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Araneae: Salticidae The Biogeography and Age of Salticid Spider Radiations with the Introduction of a New African Group (Araneae: Salticidae). by Melissa R. Bodner B.A. (Honours) Lewis and Clark College, 2004 A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE in The Faculty of Graduate Studies (Zoology) THE UNIVERSITY OF BRITISH COLUMBIA (Vancouver) July 2009 © Melissa R. Bodner 2009 ABSTRACT Globally dispersed, jumping spiders (Salticidae) are species-rich and morphologically diverse. I use both penalized likelihood (PL) and Bayesian methods to create the first dated phylogeny for Salticidae generated with a broad geographic sampling and including fauna from the Afrotropics. The most notable result of the phylogeny concerns the placement of many Central and West African forest species into a single clade, which I informally name the thiratoscirtines. I identify a large Afro-Eurasian clade that includes the Aelurilloida, Plexippoida, the Philaeus group, the Hasarieae/Heliophaninae clade and the Leptorchesteae (APPHHL clade). The APPHHL clade may also include the Euophryinae. The region specific nature of the thiratoscirtine clade supports past studies, which show major salticid groups are confined or mostly confined to Afro-Eurasia, Australasia or the New World. The regional isolation of major salticid clades is concordant with my dating analysis, which shows the family evolved in the Eocene, a time when these three regions were isolated from each other. I date the age of Salticidae to be between 55.2 Ma (PL) and 50.1 Ma (Bayesian). At this time the earth was warmer with expanded megathermal forests and diverse with insect herbivores. The two oldest region-specific clades are the South American Amycoida (41.1 Ma PL and 33.4 Ma Bayesian) and the Afro-Eurasian APPHHL clade (44.5 Ma PL and 33.8 Ma Bayesian), while the Australasian “core” astioids are younger (36.9 Ma PL and 27.3 Ma Bayesian). Mixing of fauna from isolated regions has been limited as large clades are geographically restricted, yet some more recent long-range dispersal events, such as the arrival of the genus Habronattus to the New World, have occurred. ii TABLE OF CONTENTS ABSTRACT ...................................................................................................................................ii TABLE OF CONTENTS............................................................................................................ iii LIST OF TABLES.........................................................................................................................v LIST OF FIGURES......................................................................................................................vi ACKNOWLEDGEMENTS........................................................................................................vii DEDICATION ........................................................................................................................... viii CO-AUTHORSHIP STATEMENT............................................................................................ix CHAPTER 1 An Introduction to Salticidae, Fossil Spiders and Molecular Dating Methods ..........................................................................................................................................1 1.1 Introduction ......................................................................................................................1 1.2 Literature Review.............................................................................................................2 1.2.1 The State of the Salticid Phylogeny.............................................................................2 1.2.2 Review of African Salticid Studies..............................................................................3 1.2.3 African Salticids from Arid Environments..................................................................4 1.2.4 Salticids from the Afrotropics......................................................................................5 1.2.5 A Review of Fossil Spiders in Amber .........................................................................6 1.2.6 Fossils and Molecular Dating Analyses in Spiders......................................................6 1.3 R8s and BEAST Dating Methods ...................................................................................7 1.3.1 Penalized Likelihood Implemented in R8s..................................................................7 1.3.2 BEAST: Bayesian Evolutionary Analysis Sampling Trees........................................8 1.4 Objectives and Hypotheses..............................................................................................9 1.4.1 Phylogeny, Age and Global Distribution of Salticidae................................................9 1.4.2 The Age of Salticidae...................................................................................................9 1.5 References .......................................................................................................................11 CHAPTER 2 The Biogeography and Age of Salticid Spider Radiations with the Introduction of a New African Group (Araneae: Salticidae) .................................................18 2.1 Introduction ....................................................................................................................18 2.2 Methods...........................................................................................................................20 2.2.1 Taxon Sampling.........................................................................................................20 2.2.2 DNA Extraction .........................................................................................................21 2.2.3 PCR Amplification and Sequencing ..........................................................................21 2.2.4 Sequence Alignment ..................................................................................................23 2.2.5 Phylogenetic Reconstruction .....................................................................................26 2.2.6 Divergence Time Estimation .....................................................................................27 2.3 Results .............................................................................................................................36 2.3.1 Model choice..............................................................................................................36 2.3.2 Phylogeny from the All-Genes Analysis ...................................................................36 2.3.3 Phylogenies from Individual Gene Regions ..............................................................37 2.3.4 Estimating Divergence Times....................................................................................39 2.4 Discussion........................................................................................................................42 2.4.1 The Thiratoscirtines: an African Radiation................................................................42 2.4.2 Gabonese Salticid Fauna............................................................................................43 2.4.3 Other Aspects of the Salticid Phylogeny ...................................................................44 2.4.4 Calibration Points.......................................................................................................46 2.4.5 Age and Diversity of Salticidae in Comparison to Other Radiations ........................49 iii 2.4.6 The Family Evolved at a Time of Expanded Megathermal Forests ..........................50 2.4.7 Regional Isolation of Major Salticid Groups.............................................................51 2.4.8 Biogeographic History of Region Specific Clades ....................................................54 2.4.9 The Thiratoscirtines are an Afrotropical Forest Group..............................................59 2.4.10 Age Alone Does Not Explain the Size of Salticid Radiations.................................60 2.5 References .......................................................................................................................87 Chapter 3 Salticidae: A Framework for Evolutionary Studies..............................................95 3.1 The Salticid Radiation ...................................................................................................95 3.2 Reconstructing and Dating the Salticid Phylogeny.....................................................95 3.2.1 Strengths of the Thesis...............................................................................................95 3.2.2 Challenges of Reconstructing the Salticid Phylogeny...............................................96 3.2.3 Dating and Gaps in the Jumping Spider Fossil Record .............................................96 3.3 Exploring Community Level Convergences Using the Salticids................................97 3.4 Working Hypotheses......................................................................................................98 3.4.1 Number of Dispersals Between Isolated Regions......................................................98 3.4.2 Ecomorphology..........................................................................................................99 3.5 Continuing to Build the Salticid Tree of Life ..............................................................99
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