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Wattles and Wombats:
Molecular rate variation and Biodiversity
Xavier Goldie
The University of AucklandThe Australian National University
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Outline:• Biodiversity and Evolution
• Evolutionary Speed in Australian Plants
• Evolutionary Speed Hypothesis
• Productivity and the Water-Energy Balance
• Outcomes
• Molecular rates in Mammals
• Diversification and molecular rates: crash course
• Outcomes
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Biodiversity and Evolution
• What drives diversification?
SPECIATION - EXTINCTION
• Why are species unevenly distributed -
• Spatially?
• Phylogenetically?
• Temporally?
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Rodents
Lagomorphs
Primates +
Cetartiodactyla +
Carnivora +
Chiroptera
Shrews and stuff
Xenarthra
Afrotheria
Metatheria
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Evolutionary Speed in Australian Plants
Climate and Biodiversity
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Rohde’s Evolutionary Speed Hypothesis (ESH)
Bromham and Cardillo 2003
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Rohde’s ESH deals specifically with the relationship between global temperature and biodiversity
However…
Terrestrial biodiversity is linked most tightly to the interaction of temperature and the physical state of water
“Water-Energy Balance”(Hawkins et al 2003)
“Productivity”(Gillman and Wright, 2006)
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Is Rohde’s ESH applicable to gradients of water availability?
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Species RichnessRainfallProductivity
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Amyema congener
Amyema maidenii
“Wet”
“Dry”
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30 Phylogenetically Independent contrasts
Branch length Trait
P.A.
Branch length Trait
Branch length Trait
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Significantly more (23 of 30) comparisons resulted in longer “Wet” branch lengths
(Sign Test: P = 0.005)
The rate of nucleotide substitution in “Wet” zone taxa was on average more than twice that of
closely related “Dry” zone congeners
(Wilcoxon Signed Rank Test:
W = 93, P = 0.003).
Goldie et al., 2010
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Results are consistent with the evolutionary speed hypothesis
Optimised water-energy balance may act to increase micro-evolutionary rates
in mesic areas
Arid conditions may lead to slower tempo of micro-evolution
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Bromham and Cardillo 2003
Rohde’s Evolutionary Speed Hypothesis
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Molecular Rates
Speciation Rates
Wright et al 2006Lanfear et al In Prep
Webster et al 2003Pagel et al 2006
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Molecular Rates in Mammals
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What is the relationship between the rate of molecular evolution,
and diversification,in mammals?
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Whole Mitochondrial Genomes and Nuclear Genes
Estimated Synonymous, Non-Synonymous and total substitution rates
(MG94_REV_3x4 in HyPHY)
Mitochondrial Sister-clades at roughly ordinal, familial and generic level
Nuclear Sister-Clades in Mammalia, Eutheria, Metatheria, Euarchontoglires and Laurasiatheria
Sign-Tests and Parametric Regression
Included Body Size in Multiple Regression
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Ne and/or Selection
Mutation
Clade A Clade B Clade A Clade B
dN
dS - -
dN/dS - -
Brlen
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No consistent relationship between molecular rates and clade size - mitochondrial or nuclear
Body Size Effect detected in nuclear genes, but not in mitochondrial
Saturated Synonymous Rates in mitochondrial genes
“RY” Coded four-fold degenerate sites - record only synonymous Transversions (T or C ↔ G or A)
No relationship
(Goldie et al, In Prep)
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Eutheria
ML Brlen versus Clade Size
P = 0.00545
But not repeated in dN or dS
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Welch et al 2007
P = 0.1260
Mitochondrial dS
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BUT WHAT DOES IT ALL MEAN?
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Lack of apparent relationship between mammalian molecular rates and diversification rates?
Molecular rates are not central to the diversification process in mammals
Different process operating between plants and mammals - has been suggested
But why birds, and not mammals?
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Gillman et al 2009
Higher rates of molecular evolution in tropical mammals versus temperate PICs
Interpreted as support for Evolutionary Speed Hypothesis driving Latitudinal Species Gradients
Bassariscus astustus Bassariscus sumichrasti
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Rohde’s ESH
Bromham and Cardillo, 2003
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Acknowledgements:
Shane Wright
Len Gillman
Lindell Bromham
Rob Lanfear
Nga Pae O Te Maramatanga Maori Centre of Research Excellence
The Australian National University
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Questions?