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Pleistocene-Holocene Extinctions: Distinguishing Between Anthropic and Climatic Causes

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Pleistocene-Holocene Extinctions: Distinguishing Between Anthropic and Climatic Causes Universidade Federal do Rio de Janeiro Instituto de Biologia Programa de Pós-Graduação em Ecologia Pleistocene-Holocene extinctions: distinguishing between anthropic and climatic causes Bernardo Barros de Alvarenga Araujo Supervisor: Fernando A. dos Santos Fernandez Co-supervisor: José Alexandre F. Diniz-Filho Rio de Janeiro, RJ, Brazil - 2013 "The beauty and genius of a work of art may be reconceived, though its first material expression be destroyed; a vanished harmony may yet again inspire the composer; but when the last individual of a race of living beings breathes no more, another heaven and another earth must pass before such a one can be again." William Beebe, The Bird (1906). ACKNOWLEDGEMENTS Although this is one of the most enjoyable sections to write in a dissertation – mainly because it is a space to acknowledge the importance of many people I hold dear and because it is (hopefully) a red marker free zone – I will try to make it as quick as possible. First, I would like to thank my family for the infinite care offered throughout… well, through my whole life, but particularly in the years I’ve been in college. For eight years now I’ve had two homes filled with people who gave me nothing but love and incentive, and I couldn’t possibly begin to explain how much their support has meant (and will continue to mean) to me. In the respect of support, friends are also worth mentioning, especially the ones at the Laboratório de Ecologia e Conservação de Populações (Laboratory of Population Ecology and Conservation; LECP) of Universidade Federal do Rio de Janeiro (UFRJ), not only for the countless discussions and suggestions that were so valuable for the development of the present dissertation, but also for composing a working environment that is at the same time focused, greatly productive and so incredibly pleasant to inhabit. I would also like to extend a good extent of my gratitude to my co-supervisor José Alexandre F. Diniz-Filho, and to my dear colleagues Matheus Souza Lima-Ribeiro and Joaquín Hortal, for all the brainstorms and exchanges of ideas and data that fuelled my research. A particular acknowledgment on that respect should also go to Luiz Gustavo Rodrigues Oliveira-Santos for the enormous help provided with the conception and development of the models used in this work (and I still haven’t forgotten about that beer I owe you for all that). III A couple of agencies also deserve acknowledgement, namely the Conselho Nacional de Pesquisa e Desenvolvimento (CNPq) for providing the scholarship that aided me through the last two years, and the Programa de Pós-Graduação em Ecologia (PPGE) of UFRJ (along with its faculty members) for the chance to execute this somewhat audacious project. For last, and most certainly not least, I would like to thank Fernando Fernandez – supervisor, friend, and the very first person to open my eyes to the plight of so many fantastical creatures that became my personal object of fascination for the best part of four years. For me, this piece of (hard) work is as much yours as it is mine, and I wouldn’t have it any other way. Wherever my writings (academic or otherwise) should take me, the time spent under your tutelage will never be forgotten. And if I ever manage to build that time machine, I’ll be sure to make it a two-seater. IV ABSTRACT Through the last 50,000 years, the world lost about two thirds of its genera of large terrestrial animals, in a sequence that affected every continent except Africa. Explanations for these Quaternary extinctions, mostly based on regional studies, have been placed anywhere in a gradient ranging from climatic to anthropogenic causes. However, there have been few global quantitative analyses of this phenomenon, none of which takes full advantage of the vastly improved number and reliability of paleontological dates in recent years. Herein, we used a global approach to compare the explanatory power of the climatic and anthropogenic hypotheses. The world was divided in 19 regions where the extinctions took place at distinct times. Late Quaternary climatic variance expressed by 18O fluctuations and calibrated archeological dates of human first arrival to each region were used as predictive variables. The response variable was the last calibrated dates of occurrence of each megafaunal genus in each region. A comparison of the observed patterns with null models obtained by simulations showed that the extinctions were closer in time than expected by chance to climatic changes, human arrival, both or neither in 2, 82, 10 and 32 cases respectively (n = 126 last occurrence dates). Both cases where climate was the better explanation and most unexplained cases occurred in Eurasia, where the extinction process was longer and megafauna had a long interaction with early hominids. In a global perspective, however, the results provide strong support to the