Comparative cognition, as an interdisciplinary field, should utilize a holistic approach for studying cognitive mechanisms. We suggest that research with species of interest should employ both work with animals under human care and in the field. This complimentary approach allows for a better understanding of functional cognitive mechanisms themselves (i.e., comparative cognition regarding processes), and how these skill sets can relate to a particular species’ ecological niche. We suggest that research evidence for equivalence classification, learning by exclusion, and long-term memory in pinnipeds can provide a foundation for discussion and implementation of a two-pronged methodological approach utilizing ‘lab’ and field’ work. First, we describe evidence from research with pinnipeds under human care supporting each of these cognitive abilities, then follow this with evidence for implications of these mechanisms from complimentary field research. Lastly, we provide a brief discussion of implementation of a purposeful and two-pronged research approach as an understanding of pinnipeds’ high levels of cognitive flexibility may underlie their success for navigating the ever-changing, and often human-altered, natural environment.

Ecological learning is enriched when students can apply course content to their own communities and gain an understanding and experience for deploying sustainable practices. Such merging of content, application, and place requires an interdisciplinary approach to grapple with the ecological and cultural issues more holistically that sustainability education explores. Guided by the ecological pedagogy of David Orr, founder of the Oberlin Project, we devised a pilot study that combines course content in environmental science, cultural studies, and writing with place-based, experiential, and hands-on learning to empower students to critically analyze their lifestyles and engage in activism for change.

Identifying perceptual thresholds is critical for understanding the mechanisms that underlie signal evolution. Using computer-animated stimuli, we examined visual speed sensitivity in the Jacky dragon Amphibolurus muricatus, a species that makes extensive use of rapid motor patterns in social communication. First, focal lizards were tested in discrimination trials using random-dot kinematograms displaying combinations of speed, coherence, and direction. Second, we measured subject lizards' ability to predict the appearance of a secondary reinforcer (1 of 3 different computer-generated animations of invertebrates: cricket, spider, and mite) based on the direction of movement of a field of drifting dots by following a set of behavioural responses (e.g., orienting response, latency to respond) to our virtual stimuli. We found an effect of both speed and coherence, as well as an interaction between these 2 factors on the perception of moving stimuli. Overall, our results showed that Jacky dragons have acute sensitivity to high speeds. We then employed an optic flow analysis to match the performance to ecologically relevant motion. Our results suggest that the Jacky dragon visual system may have been shaped to detect fast motion. This pre-existing sensitivity may have constrained the evolution of conspecific displays. In contrast, Jacky dragons may have difficulty in detecting the movement of ambush predators, such as snakes and of some invertebrate prey. Our study also demonstrates the potential of the computer-animated stimuli technique for conducting nonintrusive tests to explore motion range and sensitivity in a visually mediated species.

Rapid technical advances in the field of computer animation (CA) and virtual reality (VR) have opened new avenues in animal behavior research. Animated stimuli are powerful tools as they offer standardization, repeatability, and complete control over the stimulus presented, thereby "reducing" and "replacing" the animals used, and "refining" the experimental design in line with the 3Rs. However, appropriate use of these technologies raises conceptual and technical questions. In this review, we offer guidelines for common technical and conceptual considerations related to the use of animated stimuli in animal behavior research. Following the steps required to create an animated stimulus, we discuss (I) the creation, (II) the presentation, and (III) the validation of CAs and VRs. Although our review is geared toward computer-graphically designed stimuli, considerations on presentation and validation also apply to video playbacks. CA and VR allow both new behavioral questions to be addressed and existing questions to be addressed in new ways, thus we expect a rich future for these methods in both ultimate and proximate studies of animal behavior.

It is well established that recognition of complex acoustic signals, such as bird song, is dependent upon the temporal ordering of signal units or syntax. Much less is known about functionally analogous visual displays. The jacky dragon (Amphibolurus muricatus) is a native Australian agamid lizard with a highly stereotyped visual display made up of three discrete motor patterns. We conducted a playback experiment using high-resolution computer animations of conspecifics to test the importance of temporal order for signal efficacy. Lizards were shown three different life-sized simulated animations of conspecific differing in their skin texture and morphology signatures ranging from highly natural to abnormal. We evaluated signal recognition and assessed the relative importance of syntax and morphology. Our results showed that signal recognition is highly sensitive to syntax and this largely determines the observers' behavioural responses. Stimuli with abnormal texture and shape were highly effective, as long as the natural order of motor patterns was preserved. Display recognition in jacky lizards hence depends upon syntax in just the same way as temporally constrained signals in other modalities.

