WALLACE BEIROZ IMBROSIO DA SILVA UNDERSTANDING SPATIAL AND TEMPORAL SHIFTS IN TAXONOMIC AND FUNCTIONAL DIVERSITY: ASSESSING DUNG BEETLES IN A HUMAN-MODIFIED AMAZONIAN LANDSCAPE Dual PhD Scheme LAVRAS - MG 2016 i WALLACE BEIROZ IMBROSIO DA SILVA UNDERSTANDING SPATIAL AND TEMPORAL SHIFTS IN TAXONOMIC AND FUNCTIONAL DIVERSITY: ASSESSING DUNG BEETLES IN A HUMAN- MODIFIED AMAZONIAN LANDSCAPE Thesis submitted for the degree of Doctor of Philosophy as a Dual PhD with ‘Ecologia Aplicada’ Postgraduate Program, Universidade Federal de Lavras, Brazil and Lancaster Environment Centre, Lancaster University, United Kingdom PhD Júlio Louzada (Universidade Federal de Lavras) PhD Emma Sayer (Lancaster University) Supervisors PhD Jos Barlow (Lancaster University) PhD Eleanor Slade (University of Oxford and Lancaster University) Co-Supervisors LAVRAS - MG 2016 ii Ficha catalográfica elaborada pelo Sistema de Geração de Ficha Catalográfica da Biblioteca Universitária da UFLA, com dados informados pelo próprio autor da Silva, Wallace Beiroz Imbrosio. Understanding spatial and temporal shifts in taxonomic and functional diversity: Assessing dung beetles in a human-modified Amazonian landscape / Wallace Beiroz. – Lavras: UFLA, 2016. 132 p. : il. Tese (Doutorado)–Universidade Federal de Lavras, 2016. Orientador(a): Julio Louzada (UFLA) / Emma Sayer (Lancaster) Bibliografia. 1. Scarabaeinae. 2. Functional Diversity. 3. Tropical Forest. I. Universidade Federal de Lavras. II. Título. iii WALLACE BEIROZ IMBROSIO DA SILVA UNDERSTANDING SPATIAL AND TEMPORAL SHIFTS IN TAXONOMIC AND FUNCTIONAL DIVERSITY: ASSESSING DUNG BEETLES IN A HUMAN- MODIFIED AMAZONIAN LANDSCAPE Thesis submitted for the degree of Doctor of Philosophy as a Dual PhD with ‘Ecologia Aplicada’ Postgraduate Program, Universidade Federal de Lavras, Brazil and Lancaster Environment Centre, Lancaster University, United Kingdom Approved on 25th November, 2016 PhD Paulo dos Santos Pompeu Universidade Federal de Lavras, Brazil PhD Luiz Fernando Silva Magnago Universidade Federal de Lavras, Brazil PhD Rosa Menéndez Lancaster University, UK PhD André Tavares Corrêa Dias Universidade do Estado do Rio de Janeiro, Brazil PhD Júlio Louzada (Universidade Federal de Lavras) PhD Emma Sayer (Lancaster University) Supervisors PhD Jos Barlow (Lancaster University) PhD Eleanor Slade (University of Oxford and Lancaster University) Co-Supervisors Dual PhD Scheme LAVRAS - MG 2016 iv I dedicate this thesis to my parents, Hélio Francisco and Lucy Imbrosio, whom gave me all the love where I was born and grew up. Eu dedico esta tese aos meus pais, Hélio Francisco e Lucy Imbrosio, quem me deu todo o amor onde eu nasci e cresci. Dedication v ACKNOWLEDGEMENTS / AGRADECIMENTOS I am very grateful to my supervisors, Emma Sayer, Eleanor Slade, Jos Barlow and Julio Louzada, without their precious help I could not have done this thesis. Thank you for your patience, enthusiasm and for showing me that great researchers can also be great people! Thank for the lovely, crazy, silly, friendly, weird, chaotic and warm ‘Tropical Group’! In special, Natalie ‘Cisne’, Anne, Luke, Mari, Erika and Juliana. Thank you for making my life much easier and happier in UK! Thank you the people in the best office ever!!!! Guy, James, Mounir, Deidre, Radim, Katherine. I am very thankful for Tregidgo’s (‘Tregedigdigdigo’s’) for one of the best Christmas ever and for looking after me during my stay in UK, specially my ‘white bro’ Danny ‘Lézo’, for being such a terrific (or terrible) friend! I also in debt with some amazing friends I have met in Lancaster (or even before). Specially, Tom Walker, Sam Robinson, Maria ‘Malakas’, Bobby, Hannah ‘Maritaca’, Daniel Baxendale, Katherine Baxendale and Isobel, thank you for all the serious and silly chats we had, for being such a lovely friend and always (I mean ALWAYS) being there for me!! Thank you my lovely and graceful roomates. Thank you Anni (Dr. Úrsula) for being the most kind and cute person I ever met! Thank you Kasia (‘Katarzina Milakovich’) for taking care of me and being a great company in Lancaster! I am also grateful for the lovely people I met in Oxford. Specially two great hosts, Paul Manning and Olelokan. Thank you guys for your friendship and kindness. I also thank to my friends in Liverpool, Olívia, Tom Bishop and Sofia. You made my visits in Scouser’s land very lovely and funny! Quero agradecer também à outra família que me adotou, mas esta é de brasileiros, ao meus grandes amigos Beto, Michelle ‘CatÚnda’, Daniel e Marina. Agradeço também à outros amigos brasileiros que estavam ‘turistando’ na terra da Rainha, Lígia, Lélis, Gabriel, Arlete, Caio, Pedro, Andrézão e Fábio. Agradeço muito ao meu pai Hélio, minha mãe Lucy, meu irmão Helinho e minha cunhada Aline. Obrigado por sempre