Beyond the Tricks: The Science and Comparative Cognition of Magic.

Magic is an art form that has fascinated humans for centuries. Recently, the techniques used by magicians to make their audience experience the impossible have attracted the attention of psychologists, who, in just a couple of decades, have produced a large amount of research regarding how these effects operate, focusing on the blind spots in perception and roadblocks in cognition that magic techniques exploit. Most recently, this investigation has given a pathway to a new line of research that uses magic effects to explore the cognitive abilities of nonhuman animals. This new branch of the scientific study of magic has already yielded new evidence illustrating the power of magic effects as a psychological tool for nonhuman animals. This review aims to give a thorough overview of the research on both the human and nonhuman perception of magic effects by critically illustrating the most prominent works of both fields of inquiry.

Octopus toss-up: Is debris throwing driven by intent?

In a noteworthy observation, Godfrey-Smith and colleagues report the first evidence of debris throwing in wild octopuses, including instances where they target conspecifics. Proposing parallels with behaviours observed in select social mammals, this discovery prompts inquiries into the extent of their similarity and the potential role of cognition.

Cephalopod learning and memory.

Cephalopod molluscs are renowned for their unique central nervous system - a donut-shaped brain organised around the oesophagus. This brain supports sophisticated learning and memory abilities. Between the 1950s and 1980s, these cognitive abilities were extensively studied in octopus (Figure 1A) - a now leading model for the study of memory and its neural substrates (approximately 200 papers during this period). The focus on octopus learning and memory was mainly due to their curious nature and the fact that they adapt to laboratory-controlled conditions, making them easy to test and maintain in captivity. Research on cephalopod cognition began to widen in the late 20th century, when scientists started focusing on other coleoid cephalopods (i.e., cuttlefish and squid) (Figure 1B,C), and not just on associative learning and memory per se, but other more complex aspects of cognition such as episodic-like memory (the ability to remember the what, where, and when of a past event), source memory (the retrieval of contextual details from a memory), and self-control (the ability to inhibit an action in the present to gain a more valuable future reward). Attention broadened further over the last two decades to focus on the shelled cephalopods - the nautiloids (Figure 1D). The nautiloids have relatively primitive brains compared to their soft-bodied cousins (octopus, cuttlefish, and squid) but research shows that they are still able to comparatively succeed in some cognitive tasks. In this primer, we will provide a general description of the types of memory studied in cephalopods, and discuss learning and memory experiments that address the main challenges cephalopods face during their daily lives: navigation, timing, and food selection. Determining the type of information cephalopods learn and remember and whether they use such information to overcome ecological challenges will highlight why these invertebrates evolved large and sophisticated brains.

The Inner Lives of Cephalopods.

The minds of cephalopods have captivated scientists for millennia, yet the extent that we can understand their subjective experiences remains contested. In this article, we consider the sum of our scientific progress towards understanding the inner lives of cephalopods. Here, we outline the behavioral responses to specific experimental paradigms that are helping us to reveal their subjective experiences. We consider evidence from three broad research categories, which help to illuminate whether soft-bodied cephalopods (octopus, cuttlefish, and squid) have an awareness of self, awareness of others, and an awareness of time. Where there are current gaps in the literature, we outline cephalopod behaviors that warrant experimental investigation. We argue that investigations, especially framed through the lens of comparative psychology, have the potential to extend our understanding of the inner lives of this extraordinary class of animals.

Waiting for a better possibility: delay of gratification in corvids and its relationship to other cognitive capacities.

Self-control, the ability to resist temptation and wait for better but delayed possibilities, is an important cognitive skill that underpins decision-making and planning. The capacity to exert self-control has been linked to intelligence in humans, chimpanzees and most recently cuttlefish. Here, we presented 10 Eurasian jays, Garrulus glandarius, with a delayed maintenance task, which measured the ability to choose a preferred outcome as well as the ability to sustain the delay prior to that outcome. Jays were able to wait for better possibilities, but maximum wait times varied across the subjects. We also presented them with five cognitive tasks that assessed spatial memory, spatial relationships and learning capacity. These tasks are commonly used as measures of general intelligence within an ecological context. Individual performance was correlated across the cognitive tasks, which suggests that there was a general intelligence factor underlying their performance. Performance in these tasks was correlated significantly with the jays' capacity to wait for better possibilities. This study demonstrates that self-control and intelligence are correlated in jays. The fact that this correlation exists in diverse species suggests that self-control is a fundamental feature of cognition. Our results are discussed in the context of convergent evolution. This article is part of the theme issue 'Thinking about possibilities: mechanisms, ontogeny, functions and phylogeny'.

