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Sex is a crucial process that has molecular, genetic, cellular, organismal, and population‐level consequences for eukaryotic evolution. Eukaryotic life cycles are composed of alternating haploid and diploid phases but are constrained by the need to accommodate the phenotypes of these different phases. Critical gaps in our understanding of evolutionary drivers of the diversity in algae life cycles include how selection acts to stabilize and change features of the life cycle. Moreover, most eukaryotes are partially clonal, engaging in both sexual and asexual reproduction. Yet, our understanding of the variation in their reproductive systems is largely based on sexual reproduction in animals or angiosperms. The relative balance of sexual versus asexual reproduction not only controls but also is in turn controlled by standing genetic variability, thereby shaping evolutionary trajectories. Thus, we must quantitatively assess the consequences of the variation in life cycles on reproductive systems. Algae are a polyphyletic group spread across many of the major eukaryotic lineages, providing powerful models by which to resolve this knowledge gap. There is, however, an alarming lack of data about the population genetics of most algae and, therefore, the relative frequency of sexual versus asexual processes. For many algae, the occurrence of sexual reproduction is unknown, observations have been lost in overlooked papers, or data on population genetics do not yet exist. This greatly restricts our ability to forecast the consequences of climate change on algal populations inhabiting terrestrial, aquatic, and marine ecosystems. This perspective summarizes our extant knowledge and provides some future directions to pursue broadly across micro‐ and macroalgal species. [ABSTRACT FROM AUTHOR]

The relative frequency of sexual versus asexual reproduction governs the distribution of genetic diversity within and among populations. Most studies on the consequences of reproductive variation focus on the mating system (i.e., selfing vs. outcrossing) of diploid‐dominant taxa (e.g., angiosperms), often ignoring asexual reproduction. Although reproductive systems are hypothesized to be correlated with life‐cycle types, variation in the relative rates of sexual and asexual reproduction remains poorly characterized across eukaryotes. This is particularly true among the three major lineages of macroalgae (green, brown, and red). The Rhodophyta are particularly interesting, as many taxa have complex haploid–diploid life cycles that influence genetic structure. Though most marine reds have separate sexes, we show that freshwater red macroalgae exhibit patterns of switching between monoicy and dioicy in sister taxa that rival those recently shown in brown macroalgae and in angiosperms. We advocate for the investigation of reproductive system evolution using freshwater reds, as this will expand the life‐cycle types for which these data exist, enabling comparative analyses broadly across eukaryotes. Unlike their marine cousins, species in the Batrachospermales have macroscopic gametophytes attached to filamentous, often microscopic sporophytes. While asexual reproduction through monospores may occur in all freshwater reds, the Compsopogonales are thought to be exclusively asexual. Understanding the evolutionary consequences of selfing and asexual reproduction will aid in our understanding of the evolutionary ecology of all algae and of eukaryotic evolution generally. [ABSTRACT FROM AUTHOR]

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]

Summary: Rhodophyta (or red algae) are a diverse and species‐rich group that forms one of three major lineages in the Archaeplastida, a eukaryotic supergroup whose plastids arose from a single primary endosymbiosis. Red algae are united by several features, such as relatively small intron‐poor genomes and a lack of cytoskeletal structures associated with motility like flagella and centrioles, as well as a highly efficient photosynthetic capacity. Multicellular red algae (or macroalgae) are one of the earliest diverging eukaryotic lineages to have evolved complex multicellularity, yet despite their ecological, evolutionary, and commercial importance, they have remained a largely understudied group of organisms. Considering the increasing availability of red algal genome sequences, we present a broad overview of fundamental aspects of red macroalgal biology and posit on how this is expected to accelerate research in many domains of red algal biology in the coming years. [ABSTRACT FROM AUTHOR]

