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Spatial Transposition Tasks in Indian Sloth Bears (Melursus Ursinus) and Bornean Sun Bears (Helarctos Malayanus Euryspilus)

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Spatial Transposition Tasks in Indian Sloth Bears (Melursus Ursinus) and Bornean Sun Bears (Helarctos Malayanus Euryspilus) Journal of Comparative Psychology © 2017 American Psychological Association 2017, Vol. 131, No. 4, 290–303 0735-7036/17/$12.00 http://dx.doi.org/10.1037/com0000077 Spatial Transposition Tasks in Indian Sloth Bears (Melursus ursinus) and Bornean Sun Bears (Helarctos malayanus euryspilus) Daniela Hartmann Marina Davila-Ross University of Portsmouth and Justus-Liebig University Gießen University of Portsmouth Siew Te Wong Josep Call Bornean Sun Bear Conservation Centre, Sabah, Malaysia Max Planck Institute for Evolutionary Anthropology, Leipzig, Germany, and University of St. Andrews Marina Scheumann Max Planck Institute for Evolutionary Anthropology, Leipzig, Germany, and University of Veterinary Medicine Hannover Spatial transposition tasks assess individuals’ ability to represent nonvisible spatial object displacements. Several nonhuman mammal species have been tested on this task including primates, cats, and dogs, but to date, great apes seem the only taxon that has repeatedly and consistently solved spatial transpo- sition tasks. The authors investigated the ability of captive sloth and sun bears to solve spatial transposition tasks. Both species belong to the same taxonomic group as cats and dogs, but unlike them and similar to apes, they have an omnivorous diet that requires them to keep track of fruit sources in space and time. The bears were first tested on a visible displacement task and those that succeeded were further tested on a spatial transposition task that involved a 180° transposition, followed by 2 tasks with two 360° transpositions. All 7 sloth bears and 7 out of 9 sun bears solved the visible displacement task. The 180° transposition task was solved by 6 out of 7 sloth bears and 1 out of the 5 tested sun bears. Three sloth bears were tested on all 4 experiments and even solved 2-chained 360° transpositions. Control conditions were conducted showing that the bears’ performance did not rely on olfactory or auditory cues. The results provide the first indication that bears might be able to track invisible objects. Further studies will be necessary to confirm these results and to control the influence of associative learning. The present study emphasizes the importance of including different animal species in the investigation of what underlies the evolution of different cognitive skills. Keywords: spatial cognition, bears, object permanence, comparative cognition, spatial memory Supplemental materials: http://dx.doi.org/10.1037/com0000077.supp In their natural environment, animals are confronted with a sources and nests, following social partners or keeping track of variety of spatial problems like remembering the location of food prey/predators even if they are temporarily occluded by an obsta- Editor’s Note. Gordon M. Burghardt served as the action editor for this allowing us to conduct this study. We are indebted to the zoo managers of This document is copyrighted by the American Psychological Association or one of its alliedarticle.—JC publishers. Leipzig Zoo, Dr. Junhold and Gerd Nötzold, for facilitating our task This article is intended solely for the personal use of the individual user and is not to be disseminated broadly. tremendously. We thank all the bear keepers of Leipzig Zoo as well as all the wonderful staff and interns of BSBCC who helped us during the course This article was published Online First June 26, 2017. of the experiments. Special thanks go to Christian Patzer for his patient Daniela Hartmann, Psychology Department, University of Portsmouth help and bear knowledge as well as Kate Bond for her support and valuable Department of Clinical Child, and Adolescent Psychology, Justus-Liebig suggestions. We also are very grateful to Sailun Aris for his kind support University Gießen; Marina Davila-Ross, Psychology Department, Univer- during the field study. Furthermore, we would like to acknowledge Sönke sity of Portsmouth; Siew Te Wong, Bornean Sun Bear Conservation von den Berg for his support in preparing the figures, Birgit Haßfurther and Centre, Sabah, Malaysia; Josep Call, Department of Comparative Psychol- Felix Vogel for the reliability coding, Frances Sherwood-Brock for proof- ogy, Max Planck Institute for Evolutionary Anthropology, Leipzig, Ger- reading the English and three anonymous reviewers for their comments on many, and School of Psychology and Neuroscience, University of St. a former version of the manuscript. The field study received ethics ap- Andrews; Marina Scheumann, Department of Comparative Psychology, proval from the Animal Welfare and Ethical Review Body of the Univer- Max Planck Institute for