Jays are sensitive to cognitive illusions.

Jays hide food caches, steal them from conspecifics and use tactics to minimize cache theft. Jays are sensitive to the content of their own caches, retrieving items depending on their preferences and the perishability of the cached item. Whether jays impose the same content sensitivity when they steal caches is less clear. We adapted the 'cups-and-balls' magic routine, creating a cognitive illusion to test whether jays are sensitive to the (i) content of hidden items and (ii) type of displacement. Subjects were presented with two conditions in which hidden food was consistent with their expectations; and two conditions in which food was manipulated to violate their expectations by switching their second preferred food for their preferred food (up-value) or vice versa (de-value). Subjects readily accepted food when it was consistent with their expectations but were more likely to re-inspect the baited cup and alternative cup when their expectations were violated. In the de-value condition, jays exhibited longer latencies to consume the food and often rejected it. Dominant subjects were more likely to reject the food, suggesting that social factors influence their responses to cognitive illusions. Using cognitive illusions offers innovative avenues for investigating the psychological constraints in diverse animal minds.

Episodic-like memory is preserved with age in cuttlefish.

Episodic memory, remembering past experiences based on unique what-where-when components, declines during ageing in humans, as does episodic-like memory in non-human mammals. By contrast, semantic memory, remembering learnt knowledge without recalling unique what-where-when features, remains relatively intact with advancing age. The age-related decline in episodic memory likely stems from the deteriorating function of the hippocampus in the brain. Whether episodic memory can deteriorate with age in species that lack a hippocampus is unknown. Cuttlefish are molluscs that lack a hippocampus. We test both semantic-like and episodic-like memory in sub-adults and aged-adults nearing senescence (n = 6 per cohort). In the semantic-like memory task, cuttlefish had to learn that the location of a food resource was dependent on the time of day. Performance, measured as proportion of correct trials, was comparable across age groups. In the episodic-like memory task, cuttlefish had to solve a foraging task by retrieving what-where-when information about a past event with unique spatio-temporal features. In this task, performance was comparable across age groups; however, aged-adults reached the success criterion (8/10 correct choices in consecutive trials) significantly faster than sub-adults. Contrary to other animals, episodic-like memory is preserved in aged cuttlefish, suggesting that memory deterioration is delayed in this species.

Exploring the perceptual inabilities of Eurasian jays (Garrulus glandarius) using magic effects.

In recent years, scientists have begun to use magic effects to investigate the blind spots in our attention and perception [G. Kuhn, Experiencing the Impossible: The Science of Magic (2019); S. Macknik, S. Martinez-Conde, S. Blakeslee, Sleights of Mind: What the Neuroscience of Magic Reveals about Our Everyday Deceptions (2010)]. Recently, we suggested that similar techniques could be transferred to nonhuman animal observers and that such an endeavor would provide insight into the inherent commonalities and discrepancies in attention and perception in human and nonhuman animals [E. Garcia-Pelegrin, A. K. Schnell, C. Wilkins, N. S. Clayton, Science 369, 1424-1426 (2020)]. Here, we performed three different magic effects (palming, French drop, and fast pass) to a sample of six Eurasian jays (Garrulus glandarius). These magic effects were specifically chosen as they utilize different cues and expectations that mislead the spectator into thinking one object has or has not been transferred from one hand to the other. Results from palming and French drop experiments suggest that Eurasian jays have different expectations from humans when observing some of these effects. Specifically, Eurasian jays were not deceived by effects that required them to expect an object to move between hands when observing human hand manipulations. However, similar to humans, Eurasian jays were misled by magic effects that utilize fast movements as a deceptive action. This study investigates how another taxon perceives the magician's techniques of deception that commonly deceive humans.

Buoys with looming eyes deter seaducks and could potentially reduce seabird bycatch in gillnets.

