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Determining the relative importance of sexual and asexual reproduction is critical for determining how genetic diversity is partitioned within and among populations. Macroalgae exhibit reproductive system variability, with many species capable of both sexual and asexual reproduction. Thus, rates of fragmentation (asexual) versus sporic recruitment (sexual) must be quantified. This is critical during range expansions when shifts to uniparental reproduction are common. Chondria tumulosa is a cryptogenic red alga that was discovered forming thick mats at Manawai, in the Northwestern Hawaiian Islands, during opportunistic sampling in 2019. It reduced the biodiversity and abundance of associated coral reef organisms. To characterize the reproductive system of C. tumulosa in this atoll, we visually assessed 124 thallus clumps for reproductive structures and used five microsatellite loci to genotype 41 thalli. All thalli bearing reproductive structures (20%) and all thalli that were genotyped were tetrasporophytes. No gametophytic thalli were observed using microscopy or microsatellite genotyping, though 80% of thalli observed under the microscope were vegetative. We observed in situ thallus fragmentation supported by the presence of 17 repeated genotypes and other genetic signatures, suggesting high rates of asexual reproduction. These results suggest that C. tumulosa is partially clonal in this atoll and has the potential for spreading to other nearby atolls. Rapid establishment, thallus fragmentation and tetrasporophytic dominance are consistent with other red macroalgal invasions studied to date. Understanding the reproductive system is an important component of developing management strategies for this alga and expanding our understanding of the population genetics of partially clonal taxa. [ABSTRACT FROM AUTHOR]

Partial clonality is known to affect the genetic composition and evolutionary trajectory of diplontic (single, free-living diploid stage) populations. However, many partially clonal eukaryotes exhibit life cycles in which somatic development occurs in both haploid and diploid individuals (haplodiplontic life cycles). Here, we studied how haplodiplontic life cycles and partial clonality structurally constrain, as immutable parameters, the reshuffling of genetic diversity and its dynamics in populations over generations. We assessed the distribution of common population genetic indices at different proportions of haploids, rates of clonality, mutation rates, and sampling efforts. Our results showed that haplodiplontic life cycles alone in finite populations affect effective population sizes and the ranges of distributions of population genetic indices. With nonoverlapping generations, haplodiplonty allowed the evolution of 2 temporal genetic pools that may diverge in sympatry due to genetic drift under full sexuality and clonality. Partial clonality in these life cycles acted as a homogenizing force between those 2 pools. Moreover, the combined effects of proportion of haploids, rate of clonality, and the relative strength of mutation versus genetic drift impacts the distributions of population genetics indices, rendering it difficult to transpose and use knowledge accumulated from diplontic or haplontic species. Finally, we conclude by providing recommendations for sampling and analyzing the population genetics of partially clonal haplodiplontic taxa. [ABSTRACT FROM AUTHOR]

Worldwide, kelp populations are stressed by warming, increased storms and other anthropogenic disturbances. Marine population distributions are projected to retreat poleward with climate change if they cannot adapt to changing conditions, which would potentially lead to a regime shift in subtidal habitats. In Northern Europe, Laminaria hyperborea is a subtidal ecosystem engineer whose distribution has shifted over millennia, leaving predicted areas of high genetic diversity from the last glacial maximum (LGM) near its southern distribution limit in the Iberian Peninsula. In Ireland, L. hyperborea structures communities by supporting diverse faunal assemblages and producing large quantities of organic carbon throughout the year. We investigated the genetic diversity of eight populations, ranging from the southern coast to the north-west of Ireland, using nine microsatellite loci. Diversity was found to be highest in Lough Hyne, a Special Area of Conservation (SAC), near the predicted climate refugium. We found evidence of isolation by distance, with high connectivity between populations that were geographically close, probably driven by short range dispersal of L. hyperborea propagules. Genetic diversity (measured as expected heterozygosity and allelic richness) was highest at Lough Hyne, and decreased northwards, as predicted from past range shifts. Expected heterozygosity was highest at Lough Hyne (0.706) and decreased northward, with the lowest value at Bridges of Ross (0.283). Based on these patterns, further fine-scale investigation into population diversity, dispersal and potential resilience in Irish kelp forests are necessary as warming and non-native species are observed more and more frequently. [ABSTRACT FROM AUTHOR]

