Introduction
Topic: How reproductive isolation evolves and why it matters for speciation.
Central question: Why would natural selection favor alleles that reduce mating with others (i.e., reproductive isolation)?
Natural selection and the puzzle of reproductive isolation
Natural selection increases alleles that improve individual reproductive success.
Reproductive isolation appears to reduce mating opportunities, so its evolution needs explanation.
Allopatric speciation (geographic isolation)
Mechanism: A single interbreeding population becomes geographically split (example: lake split into two by land/sea change).
Consequence: Gene flow stops, allowing independent evolution via natural selection and genetic drift.
Outcomes: Divergence in morphology, behavior, development, etc., can produce reproductive isolation when populations re-contact.
Implication: Reproductive isolation can evolve as a byproduct (not directly selected for) during isolation.
Evidence: Regions with high potential for geographic isolation (e.g., Galápagos) show rapid speciation (Darwin's finches).
Conclusion: Allopatric speciation is regarded as the dominant mode in animals and many plants.
Sympatric speciation (no geographic isolation)
Challenge: With ongoing gene flow, divergence is impeded; reproductive isolation is hard to evolve without blocking gene flow.
Polyploid speciation (plants): Instantaneous reproductive isolation can occur when genome duplication produces polyploid individuals whose gametes are incompatible with the parent diploids.
Can occur via self-fertilization or hybrid polyploid formation; common in plants, rare in animals (some amphibians).
Ecological/behavioral sympatric speciation (non-polyploid): Rare but documented example — Rhagoletis (fruit flies).
Scenario: Ancestral hawthorn-feeding flies shifted to apples; mating occurs on host plant, so assortative mating by host reduces gene flow.
Additional differences: Divergence in physiology, behavior, lifecycle timing; generally considered subspecies but reproductively isolated in nature.
Caveat: Hard to rule out short historical geographic isolation; proving strict sympatry is difficult.
Summary conclusions
Most species-level reproductive isolation likely evolved after periods of geographic isolation (allopatry).
Sympatric speciation is possible: commonly via polyploidy in plants; in animals it is rare and requires unusual ecological or behavioral conditions.
Demonstrating true sympatric speciation is challenging because historical geographic events are difficult to exclude.
Unresolved questions / caveats
How frequent is true sympatric speciation in animals beyond a few special cases?
For cases like Rhagoletis, absolute exclusion of past geographic isolation is hard, leaving some uncertainty.