me darem muito amor, apoio e ajuda; dando conselhos ou apenas ouvindo. Nunca vou esquecer a frase “vai fundo cara, olha onde suas decisões já te levaram”. vi Agradeço também ao Ronaldo, Sílvia e Ramine, por terem entrado na minha vida com dois pés direitos e também terem sempre me dado muito apoio e carinho. Agradeço imensamente meu bem Ananza, por todos esses anos de companheirismo, diversão, amor e alegria. Muito obrigado por sempre “stand by me”, até nos meus maiores momentos de estresse. Te amo! Aos meus amigos de Lavras, obrigado pelos anos de amizade e apoio. Especialmente Rafael ‘Peixe’, Ruanny ‘Tatau’, Ivo, Yuri, Tonho, Ernesto, Lisi, Chaim, Carla, Danilo, Grazi, Poly, Nat, Guilherme, Gabi, Thalles. Tem duas pessoas de Lavras que quero agradecer separadamente por terem me ajudado bastante na tese. Muito obrigado Julius pela ajuda na identificação dos bichos e Marina por analisar as fotos do projeto, sem vocês eu não conseguiria. Claro, muito obrigado pela amizades de vocês, sem isso eu também não conseguiria. Agradeço também à todos os Rola-bosteiros que me ajudaram a descontrair no laboratório, discutir as ideias, ou com um simples abraço. Especialmente Cris ‘Bahia’, Toru, Lívia ‘Bagé’. Obrigado aos animais do extinto chiqueiro 1/CADA por todos os momentos de amizade e de descontração, muito obrigado. Calorada, Tonho, André, Vini. Obrigado Edvar, Irmão e Maria, pela grande (essencial) ajuda no meu campo no Jari! Obrigado ao Programa de Pós-Graduação em Ecologia Aplicada da UFLA pela oportunidade. Obrigado à CAPES pelas bolsas de doutorado no Brasil e sanduíche no exterior e ao CNPq por ter financiado o projeto que eu estive envolvido. Thank you Lancaster Univeristy for the great opportunity to study abroad. Obrigado à todos que de alguma forma ajudaram no processo desta tese, direta ou diretamente. Thank you the people that somehow was envolved on this Thesis and helped me to conclude it. vii ABSTRACT Tropical forests are under anthropogenic pressure, causing the replacement of native areas by modified habitats. These landscapes are composed of natural and modified components. Knowledge of community dynamics in these modified habitats is essential to ensure long-term conservation of species and ecosystem services. Many studies had evaluated the conservation value and impacts of anthropogenic modifications on biodiversity. However, the natural variation of communities in reference sites could obscure responses of biodiversity to disturbance. I evaluated the spatial and temporal shifts in dung beetle communities in undisturbed forest, and used these results to assess the influence of inter-annual variation on the evaluation of anthropogenic changes. Soil texture was the main factor affecting spatial variation in dung beetle communities, mainly due to different responses among beetles with distinct nesting behaviour. I found high variation in almost all community metrics after a severe dry season in the previous year, which also influenced the response of the community in modified habitats. Thus, future studies should consider inter-annual variation when studying the effect of anthropogenic change on dung beetle communities and use information about factors that could influence community metrics over time. In addition, many studies evaluate community responses to change from either taxonomic or functional perspectives, but little is known about the relationship between these components of biodiversity. I also addressed this by using conceptually similar taxonomic and functional metrics to evaluate the influence of anthropogenic changes on inter-annual variation in dung beetle communities and to compare taxonomic and functional metrics responses to modifications in forest structure. I did not find strong evidence for the influence of anthropogenic change on the inter-annual variation in the studied metrics. However, functional metrics varied less among years than taxonomic. Additionally, despite similar average effect sizes of functional and taxonomic metrics in response to forest modification, I found high variation among years for each metric and among metrics within each year. I therefore suggest that using taxonomic and functional metrics concomitantly could improve studies focusing on evaluating anthropogenic changes. Finally, I assessed the factors influencing taxonomic and functional diversity of dung beetles in plantations at the local and landscape scale. Understanding the factors modulating biodiversity in modified habitats has important applications, as these areas are likely to be the main component of tropical landscapes in the near future. I evidenced that dung beetle diversity increased with the extent of native forests surrounding the plantation, but there was no relationship between beetle