Phenology, or seasonal variation in life cycle events, is poorly described for many macroalgal species. We describe the phenology of a non‐native population of Gracilaria vermiculophylla whose thalli are free‐living or anchored by decorating polychaetes to tube caps. At a site in South Carolina, USA, we sampled 100 thalli approximately every month from January 2014 to January 2015. We assessed the reproductive state and measured thallus size based on wet weight, thallus length, and thallus surface area from herbarium mounts. Because life cycle stage cannot be assigned using morphology, we implemented a PCR assay to determine the life cycle stage—tetrasporophyte, female gametophyte, or male gametophyte—of each thallus. Tetrasporophytes dominated throughout the year, making up 81%–100% of thalli sampled per month. Reproductive tetrasporophytes varied between 0% and 65% of monthly samples and were most common in warm summer months (July through September) when thalli also tended to be larger. The vast majority of the reproductive thalli were worm‐anchored and not fixed to hard substratum via a holdfast. Thus, free‐living thalli can be reproductive and potentially seed new non‐native populations. Given G. vermiculophylla reproduction seems tied closely to temperature, our work suggests phenology may change with climate‐related changes in seawater temperatures. We also highlight the importance of understanding the natural history of macroalgae to better understand the consequence of range expansions on population dynamics. [ABSTRACT FROM AUTHOR]

Macroalgal (seaweed) genomic resources are generally lacking as compared with other eukaryotic taxa, and this is particularly true in the red algae (Rhodophyta). Understanding red algal genomes is critical to understanding eukaryotic evolution given that red algal genes are spread across eukaryotic lineages from secondary endosymbiosis and red algae diverged early in the Archaeplastids. The Gracilariales is a highly diverse and widely distributed order including species that can serve as ecosystem engineers in intertidal habitats and several notorious introduced species. The genus Gracilaria is cultivated worldwide, in part for its production of agar and other bioactive compounds with downstream pharmaceutical and industrial applications. This genus is also emerging as a model for algal evolutionary ecology. Here, we report new whole-genome assemblies for two species (Gracilaria chilensis and Gracilaria gracilis), a draft genome assembly of Gracilaria caudata , and genome annotation of the previously published Gracilaria vermiculophylla genome. To facilitate accessibility and comparative analysis, we integrated these data in a newly created web-based portal dedicated to red algal genomics (https://rhodoexplorer.sb-roscoff.fr). These genomes will provide a resource for understanding algal biology and, more broadly, eukaryotic evolution. [ABSTRACT FROM AUTHOR]

Accurate species delimitation is crucial to understanding biodiversity. In the northern Gulf of Mexico, recent genetic evidence has suggested that the tricolor Luidia lawrencei is not a species distinct from the gray Luidia clathrata. We collected Luidia specimens from Apalachee Bay, Florida, and morphologically identified 11 as L. clathrata and 16 as L. lawrencei. We sequenced 1074 bp of the cytochrome c oxidase subunit I (COI) and found ~14% divergence between L. clathrata and L. lawrencei , suggesting two distinct species (within-species divergence was <1%). Two specimens were phenotypically L. lawrencei (i.e. , tricolor morph) but mitochondrially were L. clathrata. Our findings lend support to maintaining L. clathrata and L. lawrencei as distinct species. However, the species boundary between these two taxa may be porous, and ongoing hybridization may occur when the two species are found in sympatry. Future work with nuclear markers is warranted to determine the frequency of hybridization and the extent of introgression. Clarifying the genetic relationship between these species will provide a baseline for assessing ongoing changes in connectivity of these two highly abundant sea stars in the rapidly warming northern Gulf of Mexico. [ABSTRACT FROM AUTHOR]

Novel facultative mutualisms that develop between native and non-native ecosystem engineers can lead to the retention of the non-native partner. In some cases, behavior plays an additional, but less understood, role in the development and persistence of mutualisms. In soft-sediment marine habitats along the western Atlantic, the native decorator worm Diopatra cuprea anchors the non-native red alga Gracilaria vermiculophylla to its tube cap in a mutualism. To understand whether the worm's usage of G. vermiculophylla could represent a preference, we first surveyed the species composition of macrophytes affixed to worm tube caps at three sites in coastal Virginia, USA using transect and quadrat sampling. These unmanipulated field surveys supported previous work revealing variable, but often high frequencies (31–98%) of D. cuprea decoration with G. vermiculophylla. We next used field manipulations and controlled laboratory experiments to test the consistency of individual D. cuprea decoration with G. vermiculophylla versus three common macrophytes (Ulva sp., Agardhiella sp., and Spartina alterniflora) found in our field surveys. Twenty-four hours after removing the worm's tube cap in the field, D. cuprea decoration was dominated by both G. vermiculophylla (39.6%) and S. alterniflora (25.9%). When provided a choice of macrophytes in the laboratory, individual D. cuprea consistently decorated with G. vermiculophylla (58.7%) over the other macrophytes, showing a preference for the non-native macrophyte. Our study suggests that preference can drive strong and steadfast interactions between native and non-native organisms, facilitating the latter's persistence and spread, change available habitat, and alter community interactions. [ABSTRACT FROM AUTHOR]