Evolutionary Anthropology, and Institute of sity of Portsmouth, United Kingdom. Zoology, University of Veterinary Medicine Hannover. Correspondence concerning this article should be addressed to Daniela We thank Leipzig Zoo, Sabah Biodiversity Center, Sabah Wildlife Hartmann, Haus C1, Raum C13, Otto-Behaghel-Straße 10C, D-35394 Department and the Bornean Sun Bear Conservation Center (BSBCC) for Gießen, Germany. E-mail: [email protected] 290 SPATIAL TRANSPOSITION TASK IN BEARS 291 cle. The ability to keep track of animate and inanimate objects for apes. She states that the performance of other animal species in moving in space is one way, perhaps the main way, to deal with invisible displacement tasks might be explained by three common such problems. Piaget’s (1954) pioneering work on the develop- problems: sensory cues, experimenter-given cues, and associative ment of object tracking in human infants has received considerable learning. In line with this, others have argued that the performance attention from a developmental (e.g., Sophian & Sage, 1983) and of dogs (e.g., Gagnon & Doré, 1992, 1993, 1994; Pasnak, comparative perspective (e.g., Tomasello & Call, 1997). Piaget Kurkjian, & Triana, 1988) can be explained by low-level associa- distinguished six stages characterizing the development of object tive rules rather than an understanding of object concepts (Collier- concept in human children (Piaget, 1954). At Stages 1 to 3, human Baker et al., 2004; Fiset & LeBlanc, 2007). In a reply, Pepperberg infants start to follow visible objects with their gaze. From Stage (2015) refuted Jaakkola’s critique and argued that the literature 4 onward, they seem to understand that an object still exists even provides sufficient evidence to show that Gray parrots demonstrate though it is no longer visible (object permanence). At Stage 4, 8- full object permanence. to 10-month-old infants can find a hidden object behind several It is interesting to note that all tested nonhuman mammalian screens but cannot overcome the A-not-B error, which means that species, with the exception of apes and some monkey species, have they continue to search under the screen where they found the failed the spatial transposition task (carnivora: Doré et al., 1996; object previously. At Stage 5, 12- to 16-month-old infants over- Fiset & Plourde, 2013; Rooijakkers et al., 2009; artiodactyla: come the A-not- B error and can find the object after multiple Albiach-Serrano, Brauer, Cacchione, Zickert, & Amici, 2012; visible object displacements. Reaching Stage 6, which is the high- Nawroth, von Borell, & Langbein, 2015; cetacea: Jaakkola, Gua- est level, means that 18- to 24-month-old infants have mastered the rino, Rodriguez, Erb, & Trone, 2010). It was argued that the poor invisible displacement task. Here the object is covered by a cup (or performance of dogs in the 180° rotation task might be due to hidden in the experimenter’s hand), moved in and out several conflicting contextual cues (the empty cup was placed where the screens (or containers) and left behind (inside) one of them. Once baited cup originally was; Miller, Rayburn-Reeves, & Zentall, the displacement is completed the now empty cup (hand) is shown 2009) while they were solving the 90° rotation tasks (Miller, to the infant who is then allowed to search for the object. Mastery Gipson, Vaughan, Rayburn-Reeves, & Zentall, 2009; Miller, of this stage implies that the infant understands that the object that Rayburn-Reeves, et al., 2009). However, Rooijakkers et al. (2009) was hidden under the cup moved out of sight (which is why this showed that dogs also failed the 180° spatial transposition task task is called invisible displacement) with the cup and that it is now when this conflict was avoided (by moving the cups to a new located behind one of the screens. location). Furthermore, Rooijakkers et al. (2009) demonstrated that A variation of the traditional invisible displacement task is the apes outperformed dogs in all tested spatial transposition tasks. spatial transposition task (e.g., Doré, Fiset, Goulet, Dumas, & Rooijakkers and colleagues (2009) argued that the poor perfor- Gagnon, 1996; Sophian, 1984; Sophian & Sage, 1983). In a mance in dogs might be explained by them having lost the cog- transposition task, the food is visibly hidden underneath one of at nitive skills to solve transposition tasks during the process of least two containers, which then switch positions (also known as domestication (domestication hypothesis). Contradicting this hy- the “shell game”). Interestingly, it was shown that human infants pothesis, a comparative study showed that dogs and wolves ex- were able to solve invisible displacement tasks of Stage 6 at 13 hibited similar search patterns when they were confronted with a months of age (Sophian & Sage, 1983) but solved spatial trans- transposition task (Fiset & Plourde, 2013). Interestingly,
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