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Species Richness and Diversity in the Atlantic Rainforest and Associated Ecosystems: Niche Assembly Theories and Neutral Forces

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Species Richness and Diversity in the Atlantic Rainforest and Associated Ecosystems: Niche Assembly Theories and Neutral Forces MARKUS GASTAUER SPECIES RICHNESS AND DIVERSITY IN THE ATLANTIC RAINFOREST AND ASSOCIATED ECOSYSTEMS: NICHE ASSEMBLY THEORIES AND NEUTRAL FORCES Tese apresentada à Universidade Federal de Viçosa, como parte das exigências do Programa de Pós-Graduação em Botânica, para obtenção do título de Doctor Scientiae . VIÇOSA MINAS GERAIS – BRASIL 2012 Delight itself, however, is a weak term to express the feelings of a naturalis who, for the first time, has wandered by himself in a Brazilian forest. Charles Darwin, Feb. 29th, 1832 À Patrícia e Emmily, dedico... ii AGRADECIMENTOS À minha esposa Patrícia por ser um ótimo cônjuge, mas também pela convivência de todas as decepções e dificuldades, bem como as conquistas e descobertas enfrentadas durante esse doutoramento, pela correção da minha produção textual e pelo preparo de um coquetel delicioso e inesquecível para comemorar a conclusão do meu curso. À minha filha Emmily, que indica o que realmente importa na vida quando fala “Papai, brincar!”. Aos meus pais, por tudo que me ofereceram na vida e pela formação excelente que me habilitou concluir esse curso num pais que não é o meu. Aos orientadores, professores, colegas dos diversos programas que cursei, estagiários, ajudantes de campo e os amigos que contribuíram para esse tese e para minha formação. iii Sumário Resumo vi Abstract viii General Introduction 1 Chapter 1: Floristic Composition, Species Richness and Diversity of campo rupestre vegetation from Itacolomi State Park, Minas Gerais, Brazil 8 Abstract 10 Resumo 11 1. Introduction 13 2. Material and Methods 14 3. Results 17 4. Discussion 28 5. Conclusions 30 6. References 31 Chapter 2: How many species are there in the Itacolomi State Park? 40 Abstract 42 Introduction 43 Methods 44 Results 45 Discussion 47 Conclusions 51 Notes on Contributers 52 References 53 Chapter 3: Primary forests diagnosis in Brazilian Atlantic Rainforest 61 Abstract 63 Introduction 64 Material and Methods 66 Results 67 Discussion 74 Acknowledgements 77 Resumo 78 References 79 Supplement Material 91 iv Chapter 4: Community Dynamics in a Species-Rich Old- growth Forest Patch from Viçosa, Minas Gerais, Southeastern Brazil 99 Resumo 101 Abstract 102 Introduction 103 Materials and Methods 104 Results 106 Discussion 118 Acknowledgments 121 Bibliografic references 121 Chapter 5: Hubbell´s Theory at Age Ten: (Counter-) Evidence for Neutral Species Distribution in a Primary Forest Patch from Southeastern Brazil? 129 Abstract 131 Introduction 132 Methods ‚134 Results 138 Discussion 149 BOX 1. What do the Fitting Parameters of Hubbell´s Algorithm mean? 154 Acknowledgements 155 References 155 Supplement Material 163 Main Conclusion 166 v RESUMO GASTAUER, Markus, D.Sc., Universidade Federal de Viçosa, abril de 2012. Riqueza das espécies e diversidade na Mata Atlântica e ecossistemas associados: Teorias de nichos e forças neutras. Orientador: João Augusto Alves Meira Neto. Coorientadora: Andreza Viana Neri. Na primeira parte desta tese, dois levantamentos fitossociológicos realizados em vegetação de campo rupestre são apresentados e comparados. A primeira área de estudo fica bem protegida dentro do Parque Estadual do Itacolomi. Asegunda área é afetada por agropecuária, fogo, e demais atividades de urbanização. Como a área impactada mostra maior riqueza e diversidade, outros indicadores para a avaliação dessa vegetação são apresentados como o número de espécies invasoras, endêmicas ou ameaçadas de extinção. A partir desses dois estudos fitossociológicos, o número de espécies de plantas de todo campo rupestre do Parque Estadual do Itacolomi é concordante estimado para 1100 por meio de três abordagens diferentes. As três abordagens aplicadas são (i) extrapolação da curva espécie-área com a inclinação de 0,25, que é geralmente bem aceito para plantas terrestres, (ii) o modelo de Krishnamani e (iii) o modelo de relação dos táxons. Na segunda parte da tese, uma comunidade arbórea de um hectare da Floresta de Seu Nico (FSN) é apresentado. Segundo seus proprietários, este fragmento florestal nunca foi cortado e sofreu somente extração seletiva de madeira. Portanto, a hipótese é testada que a FSN ainda apresenta características de floresta primária, como são percentuais elevados de espécies não-pioneiras, espécies dispersas por animais, espécies de sub-bosque e espécies endêmicas. A FSN cumpre todas essas premissas, especialmente a sua riqueza de espécies é notável para a região. No entanto, a dinâmica desta comunidade arbórea mostra uma perda de espécies alarmante e aumento do número de indivíduos bem como a biomassa. Perda de espécies poderia ser uma resposta