Single‐gene markers, such as the mitochondrial cox1, microsatellites, and single‐nucleotide polymorphisms are powerful methods to describe diversity within and among taxonomic groups and characterize phylogeographic patterns. Large repositories of publicly‐available, molecular data can be combined to generate and evaluate evolutionary hypotheses for many species, including algae. In the case of biological invasions, the combination of different molecular markers has enabled the description of the geographic distribution of invasive lineages. Here, we review the phylogeography of the widespread invasive red macroalga Agarophyton vermiculophyllum (synonym Gracilaria vermiculophylla). The cox1 barcoding provided the first description of the invasion history and hinted at a strong genetic bottleneck during the invasion. Yet, more recent microsatellite and SNP genotyping has not found evidence for bottlenecks and instead suggested that genetically diverse inocula arose from a highly diverse source population, multiple invasions, or some mix of these processes. The bottleneck evident from cox1 barcoding likely reflects the dominance of one mitochondrial lineage, and one haplotype in particular, in the northern source populations in Japan. Recent cox1 sequencing of A. vermiculophyllum has illuminated the complexity of phylogeographic structure in its native range of the northwest Pacific Ocean. For example, the western coast of Honshu in the Sea of Japan displays spatial patterns of haplotypic diversity with multiple lineages found together at the same geographic site. By consolidating the genetic data of this species, we clarify the phylogenetic relationships of a well‐studied macroalga introduced to virtually every temperate estuary of the Northern Hemisphere. [ABSTRACT FROM AUTHOR]

The genomic variation of an invasive species may be affected by complex demographic histories and evolutionary changes during the invasion. Here, we describe the relative influence of bottlenecks, clonality, and population expansion in determining genomic variability of the widespread red macroalga Agarophyton vermiculophyllum. Its introduction from mainland Japan to the estuaries of North America and Europe coincided with shifts from predominantly sexual to partially clonal reproduction and rapid adaptive evolution. A survey of 62,285 SNPs for 351 individuals from 35 populations, aligned to 24 chromosome‐length scaffolds indicate that linkage disequilibrium (LD), observed heterozygosity (Ho), Tajima's D, and nucleotide diversity (Pi) were greater among non‐native than native populations. Evolutionary simulations indicate LD and Tajima's D were consistent with a severe population bottleneck. Also, the increased rate of clonal reproduction in the non‐native range could not have produced the observed patterns by itself but may have magnified the bottleneck effect on LD. Elevated marker diversity in the genetic source populations could have contributed to the increased Ho and Pi observed in the non‐native range. We refined the previous invasion source region to a ~50 km section of northeastern Honshu Island. Outlier detection methods failed to reveal any consistently differentiated loci shared among invaded regions, probably because of the complex A. vermiculophyllum demographic history. Our results reinforce the importance of demographic history, specifically founder effects, in driving genomic variation of invasive populations, even when localized adaptive evolution and reproductive system shifts are observed. [ABSTRACT FROM AUTHOR]

