Anglia Ruskin Thesis 2019 Template
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ANGLIA RUSKIN UNIVERSITY FACULTY OF SCIENCE AND ENGINEERING BIRDS IN THE NEANDERTHAL ECOLOGICAL NICHE – USING BIRDS FROM PLEISTOCENE CAVE ARCHIVES TO CHARACTERISE THE RELATIONSHIP BETWEEN NEANDERTHALS AND BIRDS STEWART FINLAYSON A thesis in partial fulfilment of the requirements of Anglia Ruskin University for the degree of Doctor of Philosophy Submitted: October 2019 To my sons Aiden and Tyler No matter how complicated life gets, always believe in yourselves. …You can do it. To Mum and Dad For always believing in me, when everyone else gave up ii Acknowledgements Back in 1989, when I was six years old, I first walked down with my parents to Gorham’s Cave and explored this wonderful site that had been occupied by Neanderthals in Gibraltar for thousands of years. It was an abrupt introduction into this world of the Neanderthals. In parallel, my parents also introduced me to the magical world of birds and their migrations across the Strait of Gibraltar. When in 1993 I first stepped into the Biological Reserve of Doñana in Southern Spain, my life changed – it was like stepping into the world of the Neanderthals. Many of the animals and plants I knew from fossils in the caves, suddenly came to life. I must therefore start by thanking these Neanderthal people, and the birds for allowing me to peek into their world. I also have to thank my parents who introduced me into the world of the naturalist, and encouraged me and have been my mentors throughout. My granparents: Tommy, Angie, Ernest and Maite, and my uncles Tom and John for your positive influence. A very special thanks to my principal supervisor, Dawn Hawkins, who gave me the opportunity embark on this thesis and believed in me and my work. She has been a pillar of support throughout, and has encouraged me even in the most difficult moments. Dawn has been a perfect guide through the process which has culminated in this thesis and cannot thank her enough. Phillip Pugh has provided me with the statistical and theoretical support and I’m grateful for his patience whilst going over multivariate statistics with me. It’s always easier when you have such a naturally gifted teacher, and I will always enjoy statistics thanks to him. Darren Fa co-supervised this thesis. I have known him since my early days in the caves in Gibraltar, and he has always been there to provide advice and give me the benefit of his knowledge. More than a supervisor, he has been a friend throughout. Nancy Harrison co-supervised this thesis at the early stages, and I particularly enjoyed my chats with her on all matters sea-birds, a passion that we share. iii I would like to acknowledge the support received from the Gibraltar National Museum, who also funded this thesis, and the Gorham’s Cave Complex, UNESCO World Heritage Site Office for allowing me access to the sites and collections. I would also like to thank my friends and colleagues at the Gibraltar National Museum, for their support throughout. In particular, I would like to thank Francisco Giles Guzman, Tyson Lee Holmes, and Sue Davies. Among the many researchers with whom I have shared many wonderful moments in the field and in the lab, I would like to highlight Nick Barton, Ruth Blasco, Andy Currant, Francisco Giles Pacheco, Jose Maria Gutierrez Lopez, Carlos Ibañez, Javier Juste, Richard Jennings, Alex Menez, Juan Jose Negro, Cristina Reinoso del Rio, Joaquin Rodriguez Vidal, Jordi Rossell, Larry Sawchuk, James Shipman, Chris Stringer. I would like to thank my good friends Alicia Montado and Giovanni Santini in particular for their cheerful support, no matter how peculiar the hours I worked and the places I travelled to. I wish to record my acknowledgement of the Victorian naturalists who I met through their writings, which inspired me through the outset: Howard Irby, William Willoughby Cole Verner, Abel Chapman and Walter Buck, and also Guy Mountfort for his Portrait of a Wilderness. iv ANGLIA RUSKIN UNIVERSITY Abstract FACULTY OF SCIENCE AND ENGINEERING DOCTOR OF PHILOSOPHY BIRDS IN THE NEANDERTHAL ECOLOGICAL NICHE – USING BIRDS FROM PLEISTOCENE CAVE ARCHIVES TO CHARACTERISE THE RELATIONSHIP BETWEEN NEANDERTHALS AND BIRDS STEWART FINLAYSON October 2019 Aspects of Neanderthal and modern human behaviour and ecology, particularly their relationship with birds are investigated. Birds are good indicators of climate and habitat and are frequently present in Neanderthal sites. This permitted the characterisation of Neanderthal habitats and climate, important given current questions about their ability to tolerate cold climates and tundra, which has implications when explaining their extinction as climate-caused. In addition, the inability of Neanderthals to catch birds has been used to support models of superior modern human cognitive capacity. Given recent published results showing Neanderthal exploitation of birds, a reassessment was considered necessary. A database of hominin sites and associated bird species was created. Bird species were assigned to specific climatic and habitat categories which permitted analyses of the climates and habitats occupied by Neanderthals and modern humans during the last glacial cycle (125 – 10 thousand years ago), the period when the two hominin taxa came into contact in Eurasia. The principal bird species associated with Neanderthals were identified and their taxonomic, behavioural and ecological features recognised. Finally, a database of bird species known to have been processed by Neanderthals was created from published sources. Both multivariate and univariate statistics were used to analyse the data including novel applications of PCA in this area. The results showed that Neanderthals were temperate climate hominins that rarely occupied extremely cold climates and open habitats. They occupied habitat mosaics where they regularly exploited a non-random suite of bird species, characterised by particular taxonomic, behavioural and ecological characteristics. The evidence presented here shows that, in terms of climate tolerance, habitat occupation and bird species exploitation, Neanderthals overlapped broadly with modern humans. Neanderthals had skills and abilities capable of despatching a wide range of birds and they were selective in the species that they took. This evidence calls into question models of modern human superiority as the cause of the Neanderthal extinction, and appears to support climate factors instead. Keywords: Neanderthal, Mousterian, Birds, Palaearctic, Pleistocene, Taphonomy v Table of Contents Acknowledgements ..................................................................................................... iii Abstract .............................................................................................................. v Table of Contents ........................................................................................................ vi List of Figures ............................................................................................................ xii List of Tables ......................................................................................................... xviii List of Appendices ..................................................................................................... xx Copyright ........................................................................................................... xxi Chapter 1 Introduction ............................................................................................ 1 1.1 Introduction ................................................................................................... 1 1.2 Neanderthals and Models of Recent Human Evolution ................................ 3 1.2.1 Behavioural Modernity and the Replacement Model ............................. 5 1.3 Neanderthals and behavioural modernity ...................................................... 6 1.3.1 Gibraltar, the Neanderthals and behavioural modernity ......................... 7 1.4 Birds in the Neanderthal ecological niche ................................................... 11 1.4.1 Birds as indicators ................................................................................. 14 1.4.2 Climate .................................................................................................. 16 1.4.3 Habitat ................................................................................................... 17 1.4.4 Birds as Resources ................................................................................ 18 1.5 Neanderthal as bird hunters ......................................................................... 19 1.6 Direct Evidence of Neanderthal Exploitation of Birds and its significance ................................................................................................ 22 1.7 Supporting sources of information .............................................................. 23 1.8 Research Questions and Thesis Structure .................................................... 25 1.8.1 Research Questions ............................................................................... 25 1.8.2 Thesis Structure .................................................................................... 25 vi Chapter 2 General Methods .................................................................................. 29 2.1 Introduction to the main dataset .................................................................. 29 2.2 The Study Sites ...........................................................................................