Reproductive isolation Reproductive isolating barriers Extrinsic and intrinsic isolation: how to tell apart [Blackboard: population genetics of assortative mating] SPECIATION / EVA KISDI / 2014 FALL / LECTURE 2 1 Reproductive isolating barriers n Prezygotic isolation: no hybrid zygote is formed q Behavioural isolation: song, pheromone, courtship, size, ... Darwin's finches, Podos 2001 Geospiza magnirostris Geospiza fuliginosa kHz 0.5 s q Mechanical isolation: insect genitalia, flower morphology SPECIATION / EVA KISDI / 2014 FALL / LECTURE 2 2 Reproductive isolating barriers n Prezygotic isolation q Gametic isolation: cross-fertilization impossible or conspecific sperm / pollen precedence Sea urchin spp: sympatric broadcast spawners Echinometra bindin: released to enter the vitelline envelope of the egg, rapidly diverges between species (high replacement / silent substitution ratio) SPECIATION / EVA KISDI / 2014 FALL / LECTURE 2 3 Reproductive isolating barriers n Prezygotic isolation q Ecological isolation (byproducts of adaptation) n n Habitat isolation: breed in different habitats within cruising range Allochronic isolation: breed at different times hawthorn race apple race apple maggot fly Rhagoletis pomonella n Aquilegia formosa (vs A. pubescens) Pollinator (floral) isolation: different insect spp or different individuals (!) SPECIATION / EVA KISDI / 2014 FALL / LECTURE 2 4 Reproductive isolating barriers n Postzygotic isolation: hybrid inviability or sterility q Extrinsic postzygotic isolation: environment-dependent n n q Ecological: hybrids are selected against in nature (no niche) Behavioural: hybrids cannot find appropriate mates Intrinsic postzygotic isolation: in any / in stressful environment n n Hybrid inviability: death during development or reduced survival later Hybrid sterility: no functional reproductive organs or incapable of courtship/parental care Agapornis roseicollis x A. personata fischeri female hybrids fail to build a nest / do not incubate SPECIATION / EVA KISDI / 2014 FALL / LECTURE 2 5 Extrinsic postzygotic isolation H. cydno hybrid proportion attacked by birds Heliconius melpomene ME F1 CY Merrill et al. (2012) SPECIATION / EVA KISDI / 2014 FALL / LECTURE 2 Extrinsic vs intrinsic isolation How to tell apart extrinsic and intrinsic postzygotic isolation? Threespine stickleback (Gasterosteus aculeatus) invaded freshwater lakes in Canada after the ice age Benthic and limnetic morphs differ in morphology (large/small; jaws, gill rakers) diet (benthic invertebrates/zooplankton) Trade-off between habitats: Schluter (1995) SPECIATION / EVA KISDI / 2014 FALL / LECTURE 2 7 Extrinsic vs intrinsic isolation Hypothesis: hybrids are not efficient in either habitat = extrinsic postzygotic isolation 1. F1 hybrids are viable and fertile in the laboratory Hatfield & Schluter 1999 SPECIATION / EVA KISDI / 2014 FALL / LECTURE 2 8 Extrinsic vs intrinsic isolation 2. F1 hybrids do poorly in both natural habitats benthic limnetic F1 hybrid Hatfield & Schluter 1999 - but is this because they are not adapted to either niche (extrinsic) or because they have intrinsic genetic incompatibilities that manifest only in stressful environments? SPECIATION / EVA KISDI / 2014 FALL / LECTURE 2 9 Extrinsic vs intrinsic isolation 3. Backcrosses: F1 x benthic ® BB, F1 x limnetic ® BL phenotypically closer to the parental forms: if extrinsic, BB should do better in the littoral habitat BL should do better in open water equal genetic incompatibilities: if intrinsic, BB and BL should do equally badly in both habitats Rundle 2002 SPECIATION / EVA KISDI / 2014 FALL / LECTURE 2 10