retardada ao desmatamento do redor e o alto grau de fragmentação da paisagem que reduz a imigração de propágulos de espécies novas para a FSN. O aumento vi da biomassa e do número de indivíduos indica aumento de disponibilidade de recursos, talvez devido às mudanças climáticas. Na terceira parte dessa tese, os mecanismos neutros e não-neutros são testados para explicar a distribuição espacial das espécies. Na FSN, as variáveis de solo e disponibilidade de luz no sub-bosque explicam pouco da distribuição espacial das espécies. A distribuição de abundância das espécies é ajustada melhor pelos modelos derivados de pressupostos neutros. A análise filogenética da comunidadena FSN, no entanto, indica sobredistribuição filogenética significativa devido à força alta de nichos, competição interespecífica e fatores de dependenciade densidade negativa. Como variáveis edáficas mostram um gradientepequeno e explicam apenas pouca variação na distribuição espacial das espécies, fatores bióticos como formuladas na hipótese de Janzen-Connell devem responder por essa distribuição formando um mosaico de diferentes nichos ecológicos. vii ABSTRACT GASTAUER, Markus, D.Sc., Universidade Federal de Viçosa, April, 2012. Species richness and diversity in the Atlantic Rainforest and associated ecosystems: Niche assembly theories and neutral forces. Adviser: João Augusto Alves Meira Neto. Co-adviser: Andreza Viana Neri. In the first part of this thesis, two phytosociological surveys carried out in campo rupestre vegetation are presented and compared. One study site lies well-protected within the Itacolomi State Park. The other one is impacted by extensive pasture, fire, and settling activities. As the impacted area shows higher species richness and diversity, further indicators for the intactness of this vegetation are presented like number of invasive, endemic or endangered species. From both surveys, the number of plant species from the Itacolomi State Park´s campos rupestre is concordantly estimated to 1100 using three different approaches, the species-area realtionship with the generally accepted slope of 0.25 for terrestrial plants, the Krishnamani model and the taxon ratio model. In the second part, a one hectare tree community from the Forest of Seu Nico (FSN) is presented. According to its owners, this forest fragment never has been logged. Therefore, the hypothesis was tested that the FSN still shows characteristics as a primary forest, as they are elevated percentages of non-pioneer, animal-dispersed, understory and endemic species. The FSN fulfills all requirements, especially its species richness is outstanding for the region. Nevertheless, the dynamics of this tree community show an alarming loss of species and increase of number of individuals as well as biomass. Species loss might be a time-delayed answer to comprehensive logging and high degree of fragmentation and isolation in the landscape reducing immigration of propagules from new species to the FSN. Increase of biomass and number of individuals is a hint for increased resource availability, perhaps due to climate change. In the third part, neutral and non-neutral mechanisms are tested to explain spatial species distributions. In the FSN, soil variables and understory light availability explain little spatial species distribution, species abundance distribution is viii furthermore best fit by models derived from neutral assumptions. A phylogenetic analysis of the community nevertheless indicates significant phylogenetic clustering due to niche assembly, interspecific competition and negative density-dependence factors. As edaphic variables show a short gradient only and explain little variance, biotic factors as formulated in the Connell-Janzen hypothesis should account for spatial species distribution forming a small-scaled mosaic of different ecological niches. ix General Introduction 1 2 Nevertheless species richness and diversity from all over the world are threatened and influenced by climate change (IPCC 2007), habitat loss and landscape´s fragmentation (Wright 2010), the discussion about what maintains species richness and diversity is still not at end. This lack of knowledge comprises especially the underlying – ecological – mechanisms like neutrality (Hubbell 2001), environmental sorting (Carvender-Bares et al . 2004) or niche assembly due to environmental heterogeneity (Hutchinson 1957) and negative density-dependant factors (Janzen 1971, Connell 1971) that may form and maintain species rich and diverse ecosystems. Before the Europeans arrived in South America, the Atlantic Rainforest, once classified as a hotspot of biodiversity (Myers et al. 2001) due to high species richness and a high degree of endemism (Stehmann et al. 2010), covered between 1.300.000 and 1.500.000 km 2 (Câmara 2005). Much of that has been logged and until today only 11 to 16 % of the original vegetation cover remainedin isolated fragments
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