The rapid evolution of non-native species can facilitate invasion success, but recent reviews indicate that such microevolution rarely yields expansion of the climatic niche in the introduced habitats. However, because some invasions originate from a geographically restricted portion of the native species range and its climatic niche, it is possible that the frequency, direction, and magnitude of phenotypic evolution during invasion have been underestimated. We explored the utility of niche shift analyses in the red seaweed Gracilaria vermiculophylla, which expanded its range from the northeastern coastline of Japan to North America, Europe, and northwestern Africa within the last 100 years. A genetically informed climatic niche shift analysis indicates that native source populations occur in colder and highly seasonal habitats, while most non-native populations typically occur in warmer, less seasonal habitats. This climatic niche expansion predicts that non-native populations evolved greater tolerance for elevated heat conditions relative to native source populations. We assayed 935 field-collected and 325 common-garden thalli from 40 locations, and as predicted, non-native populations had greater tolerance for ecologically relevant extreme heat (40°C) than did Japanese source populations. Non-native populations also had greater tolerance for cold and low-salinity stresses relative to source populations. The importance of local adaptation to warm temperatures during invasion was reinforced by evolution of parallel clines: Populations from warmer, lower-latitude estuaries had greater heat tolerance than did populations from colder, higher-latitude estuaries in both Japan and eastern North America. We conclude that rapid evolution plays an important role in facilitating the invasion success of this and perhaps other non-native marine species. Genetically informed ecological niche analyses readily generate clear predictions of phenotypic shifts during invasions and may help to resolve debate over the frequency of niche conservatism versus rapid adaptation during invasion. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

The mating system partitions genetic diversity within and among populations and the links between life history traits and mating systems have been extensively studied in diploid organisms. As such most evolutionary theory is focused on species for which sexual reproduction occurs between diploid male and diploid female individuals. However, there are many multicellular organisms with biphasic life cycles in which the haploid stage is prolonged and undergoes substantial somatic development. In particular, biphasic life cycles are found across green, brown and red macroalgae. Yet, few studies have addressed the population structure and genetic diversity in both the haploid and diploid stages in these life cycles. We have developed some broad guidelines with which to develop population genetic studies of haploid-diploid macroalgae and to quantify the relationship between power and sampling strategy. We address three common goals for studying macroalgal population dynamics, including haploid-diploid ratios, genetic structure and paternity analyses. [ABSTRACT FROM AUTHOR]

Haploid-diploid life cycles impose unique eco-evolutionary consequences, rendering commonly used proxies difficult to use (e.g. separate sexes prevent selfing). Population genetic analyses are therefore required to explore patterns of reproductive system variation. However, there are still few haploiddiploid species for which polymorphic, nuclear loci exist. This problem is particularly acute for algae. Here, we describe the development of the first microsatellite loci in a freshwater red alga. We tested 73 candidate loci against a panel of Batrachospermum gelatinosum (L.) De Candolle gametophytes that encompass much of its North American range. Ten loci consistently amplified and were characterized by clean peak architectures on a capillary sequencer with one allele per locus, as expected in a haploid gametophyte. We then explored some basic population genetic indices in gametophytes collected from one site and obtained good resolution based on the probability of identity (pid). Yet, we observed a pattern of clumped repeated genotypes throughout the stream reach sampled. The pattern of moderate genotypic richness could be due to intragametophytic selfing resulting in the complete loss of genetic diversity from a single gamete union. Future studies will need to sample more populations to determine if intragametophytic selfing is the dominant reproductive mode in this monoicous taxon. The loci developed here represent an important tool for studying freshwater red algal populations in specific as well as enhancing our understanding of reproductive system variation and the haploid-diploid life cycle of algae in general. [ABSTRACT FROM AUTHOR]