hypothesis that anthropogenic impacts are the best explanation for the extensive megafaunal extinctions that ravaged the planet in the Quaternary. Such realization should prompt us to rescue the true baselines of the world’s megafauna and ecosystem structure, and to reevaluate how we perceive and manage our remaining biodiversity and ecological processes. KEY-WORDS: Quaternary extinction, megafauna, anthropogenic impacts, climatic variance, global analysis. V RESUMO Ao longo dos últimos 50.000 anos, o mundo perdeu por volta de dois terços dos seus gêneros de grandes animais terrestres, em uma sequência que afetou todos os continentes exceto a África. As explicações para estas extinções do Quaternário, em sua maioria baseadas em estudos regionais, têm sido colocadas em qualquer ponto de um gradiente que vai de causas climáticas a antropogênicas. No entanto, têm havido poucas análises quantitativas globais deste fenômeno, nenhuma das quais tira completo proveito do vasto aperfeiçoamento no número e confiabilidade das datas paleontológicas nos anos recentes. Aqui, nós usamos uma abordagem global para comparar o poder explicador das hipóteses climática e antropogênica. O mundo foi dividido em 19 regiões onde as extinções aconteceram em tempos diferentes. A variância climática do final do Quaternário expressa por flutuações em 18O e datas arqueológicas calibradas de primeira chegada humana a cada região foram utilizadas como variáveis preditoras. A variável resposta foi a última data calibrada de ocorrência de cada gênero de megafauna em cada região. Uma comparação dos padrões observados com os modelos nulos obtidos por simulações mostrou que as extinções eram mais próximas no tempo do que o esperado ao acaso de mudanças climáticas, chegada humana, ambas ou nenhum dos dois em 2, 82, 10 e 32 casos respectivamente (n = 126 datas de última ocorrência). Ambos os casos nos quais o clima foi a melhor explicação e a maior parte dos casos não explicados ocorreram na Eurásia, onde o processo de extinção foi mais longo e a megafauna teve uma longa interação com hominídeos mais antigos. Numa perspectiva global, no entanto, os resultados fornecem um forte apoio para a hipótese de que impactos antropogênicos são a melhor explicação para as extensivas extinções da megafauna que assolaram o planeta no Quaternário. Essa percepção deve nos impelir a recuperar as verdadeiras linhas de base da megafauna e estrutura dos ecossistemas do mundo, e a reavaliar como nós percebemos e manejamos nossa biodiversidade e nossos processos ecológicos remanescentes. PALAVRAS-CHAVE: extinções do Quaternário, megafauna, impactos antropogênicos, variância climática, análise global. VI TABLE OF CONTENTS Acknowledgements III Abstract V Resumo VI List of Tables VIII List of Figures IX Introduction 1 Methods data 5 Methods analyses 10 Results 16 Discussion 25 References 32 Appendix I supplementary data a1 Appendix I supplementary data references a82 Appendix II supplementary figure a91 VII LIST OF TABLES Table 1 Mead-Meltzer Scale, as modified by Lindsey and Barnosky (2010) 7 Table 2 Scoring systems created to assess the reliability of U/Th and OSL dates 8 Table 3 Dates for extinct genera, human arrival, and summarized results from the null models 19 Table 4 Dates corrected by bootstrapping and comparisons between results 24 Table S1 Megafaunal dates from South America that ranked 11+ on the Mead-Meltzer Scale a2 Table S2 Megafaunal dates from Caribbean that ranked 11+ on the Mead-Meltzer Scale a5 Table S3 Megafaunal dates from North America that ranked 11+ on the Mead-Meltzer Scale a6 Table S4 Megafaunal dates from Eurasia that ranked 11+ on the Mead-Meltzer Scale a14 Table S5 Megafaunal dates from Japan that ranked 11+ on the Mead-Meltzer Scale a42 Table S6 Megafaunal dates from Madagascar that ranked 11+ on the Mead-Meltzer Scale a44 Table S7 Megafaunal dates from Tasmania that ranked 11+ on the Mead-Meltzer Scale a47 Table S8 Megafaunal dates from New Zealand that ranked 11+ on the Mead-Meltzer Scale a48 Table S9 Megafaunal dates from Australia that ranked 11+ on the Mead-Meltzer Scale a52 Table S10 Human dates from South America that ranked 13+ on the Mead-Meltzer Scale a60 Table S11 Human dates from North America that ranked 13+ on the Mead-Meltzer Scale a66 Table S12 Human dates from Eurasia that ranked 13+ on the Mead-Meltzer Scale a70 Table S13 Human dates from Japan that ranked 13+ on the Mead-Meltzer Scale a78 Table S14 Human dates from Tasmania that ranked 13+ on the Mead-Meltzer Scale a79 Table S15 Well ranked Rattus exulans dates from New Zealand as proxies for human arrival a80 VIII LIST OF FIGURES Figure 1 Regions considered on the analyses 11 Figure 2 Sampled dates of Mylodon from Cueva del Milodon, replacement for 1k iterations 15 Figure 3 Null model results - effects of climatic variance over extinct megafauna genera 17 Figure 4 Null model results - effects of human arrival over extinct megafauna genera 18 Figure 5 Null model results - Geographical distribution of all extinction causes 18 Figure S1 Effects of climatic variance and human arrival time-lapse over No.
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