Despite variations of environmental noise, signals are designed to be effective and conspicuous over an appreciable distance. In particular, visual signals must be perceptible against interference caused by natural elements, such as windblown vegetation. We examined the efficiency of aggressive and submissive displays to elicit behavioural responses from observers in the Jacky dragon (Amphibolurus muricatus) across relative environmental noise. Both displays have been reported to play an important role during social interactions in this species. We conducted two video playback experiments that utilised a high-resolution computer-generated lizard animation to produce social displays that were embedded within simulated windblown vegetation. First, we compared the efficiency a full aggressive display action pattern (comprised of a tail-flick, backward-forward arm wave, and push-up body rock) to a slow arm wave submissive display, against identical background of windblown vegetation. Second, we compared the tail-flick (alerting component) to the slow arm wave across three varying natural conditions, in which the vegetation behind the displays acted as simulated noise: calm, typical, and windy. We found that aggressive displays were more efficient to elicit an observer's response than submissive signals. Furthermore, the tail-flick display is more efficient than submissive displays across a range of natural variation in windblown vegetation movement, but both signals remain efficient in the face of environmental motion noise. Our results suggest that constraints from the environmental background scene play may have a critical role in the evolution of signal design used in Jacky dragon communication.

Design characteristics of signals, such as their duration, may have evolved to maximize signal efficiency. It is commonly assumed that constraints on signal design have usually shaped the most optimal display characteristics to improve signal transmission and information transfer of the signaller, and detection by intended receivers. In this study, we tested whether the characteristics (duration, speed and frequency) of an aggressive display, the push-up body rock, exhibited by the Jacky dragon (Amphibolurus muricatus) have likely evolved for optimal signal efficiency, as it is able to draw attention to the signaller. We performed two video playback experiments using high-resolution 3D animations testing the effect of variation in push-up body rock structure. In experiment 1, we manipulated push-up body rock display structure. We gradually increased the number of push-ups exhibited by a digitally animated Jacky dragon increasing the overall display duration. In experiment 2, we developed four stimuli based on population-typical push-up body rock display for duration (short and long), and frequency of push-ups (1 or 5 consecutive push-ups) by manipulating push-ups speed. In both experiments, we measured the probability of an orienting response and response latency of focal lizards when being exposed to the different stimuli. Our results showed that display duration is critically important for signal efficiency in the aggressive push-up body rock display. If we are to understand the design characteristics of signals used in animal communication, then it appears important to consider the possible trade-off between signal efficiency and costs.

Traditionally, studies that explored animal communication have been directed towards the observation of natural interactions between individuals. Over the years, researchers have long championed the use of artificial stimuli in place of natural ones in behavioral experiments to precisely control what the observers get to see or experience. The employment of diverse techniques to stage animal interactions has provided an alternative to observations and intrusive experimental methods. Technological advances now allow researchers to develop realistic computer animations of social partners that mimic behaviors with a high degree of fidelity for morphological and behavioral characteristics of tutors. The increasing use of the computer-generated animations technique reveals a desire to deliver standardized visual stimuli and to limit the variable behavior of demonstrators across experimental sessions. In the following review, we provide an appraisal of the computer-generated animations efficiency to stage animal interactions, and consider experimental studies in which this technique has been employed to simulate social interactions. We also present alternative methods that are used for designing animation models. Our aim is to evaluate the merits of computer-generated animations and how this technique may be more appropriate for certain types of staged interactions when compared to other classically employed approaches. We advocate that computer-generated animations appear to be the most flexible technique to date, and offers better control of visual cues that are presented, thus allowing researchers to program a large variety of stimuli. Finally, we suggest improvements of this technique, and especially how it may be used to study signal design in multimodal systems.

The complex ritualized displays of males in many territorial species suggest that selection has shaped male behaviors in ways that affect fitness. In this study, we evaluated the link between display behavior during male–male interactions and reproductive success in the Australian jacky dragon ( Amphibolurus muricatus ), a lizard species that uses a complex series of movement patterns for communication. We quantified variation in male display behaviors by using video playback experiments in the laboratory, and subsequently assessed variation in male reproductive success by paternity analyses of offspring. Because the lizards used in this study came from eggs incubated under three thermal environments, we also could evaluate the impact of developmental temperature on adult behavior and reproductive success. Incubation temperature had a strong effect on male reproductive success; males produced under intermediate temperatures sired more offspring than those produced under extreme developmental temperatures. However, incubation temperature did not affect male display behavior, nor was male behavior associated with reproductive success. Our findings do not support the common assumption that display behaviors used during male–male interactions affect reproductive success.

By investigating the mechanisms that underlie the perception of environmental cues, we may begin to understand how the sensory system governs behavioral responses. This is the first empirical study to examine learning and visual sensitivity in a reptile species, the tuatara (Sphenodon punctatus). We established a non-intrusive psychophysical method by employing an instrumental paradigm in order to examine discrimination learning and the ability to distinguish different flicker frequencies in the tuatara. Seventeen tuatara were trained under an operant conditioning task to respond to various discriminative stimuli flickering between 2.65 and 65.09 Hz. Tuatara were able to learn the operant task and discriminate between a constant light and flicker frequency rates between 2.65 and 45.61 Hz, but not at 65.09 Hz. We demonstrated a reliable psychophysical method where these reptiles could learn a basic operant task and discriminate visual stimuli in the form of flicker frequency rates. The tuatara's ability to perceive flickering light is comparable to that of avian, mammalian, and other reptilian species. This method is thus suitable for more comprehensive examinations of vision and additional sensory abilities in other reptiles.

