Baker's Law predicts uniparental reproduction will facilitate colonization success in novel habitats. While evidence supports this prediction among colonizing plants and animals, few studies have investigated shifts in reproductive mode in haplo-diplontic species in which both prolonged haploid and diploid stages separate meiosis and fertilization in time and space. Due to this separation, asexual reproduction can yield the dominance of one of the ploidy stages in colonizing populations. We tested for shifts in ploidy and reproductive mode across native and introduced populations of the red seaweed Gracilaria vermiculophylla. Native populations in the northwest Pacific Ocean were nearly always attached by holdfasts to hard substrata and, as is characteristic of the genus, haploid-diploid ratios were slightly diploid-biased. In contrast, along North American and European coastlines, introduced populations nearly always floated atop soft-sediment mudflats and were overwhelmingly dominated by diploid thalli without holdfasts. Introduced populations exhibited population genetic signals consistent with extensive vegetative fragmentation, while native populations did not. Thus, the ecological shift from attached to unattached thalli, ostensibly necessitated by the invasion of soft-sediment habitats, correlated with shifts from sexual to asexual reproduction and slight to strong diploid bias. We extend Baker's Law by predicting other colonizing haplo-diplontic species will show similar increases in asexuality that correlate with the dominance of one ploidy stage. Labile mating systems likely facilitate colonization success and subsequent range expansion, but for haplo-diplontic species, the long-term eco-evolutionary impacts will depend on which ploidy stage is lost and the degree to which asexual reproduction is canalized.

The identification of native sources and vectors of introduced species informs their ecological and evolutionary history and may guide policies that seek to prevent future introductions. Population genetics provides a powerful set of tools to identify origins and vectors. However, these tools can mislead when the native range is poorly sampled or few molecular markers are used. Here, we traced the introduction of the Asian seaweed Gracilaria vermiculophylla (Rhodophyta) into estuaries in coastal western North America, the eastern United States, Europe, and northwestern Africa by genotyping more than 2,500 thalli from 37 native and 53 non‐native sites at mitochondrial cox1 and 10 nuclear microsatellite loci. Overall, greater than 90% of introduced thalli had a genetic signature similar to thalli sampled from the coastline of northeastern Japan, strongly indicating this region served as the principal source of the invasion. Notably, northeastern Japan exported the vast majority of the oyster Crassostrea gigas during the 20th century. The preponderance of evidence suggests G. vermiculophylla may have been inadvertently introduced with C. gigas shipments and that northeastern Japan is a common source region for estuarine invaders. Each invaded coastline reflected a complex mix of direct introductions from Japan and secondary introductions from other invaded coastlines. The spread of G. vermiculophylla along each coastline was likely facilitated by aquaculture, fishing, and boating activities. Our ability to document a source region was enabled by a robust sampling of locations and loci that previous studies lacked and strong phylogeographic structure along native coastlines.

Differences with respect to anti-herbivore defense were investigated in invasive and native populations of the seaweed Gracilaria vermiculophylla. Specimens from 6 native populations in East Asia and from 8 populations invasive in Europe and the Mexican Pacific coast were maintained under identical conditions and offered to herbivorous snails from both the native range (Littorina brevicula) and Europe (L. littorea) in no-choice feeding assays. L. brevicula consumed in total significantly larger amounts of G. vermiculophylla tissue than did L. littorea. Further, both snail species least consumed the seaweed specimens originating from either non-native populations or from populations native to the Korean East Sea/Sea of Japan. The Korean East Sea/Sea of Japan had previously been identified as putative donor region of all the invasive populations of G. vermiculophylla. Thus, populations in the donor region as well as non-native populations in different invaded realms feature an increased capacity to resist feeding pressure. Differences in nutrient content did not account for the observed patterns of consumption, as palatability and carbon to nitrogen (C:N) ratio were not significantly correlated. Thus, mechanical or chemical defenses or the content of feeding cues influenced the behavior of the snails. We suggest that low palatability contributed to the invasion success of the species.

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The perennial red macroalga Gracilaria vermiculophylla (Ohmi) Papenfuss has recently been introduced to the Baltic Sea and is a potential competitor to Fucus vesiculosus, the most common native perennial alga in large parts of the Baltic Sea. Gracilaria might interfere with Fucus through direct competition for resources. In addition, Gracilaria is a favoured refuge for mesograzers, which prefer to feed on Fucus. Mesocosm-experiments were conducted over one year in the Kiel Fjord in order to test the direct and indirect effects of Gracilaria on Fucus. Fucus was incubated with Gracilaria at three different densities and grazers in high or low abundances. High densities of Gracilaria inhibited the growth of Fucus adults and also reduced the half-life-time of Fucus germlings. Associated grazers also had a negative effect on Fucus adults. Our results suggest that Gracilaria is able to influence Fucus in the Baltic Sea through direct competition for resources and by exposing it to higher grazer pressure.

The source and vector of an introduced species inform its ecological and evolutionary history and may guide management that seeks to prevent future introductions. Surprisingly, few studies have successfully used genetic tools to independently inform the specific source and pathway of biological invasions. The ecological history of many introduced species, including their origins and vectors, is often based on suppositions or educated guesses. Here, we used mitochondrial and microsatellite genotyping to trace the invasion of the Asian seaweed Gracilaria vermiculophylla (Rhodophyta) along the three coastlines of the Northern Hemisphere to which it has been introduced: the western coast of North America, eastern coast of the United States and the coasts of Europe and northwest Africa. Analyzing 37 native and 53 introduced sites, we identified the Pacific coastline of northeastern Japan as the ultimate source of the Northern Hemisphere invasion. Coincidentally, most exports of the oyster Crassostrea gigas historically originated from this region and both species often grow in close proximity. Based on genetic signatures, each of the three coastlines likely received thalli directly from Japan, as well as material from another introduced coastline (i.e., a secondary invasion). Our ability to document a source region, which was enabled by a robust sampling of locations and loci that previous studies lacked, reflected strong phylogeographic structure along native coastlines. We suggest Gracilaria vermiculophylla is an important representative example of many species likely exported out of Japan by the oyster trade and its genetic signatures that may be a hallmark of oyster introduction legacies.

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