SummaryPolyploidy is viewed as an important mechanism of sympatric speciation, but few studies have documented the reproductive barriers between polyploids and their diploid progenitors or explored the significance of assortative mating for polyploid establishment. Here we synthesize new and existing data on five prezygotic (geographic isolation, flowering asynchrony, pollinator fidelity, self-pollination, gametic selection) and two postzygotic (selection against triploid hybrids, inbreeding depression) reproductive barriers between diploid and autotetraploid individuals of the perennial plant Chamerion angustifolium . We also present estimates of realized rates of between-ploidy mating and examine the impact of assortative mating on polyploid dynamics using computer simulation. Reproductive isolation (measured from 0 to 1) was enforced by each barrier, including: geographic separation (RI = 0.41), flowering asynchrony (0.13), pollinator fidelity (0.85), self-pollination (0.44), gametic selection (0.44) and postzygotic isolation (0.87). Total reproductive isolation was 0.997, with the largest relative contributions by geography (41%) and pollinator fidelity (44%). Prezygotic barriers accounted for 97.6% isolation overall; however, tetraploids were more assortatively mating (98%) than diploids (79%). Realized reproductive isolation between ploidy levels in sympatric populations was 87% and tetraploids produced significantly fewer triploids than did diploids. Simulations indicated that the observed prezygotic isolation will reduce the strength of minority disadvantage acting on tetraploids and increase the importance of differences in viability and fertility between cytotypes in regulating polyploidy establishment.© New Phytologist (2003) 161 : 703 -713

Establishment of polyploid individuals within diploid populations is theoretically unlikely unless polyploids are reproductively isolated, pre-zygotically, through assortative pollination. Here, we quantify the contribution of pollinator diversity and foraging behaviour to assortative pollen deposition in three mixed-ploidy populations of Chamerion angustifolium (Onagraceae). Diploids and tetraploids were not differentiated with respect to composition of insect visitors. However, foraging patterns of the three most common insect visitors (all bees) reinforced assortative pollination. Bees visited tetraploids disproportionately often and exhibited higher constancy on tetraploids in all three populations. In total, 73% of all bee flights were between flowers of the same ploidy (2x-2x, 4x-4x); 58% of all flights to diploids and 83% to tetraploids originated from diploid and tetraploid plants, respectively. Patterns of pollen deposition on stigmas mirrored pollinator foraging behaviour; 73% of all pollen on stigmas (70 and 75% of pollen on diploid and tetraploid stigmas, respectively) came from within-ploidy pollinations. These results indicate that pollinators contribute to high rates of pre-zygotic reproductive isolation. If patterns of fertilization track pollen deposition, pollinator-plant interactions may help explain the persistence and spread of tetraploids in mixed-ploidy populations.

Genome duplication causes phenotypic shifts and contributes directly to prezygotic isolation for some barriers (gametic isolation) but cannot account for the cumulative isolation from diploids observed in natural tetraploids. Therefore, conditions for species formation through genome duplication are not necessarily instantaneous and selection to strengthen prezygotic barriers in young polyploids is critical for the establishment of polyploid species in sympatry.

Our results indicate that tetraploids regularly coexist with diploids in the contact zone and that this coexistence is likely promoted by both strong reproductive isolation and asymmetric intercytotype mating between diploid and tetraploid C. angustifolium.