The evolution of movement-based signals is constrained by the successful segmentation of relevant movements from motion noise by the visual system of receivers. We tested five Jacky dragons, Amphibolurus muricatus, a species characterized by stereotyped movement-based social signals, for their ability to discriminate the direction of drifting dots against simulated background motion noise of varying angular speeds. Results from trials with artificial (dot) backgrounds suggested that Jacky dragons are most sensitive to high-speed movement, while background movement of similar angular speeds to target stimuli reduced performance. We also found differences in accuracy between natural backgrounds comprising footage of plant motion at two independent sites. We argue that detecting salient movement depends on the particular characteristics of the surrounding motion noise, and discuss its implications for movement-based signal design. (C) 2008 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

The increasing use of the video playback technique in behavioural ecology reveals a growing need to ensure better control of the visual stimuli that focal animals experience. Technological advances now allow researchers to develop computer-generated animations instead of using video sequences of live-acting demonstrators. However, care must be taken to match the motion characteristics (speed and velocity) of the animation to the original video source. Here, we presented a tool based on the use of an optic flow analysis program to measure the resemblance of motion characteristics of computer-generated animations compared to videos of live-acting animals. We examined three distinct displays (tail-flick (TF), push-up body rock (PUBR), and slow arm wave (SAW)) exhibited by animations of Jacky dragons (Amphibolurus muricatus) that were compared to the original video sequences of live lizards. We found no significant differences between the motion characteristics of videos and animations across all three displays. Our results showed that our animations are similar the speed and velocity features of each display. Researchers need to ensure that similar motion characteristics in animation and video stimuli are represented, and this feature is a critical component in the future success of the video playback technique.

Testing sensory characteristics on herpetological species has been difficult due to a range of properties related to physiology, responsiveness, performance ability, and the type of reinforcer used. Using the Jacky lizard as a model, we outline a successfully established procedure in which to test the visual sensitivity to motion characteristics. We incorporated modifications to traditional operant paradigms by using three video playback systems to deliver random-dot kinematogram motion stimuli coupled with salient computer-animated secondary reinforcers representative of biologically important appetitive stimuli. This procedure has the capacity to test other visual aspects in lizards as well as other nonhuman species using video playback and computer-animation techniques as experimental tools.

Communication between animals is diverse and complex. Animals may communicate using auditory, seismic, chemosensory, electrical, or visual signals. In particular, understanding the constraints on visual signal design for communication has been of great interest. Traditional methods for investigating animal interactions have used basic observational techniques, staged encounters, or physical manipulation of morphology. Less intrusive methods have tried to simulate conspecifics using crude playback tools, such as mirrors, still images, or models. As technology has become more advanced, video playback has emerged as another tool in which to examine visual communication (Rosenthal, 2000). However, to move one step further, the application of computer-animation now allows researchers to specifically isolate critical components necessary to elicit social responses from conspecifics, and manipulate these features to control interactions. Here, I provide detail on how to create an animation using the Jacky dragon as a model, but this process may be adaptable for other species. In building the animation, I elected to use Lightwave 3D to alter object morphology, add texture, install bones, and provide comparable weight shading that prevents exaggerated movement. The animation is then matched to select motor patterns to replicate critical movement features. Finally, the sequence must rendered into an individual clip for presentation. Although there are other adaptable techniques, this particular method had been demonstrated to be effective in eliciting both conspicuous and social responses in staged interactions.

Testing visual sensitivity in any species provides basic information regarding behaviour, evolution, and ecology. However, testing specific features of the visual system provide more empirical evidence for functional applications. Investigation into the sensory system provides information about the sensory capacity, learning and memory ability, and establishes known baseline behaviour in which to gauge deviations (Burghardt, 1977). However, unlike mammalian or avian systems, testing for learning and memory in a reptile species is difficult. Furthermore, using an operant paradigm as a psychophysical measure of sensory ability is likewise as difficult. Historically, reptilian species have responded poorly to conditioning trials because of issues related to motivation, physiology, metabolism, and basic biological characteristics. Here, I demonstrate an operant paradigm used a novel model lizard species, the Jacky dragon (Amphibolurus muricatus) and describe how to test peripheral sensitivity to salient speed and motion characteristics. This method uses an innovative approach to assessing learning and sensory capacity in lizards. I employ the use of random-dot kinematograms (RDKs) to measure sensitivity to speed, and manipulate the level of signal strength by changing the proportion of dots moving in a coherent direction. RDKs do not represent a biologically meaningful stimulus, engages the visual system, and is a classic psychophysical tool used to measure sensitivity in humans and other animals. Here, RDKs are displayed to lizards using three video playback systems. Lizards are to select the direction (left or right) in which they perceive dots to be moving. Selection of the appropriate direction is reinforced by biologically important prey stimuli, simulated by computer-animated invertebrates.