Bycatch of seabirds in gillnet fisheries is a global conservation issue with an estimated 400 000 seabirds killed each year. To date, no underwater deterrents trialled have consistently reduced seabird bycatch across operational fisheries. Using a combination of insights from land-based strategies, seabirds' diving behaviours and their cognitive abilities, we developed a floating device exploring the effect of large eyespots and looming movement to prevent vulnerable seabirds from diving into gillnets. Here, we tested whether this novel above-water device called 'Looming eyes buoy' (LEB) would consistently deter vulnerable seaducks from a focal area. We counted the number of birds present in areas with and without LEBs in a controlled experimental setting. We show that long-tailed duck Clangula hyemalis abundance declined by approximately 20-30% within a 50 m radius of the LEB and that the presence of LEBs was the most important variable explaining this decline. We found no evidence for a memory effect on long-tailed ducks but found some habituation to the LEB within the time frame of the project (62 days). While further research is needed, our preliminary trials indicate that above-water visual devices could potentially contribute to reduce seabird bycatch if appropriately deployed in coordination with other management measures.

Cuttlefish exert self-control in a delay of gratification task.

The ability to exert self-control varies within and across taxa. Some species can exert self-control for several seconds whereas others, such as large-brained vertebrates, can tolerate delays of up to several minutes. Advanced self-control has been linked to better performance in cognitive tasks and has been hypothesized to evolve in response to specific socio-ecological pressures. These pressures are difficult to uncouple because previously studied species face similar socio-ecological challenges. Here, we investigate self-control and learning performance in cuttlefish, an invertebrate that is thought to have evolved under partially different pressures to previously studied vertebrates. To test self-control, cuttlefish were presented with a delay maintenance task, which measures an individual's ability to forgo immediate gratification and sustain a delay for a better but delayed reward. Cuttlefish maintained delay durations for up to 50-130 s. To test learning performance, we used a reversal-learning task, whereby cuttlefish were required to learn to associate the reward with one of two stimuli and then subsequently learn to associate the reward with the alternative stimulus. Cuttlefish that delayed gratification for longer had better learning performance. Our results demonstrate that cuttlefish can tolerate delays to obtain food of higher quality comparable to that of some large-brained vertebrates.

Cephalopods: Ambassadors for rethinking cognition.

Traditional approaches in comparative cognition have a long history of focusing on a narrow range of vertebrate species. However, in recent years the range of model species has expanded. Despite this development, invertebrate taxa are still largely neglected in comparative cognition, which limits our ability to locate the origins of cognitive traits. The time has come to rethink cognition and develop a more comprehensive understanding of cognitive evolution by expanding comparative analyses to include a diverse range of invertebrate taxa. In this review, we contend that cephalopods are suitable ambassadors for rethinking cognition. Cephalopods have large complex brains, exhibit sophisticated behavioral traits, and increasing evidence suggests that they possess complex cognitive abilities once thought to be unique to large-brained vertebrates. Comparing cephalopods with vertebrates, whose cognition has evolved independently, provides prominent opportunities to circumvent current limitations in comparative cognition that have arisen from traditional vertebrate comparisons. Increased efforts in investigating the cognitive abilities of cephalopods have also led to important welfare-related improvements. These large-brained molluscs are paving the way for a more inclusive approach to investigating cognitive evolution that we hope will extend to other invertebrate taxa.

How intelligent is a cephalopod? Lessons from comparative cognition.

The soft-bodied cephalopods including octopus, cuttlefish, and squid are broadly considered to be the most cognitively advanced group of invertebrates. Previous research has demonstrated that these large-brained molluscs possess a suite of cognitive attributes that are comparable to those found in some vertebrates, including highly developed perception, learning, and memory abilities. Cephalopods are also renowned for performing sophisticated feats of flexible behaviour, which have led to claims of complex cognition such as causal reasoning, future planning, and mental attribution. Hypotheses to explain why complex cognition might have emerged in cephalopods suggest that a combination of predation, foraging, and competitive pressures are likely to have driven cognitive complexity in this group of animals. Currently, it is difficult to gauge the extent to which cephalopod behaviours are underpinned by complex cognition because many of the recent claims are largely based on anecdotal evidence. In this review, we provide a general overview of cephalopod cognition with a particular focus on the cognitive attributes that are thought to be prerequisites for more complex cognitive abilities. We then discuss different types of behavioural flexibility exhibited by cephalopods and, using examples from other taxa, highlight that behavioural flexibility could be explained by putatively simpler mechanisms. Consequently, behavioural flexibility should not be used as evidence of complex cognition. Fortunately, the field of comparative cognition centres on designing methods to pinpoint the underlying mechanisms that drive behaviours. To illustrate the utility of the methods developed in comparative cognition research, we provide a series of experimental designs aimed at distinguishing between complex cognition and simpler alternative explanations. Finally, we discuss the advantages of using cephalopods to develop a more comprehensive reconstruction of cognitive evolution.