For many taxa, including isomorphic haplodiplontic macroalgae, determining sex and ploidy is challenging, thereby limiting the scope of some population demographic and genetic studies. Here, we used double‐digest restriction site‐associated DNA sequencing (ddRAD‐seq) to identify sex‐linked molecular markers in the widespread red alga Agarophyton vermiculophyllum. In the ddRAD‐seq library, we included 10 female gametophytes, 10 male gametophytes, and 16 tetrasporophytes from one native and one non‐native site (N = 40 gametophytes and N = 32 tetrasporophytes total). We identified seven putatively female‐linked and 19 putatively male‐linked sequences. Four female‐ and eight male‐linked markers amplified in all three life cycle stages. Using one female‐ and one male‐linked marker that were sex‐specific, we developed a duplex PCR and tested the efficacy of this assay on a subset of thalli sampled at two sites in the non‐native range. We confirmed ploidy based on the visual observation of reproductive structures and previous microsatellite genotyping at 10 polymorphic loci. For 32 vegetative thalli, we were able to assign sex and confirm ploidy in these previously genotyped thalli. These markers will be integral to ongoing studies of A. vermiculophyllum invasion. We discuss the utility of RAD‐seq over other approaches previously used, such as RAPDs (random amplified polymorphic DNA), for future work designing sex‐linked markers in other haplodiplontic macroalgae for which genomes are lacking. [ABSTRACT FROM AUTHOR]

Partially clonality is an incredibly common reproductive mode found across all the major eukaryotic lineages. Yet, population genetic theory is based on exclusive sexuality or exclusive asexuality, and partial clonality is often ignored. This is particularly true in haplodiplontic eukaryotes, including algae, ferns, mosses, and fungi, where somatic development occurs in both the haploid and diploid stages. Haplodiplontic life cycles are predicted to be correlated with asexuality, but tests of this prediction are rare. Moreover, there are unique consequences of having long-lived haploid and diploid stages in the same life cycle. For example, clonal processes uncouple the life cycle such that the repetition of the diploid stage via clonality leads to the loss of the haploid stage. Here, we surveyed the literature to find studies that had genotyped both haploid and diploid stages and recalculated population genetic summary metrics for seven red algae, one green alga, three brown algae, and three mosses. We compared these data to recent simulations that explicitly addressed the population genetic consequences of partial clonality in haplodiplontic life cycles. Not only was partial clonality found to act as a homogenizing force, but the combined effects of proportion of haploids, rate of clonality, and the relative strength of mutation versus genetic drift impacts the distributions of population genetic indices. We found remarkably similar patterns across commonly used population genetic metrics between our empirical and recent theoretical expectations. To facilitate future studies, we provide some recommendations for sampling and analyzing population genetic parameters for haplodiplontic taxa. [ABSTRACT FROM AUTHOR]

Organisms across the tree of life have complex life cycles that include both sexual and asexual reproduction or that are obligately asexual. These organisms include ecologically dominant species that structure many terrestrial and marine ecosystems, as well as many pathogens, pests, and invasive species. We must consider both the evolution and maintenance of these various reproductive modes and how these modes shape the genetic diversity, adaptive evolution, and ability to persist in the species that exhibit them. Thus, having a common framework is a key aspect of understanding the biodiversity that shapes our planet. In the 2019 AGA President's Symposium, Sex and Asex: The genetics of complex life cycles , researchers investigating a wide range of taxonomic models and using a variety of modes of investigation coalesced around a common theme—understanding not only how such complex life cycles may evolve, but how they are shaped by the evolutionary and ecological forces around them. In this introduction to the Special Issue from the symposium, we give an overview of some of the key ideas and areas of investigation (a common clonal lexicon , we might say) and introduce the breadth of work submitted by symposium participants. [ABSTRACT FROM AUTHOR]

