Recently, there has been a surge in cognition research using non-avian reptile systems. As a diverse group of animals, non-avian reptiles [turtles, the tuatara, crocodylians, and squamates (lizards, snakes and amphisbaenids)] are good model systems for answering questions related to cognitive ecology, from the role of the environment on the brain, behaviour and learning, to how social and life-history factors correlate with learning ability. Furthermore, given their variable social structure and degree of sociality, studies on reptiles have shown that group living is not a pre-condition for social learning. Past research has demonstrated that non-avian reptiles are capable of more than just instinctive reactions and basic cognition. Despite their ability to provide answers to fundamental questions in cognitive ecology, and a growing literature, there have been no recent systematic syntheses of research in this group. Here, we systematically, and comprehensively review studies on reptile learning. We identify 92 new studies investigating learning in reptiles not included in previous reviews on this topic - affording a unique opportunity to provide a more in-depth synthesis of existing work, its taxonomic distribution, the types of cognitive domains tested and methodologies that have been used. Our review therefore provides a major update on our current state of knowledge and ties the collective evidence together under nine umbrella research areas: (i) habituation of behaviour, (ii) animal training through conditioning, (iii) avoiding aversive stimuli, (iv) spatial learning and memory, (v) learning during foraging, (vi) quality and quantity discrimination, (vii) responding to change, (viii) solving novel problems, and (ix) social learning. Importantly, we identify knowledge gaps and propose themes which offer important future research opportunities including how cognitive ability might influence fitness and survival, testing cognition in ecologically relevant situations, comparing cognition in invasive and non-invasive populations of species, and social learning. To move the field forward, it will be immensely important to build upon the descriptive approach of testing whether a species can learn a task with experimental studies elucidating causal reasons for cognitive variation within and among species. With the appropriate methodology, this young but rapidly growing field of research should advance greatly in the coming years providing significant opportunities for addressing general questions in cognitive ecology and beyond.

中文翻译：

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非鸟类爬行动物学习 40 年：进展和有希望的新方向

最近，使用非鸟类爬行动物系统进行的认知研究激增。作为一个多样化的动物群体，非鸟类爬行动物 [乌龟、大蜥蜴、鳄鱼和有鳞动物（蜥蜴、蛇和两栖动物）] 是回答与认知生态学相关问题的良好模型系统，从环境对大脑的作用来看，行为和学习，社会和生活史因素如何与学习能力相关。此外，鉴于爬行动物的社会结构和社会性程度多变，对爬行动物的研究表明，群居并不是社会学习的先决条件。过去的研究表明，非鸟类爬行动物不仅具有本能反应和基本认知能力。尽管他们有能力为认知生态学的基本问题提供答案，而且文献越来越多，该组最近没有系统的研究综合。在这里，我们系统地、全面地回顾了关于爬行动物学习的研究。我们确定了 92 项调查爬行动物学习的新研究，这些研究未包含在之前关于该主题的评论中——提供了一个独特的机会来更深入地综合现有工作、其分类分布、测试的认知领域类型和已使用的方法. 因此，我们的审查提供了我们当前知识状态的重大更新，并将集体证据联系在九个总括研究领域下：(i) 行为习惯，(ii) 通过条件反射训练动物，(iii) 避免厌恶刺激，(iv)空间学习和记忆，(v) 在觅食过程中学习，(vi) 质量和数量歧视，(vii) 对变化做出反应，(viii) 解决新问题，以及 (ix) 社会学习。重要的是，我们确定了知识差距并提出了提供重要未来研究机会的主题，包括认知能力如何影响健康和生存、在生态相关情况下测试认知、比较入侵和非入侵物种种群的认知以及社会学习。为了推动该领域向前发展，建立在测试一个物种是否可以学习任务的描述性方法的基础上，通过实验研究阐明物种内部和物种之间认知变化的因果关系将是非常重要的。通过适当的方法，