Dimensions of Animal Consciousness.

How does consciousness vary across the animal kingdom? Are some animals 'more conscious' than others? This article presents a multidimensional framework for understanding interspecies variation in states of consciousness. The framework distinguishes five key dimensions of variation: perceptual richness, evaluative richness, integration at a time, integration across time, and self-consciousness. For each dimension, existing experiments that bear on it are reviewed and future experiments are suggested. By assessing a given species against each dimension, we can construct a consciousness profile for that species. On this framework, there is no single scale along which species can be ranked as more or less conscious. Rather, each species has its own distinctive consciousness profile.

Cuttlefish show flexible and future-dependent foraging cognition.

Some animals optimize their foraging activity by learning and memorizing food availability, in terms of quantity and quality, and adapt their feeding behaviour accordingly. Here, we investigated whether cuttlefish flexibly adapt their foraging behaviour according to the availability of their preferred prey. In Experiment 1, cuttlefish switched from a selective to an opportunistic foraging strategy (or vice versa) when the availability of their preferred prey at night was predictable versus unpredictable. In Experiment 2, cuttlefish exhibited day-to-day foraging flexibility, in response to experiencing changes in the proximate future (i.e. preferred prey available on alternate nights). In Experiment 1, the number of crabs eaten during the day decreased when shrimp (i.e. preferred food) were predictably available at night, while the consumption of crabs during the day was maintained when shrimp availability was unpredictable. Cuttlefish quickly shifted from one strategy to the other, when experimental conditions were reversed. In Experiment 2, cuttlefish only reduced their consumption of crabs during the daytime when shrimps were predictably available the following night. Their daytime foraging behaviour appeared dependent on shrimps' future availability. Overall, cuttlefish can adopt dynamic and flexible foraging behaviours including selective, opportunistic and future-dependent strategies, in response to changing foraging conditions.

Cephalopod cognition.

Cephalopods have captivated the minds of scientists for thousands of years, dating back to approximately 330 BC when Aristotle became fascinated by their ability to rapidly change colour. This remarkable ability, however, is not the only aspect of cephalopod behaviour that has garnered attention from the scientific community. The soft-bodied cephalopods (henceforth cephalopods), namely octopus, cuttlefish, and squid, are widely considered to be the most cognitively advanced group of invertebrates. They possess highly developed perceptual, memory, and spatial learning abilities and are also capable of intriguing feats of behaviour that appear to indicate complex cognition.

Fighting and mating success in giant Australian cuttlefish is influenced by behavioural lateralization.

Behavioural lateralization is widespread. Yet, a fundamental question remains, how can lateralization be evolutionary stable when individuals lateralized in one direction often significantly outnumber individuals lateralized in the opposite direction? A recently developed game theory model predicts that fitness consequences which occur during intraspecific interactions may be driving population-level lateralization as an evolutionary stable strategy. This model predicts that: (i) minority-type individuals exist because they are more likely to adopt unpredictable fighting behaviours during competitive interactions (e.g. fighting); and (ii) majority-type individuals exist because there is a fitness advantage in having their biases synchronized with other conspecifics during interactions that require coordination (e.g. mating). We tested these predictions by investigating biases in giant Australian cuttlefish during fighting and mating interactions. During fighting, most male cuttlefish favoured the left eye and these males showed higher contest escalation; but minority-type individuals with a right-eye bias achieved higher fighting success. During mating interactions, most male cuttlefish favoured the left eye to inspect females. Furthermore, most male cuttlefish approached the female's right side during a mating attempt and these males achieved higher mating success. Our data support the hypothesis that population-level biases are an evolutionary consequence of the fitness advantages involved in intraspecific interactions.

Grow Smart and Die Young: Why Did Cephalopods Evolve Intelligence?

Intelligence in large-brained vertebrates might have evolved through independent, yet similar processes based on comparable socioecological pressures and slow life histories. This convergent evolutionary route, however, cannot explain why cephalopods developed large brains and flexible behavioural repertoires: cephalopods have fast life histories and live in simple social environments. Here, we suggest that the loss of the external shell in cephalopods (i) caused a dramatic increase in predatory pressure, which in turn prevented the emergence of slow life histories, and (ii) allowed the exploitation of novel challenging niches, thus favouring the emergence of intelligence. By highlighting convergent and divergent aspects between cephalopods and large-brained vertebrates we illustrate how the evolution of intelligence might not be constrained to a single evolutionary route.