Evolution Lecture 25 Chapter 18 Topics for today: Four modes of geographic speciation 1. Genetic factors that lead to reproductive isolation 2. Evidence from: • natural populations • experimentation Mechanisms of speciation Reproductive isolation is key Geographic factors reduce gene flow and allow reproductive isolation to evolve (4 ways) Genetic factors cause reproductive isolation 1. Genetic divergence due to: • Ecological selection • Sexual selection 2. Break up of positive epistasis (Dobzhansky-Muller) 3. Cytoplasmic incompatibility 4. Chromosome divergence (polyploidy, cytological change) 5. Recombination in hybrids Four geographical modes of speciation Fig. 18.1 new 16.1 old A. Allopatric speciation: physical barrier Most common mechanism of speciation? Defined by reduction in movement of individuals and gametes Distance not necessarily important Diverge due to drift and selection Reproductive isolation is a by-product of selection Renewed contact may result in o complete isolation o hybridization Evidence for allopatric speciation Incipient prezygotic and/or postzygotic mating barriers among geographic populations o Experimental pairing of individuals within and between populations o Do they mate? Molecular differentiation among allopatric populations o Mitochondria DNA of six freshwater fish species o Distinct eastern and western clades in all species Fig. 18.3 new 16.3 old Molecular differentiation among allopatric populations o Mitochondria DNA of six freshwater fish species o Distinct eastern and western clades in all species o Gene flow reduced by Appalachian moutains o Isolation occurred 3-4 myo Filg 18.3 new 16.3 old Increasing geographic overlap of species Allopatric species originally separated How should the geographic overlap of closely related species pairs change over time? Can only increase or stay at zero Genetic divergence used as an index for time since divergence Fig. 18.5 new 16.5 old Causes of reproductive isolation in allopatry? Genetic divergence via ecological selection o Reproductive isolation evolves as a by-product of selection on other traits (pleiotropic effects) o Example 1 o Test mating compatibility of herbivorous insects that specialize on different hosts Fig. 18.8 new 16.7 old o Example 2 Evidence is stronger if the pattern of isolation is repeated (parallel divergence) Three-spined stickleback Fig. 18.8 new 16.7 old Genetic divergence – sexual selection o Different male traits may evolve by female choice in different populations ultimately causing reproductive isolation o Two populations of bush crickets o Example 1 Ephippiger ephippiger Different male traits may evolve by female choice in different populations ultimately causing reproductive isolation Two populations of bush crickets Differences in♀ choice drive differences in ♂ song Fig. 18.12 new 16.10 old o Example 2 Sexual selection on males is strongest when there is high variance in reproductive success Fig. 18.12 new 16.9 old Dobzhansky-Muller incompatibility o Populations have same allele frequencies initially o Mutations occur that have positive epistatic effects with alleles at other loci & are fixed by selection o Hybridization breaks up positive epistasic relationships that differ among populations Fig. 18.7 new 16.6 old