And Blue and Gold Macaws (Ara Ararauna)
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Relationship Quality and Cognition in Orange-Winged Amazons (Amazona amazonica) and Blue and Gold Macaws (Ara ararauna) Alejandra Morales Picard PhD University of York Psychology March 2016 2 ABSTRACT Evidence of cognitive complexity in birds has led to the recognition that mammalian and avian species likely evolved comparable high-level cognitive capacities independently. One of the most significant findings that have emerged from this area of study is the identification of a key social trait that is found in species deemed to be intelligent – the presence of long-term, high-quality partnerships. Amongst birds, parrots (along with corvids) have shown relatively high levels of both cognitive and social complexity. However, relatively few parrot species have been the subject of empirical investigation. The original research presented in this thesis explores social behaviour and cognitive capacity in orange-winged Amazons (Amazona amazonica) and blue and gold macaws (Ara ararauna). Observational research findings revealed evidence of high-quality relationships in both species, and while levels of affiliative investment were similar in orange-winged Amazons (OWAs) and blue and gold macaws (BGMs), some potentially meaningful between species differences were found in courtship feeding, allopreening, and social tolerance. Experimental research findings revealed evidence of inhibitory control in both species (as measured by performance on a transparent cylinder task), though OWAs performed significantly better than BGMs on test trials. Social learning capacities were also found in OWAs through an open diffusion experiment. Both species showed poor performance in a means-end task, yielding no evidence of causal understanding. Birds also had difficulty acquiring the loose string task, which was aimed at testing cooperative problem solving. Poor performance of both OWAs and BGMs on these tasks is believed to have been primarily due to non-cognitive factors (e.g., motivation, motor difficulty of task). Affiliative investment and performance on the social learning task were found to be positively correlated in BGMs and there was some indication that individual variation in boldness may have been associated with inhibitory task performance in BGMs. 3 TABLE OF CONTENTS ABSTRACT…………………………………………………………………………….2 TABLE OF CONTENTS……………………………………………………………….3 LIST OF TABLES………………………………………………………………………7 LIST OF FIGURES……………………………………………………………..............9 PREFACE.......................................................................................................................11 ACKNOWLEDGMENTS..............................................................................................12 AUTHOR’S DECLARATION.......................................................................................13 CHAPTER 1: INTRODUCTION...................................................................................15 Complex cognition, convergence, and the origins of intelligence: Theoretical background................................................................................................................15 Historical context................................................................................................15 Theoretical approaches to comparative cognition..............................................17 The debate over complex cognition: What does it meant to be intelligent?.......18 Convergent evolution of complex cognition......................................................22 Cognitive convergence despite neuroanatomical divergence.............................24 Questions raised by evidence of cognitive convergence....................................25 Shared traits: Explaining why and how complex cognition evolved.................26 Sociality as a key factor in the emergence of complex cognition......................29 Re-thinking social intelligence hypotheses........................................................30 Limitations of brain size analyses.......................................................................35 Where do we go from here?................................................................................37 Parrots: an untapped resource.............................................................................37 Psittaciformes............................................................................................................37 Parrot phylogeny................................................................................................38 Psittaciform superfamilies..................................................................................39 Behaviour and socio-ecology.............................................................................41 The parrot-corvid comparison: Indicators of intelligence..................................43 The potential contribution of parrot research.....................................................45 CHAPTER 2: AVIAN PROBLEM-SOLVING CAPACITIES.....................................47 Avian cognition research in historical context..........................................................47 Solving problems in the physical realm.....................................................................49 Object permanence.............................................................................................50 4 Inhibitory control................................................................................................52 Concept formation..............................................................................................54 ‘Folk physics’ and understanding causality........................................................57 Solving problems in a social environment.................................................................64 Social learning....................................................................................................64 Cooperative problems solving............................................................................68 Conclusion.................................................................................................................69 CHAPTER 3: STUDY SPECIES...................................................................................71 Natural History..........................................................................................................72 Socio-ecology............................................................................................................74 Breeding..............................................................................................................75 In captivity.................................................................................................................76 Pet trade..............................................................................................................76 OWA research....................................................................................................77 BGM research.....................................................................................................80 Conclusion.................................................................................................................81 CHAPTER 4: RELATIONSHIP QUALITY..................................................................83 Introduction................................................................................................................85 Methods.....................................................................................................................96 Results......................................................................................................................111 Discussion................................................................................................................121 CHAPTER 5: PHYSICAL COGNITION....................................................................131 Introduction.............................................................................................................132 Study 1: Physical cognition.....................................................................................132 Introduction.......................................................................................................132 General methods...............................................................................................143 Experiment 1: Measuring inhibitory control using a cylinder task..................146 Experiment 2: Measuring inhibitory control using the A not B task................151 Experiment 3: Measuring means-end understanding using a string-pulling task....................................................................................................................156 Composite cognitive scores..............................................................................161 Study 1 discussion............................................................................................162 Study 2: Potential covariate of cognitive performance – affiliative investment and boldness...................................................................................................................168 5 Introduction.......................................................................................................168 Relationship quality and cognitive performance..............................................173 Boldness and cognitive performance................................................................175 Study 2 discussion............................................................................................181