable than when they were not. We then investigated whether socially salient cues are used preferentially over non-social ones in variable environments. Although bees clearly used social cues in highly variable environments, under the same conditions they did not use non-social cues. These results suggest that bumblebees use a 'copy-when-uncertain' strategy.