Species concepts formalize evolutionary and ecological processes, but often conflict with one another when considering the mechanisms that ultimately lead to species delimitation. Evolutionary biologists are, however, recognizing that the conceptualization of a species is separate and distinct from the delimitation of species. Indeed, if species are generally defined as separately evolving metapopulation lineages, then characteristics, such as reproductive isolation or monophyly, can be used as evidence of lineage separation and no longer conflict with the conceptualization of a species. However, little of this discussion has addressed the formalization of this evolutionary conceptual framework for macroalgal species. This may be due to the complexity and variation found in macroalgal life cycles. While macroalgal mating system variation and patterns of hybridization and introgression have been identified, complex algal life cycles generate unique eco‐evolutionary consequences. Moreover, the discovery of frequent macroalgal cryptic speciation has not been accompanied by the study of the evolutionary ecology of those lineages, and, thus, an understanding of the mechanisms underlying such rampant speciation remain elusive. In this perspective, we aim to further the discussion and interest in species concepts and speciation processes in macroalgae. We propose a conceptual framework to enable phycological researchers and students alike to portray these processes in a manner consistent with dialogue at the forefront of evolutionary biology. We define a macroalgal species as an independently evolving metapopulation lineage, whereby we can test for reproductive isolation or the occupation of distinct adaptive zones, among other mechanisms, as secondary lines of supporting evidence. [ABSTRACT FROM AUTHOR]

Worldwide, macroalgae have invaded near‐shore marine ecosystems. However, their haplodiplontic life cycles have complicated efforts to predict patterns of growth and spread, particularly since most theoretical predictions are derived from diplontic taxa (i.e., animals). To complete one revolution of the life cycle, two separate ploidy stages, often including separate haploid sexes, must pass through development and reproduction. In the case of the invasive, red macroalga Agarophyton vermiculophyllum, during the invasion of soft‐sediment estuaries throughout the Northern Hemisphere, diploid tetrasporophytes came to dominate all free‐floating populations and haploid gametophytes were consistently lost. The ecological hypothesis of nutrient limitation might contribute to an explanation of this pattern of tetrasporophytic dominance in free‐floating populations. Under this hypothesis, gametophytes should outperform tetrasporophytes under nutrient limited conditions, but tetrasporophytes should be better able to exploit optimal or even abundant nutrient conditions, such as in eutrophic estuaries. We sampled tetrasporophytes, male gametophytes, and female gametophytes from two sites each located on either side of the Delmarva Peninsula that separates the Chesapeake Bay from the Atlantic Ocean. We subjected apices excised from multiple thalli from each life cycle stage to a nutrient‐enriched and a nutrient‐poor seawater treatment and assessed growth and survival. While nutrient addition increased growth rates, there was no significant difference among ploidies or sexes. Gametophytes did, however, suffer higher mortality than tetrasporophytes. We discuss how nutrient‐dependent differences in growth and survival may contribute to observed patterns of tetrasporophytic dominance in soft‐sediment A. vermiculophyllum populations. [ABSTRACT FROM AUTHOR]

Biological invasions represent grave threats to terrestrial, aquatic, and marine ecosystems, but our understanding of the role of evolution during invasions remains rudimentary. In marine environments, macroalgae account for a large percentage of invaders, but their complicated life cycles render it difficult to move methodologies and predictions wholesale from species with a single, free‐living ploidy stage, such as plants or animals. In haplodiplontic macroalgae, meiosis and fertilization are spatiotemporally separated by long‐lived, multicellular haploid and diploid stages, and gametes are produced by mitosis, not meiosis. As a consequence, there are unique eco‐evolutionary constraints that are not typically considered in invasions. First, selfing can occur in both monoicious (i.e., hermaphroditic) and dioicious (i.e., separate sexes) haplodiplontic macroalgae. In the former, fertilization between gametes produced by the same haploid thallus results in instantaneous, genome‐wide homozygosity. In the latter, cross‐fertilization between separate male and female haploids that share the same diploid parent is analogous to selfing in plants or animals. Separate sexes, therefore, cannot be used as a proxy for outcrossing. Second, selfing likely facilitates invasions (i.e., Baker's law) and the long‐lived haploid stage may enable purging of deleterious mutations, further contributing to invasion success. Third, asexual reproduction will result in the dominance of one ploidy and/or sex and the loss of the other(s). Whether or not sexual reproduction can be recovered depends on which stage is maintained. Finally, fourth, haplodiplontic life cycles are predicted to be maintained through niche differentiation in the haploid and diploid stages. Empirical tests are rare, but fundamental to our understanding of macroalgal invasion dynamics. By highlighting these four phenomena, we can build a framework with which to empirically and theoretically address important gaps in the literature on marine evolutionary ecology, of which biological invasions can serve as unnatural laboratories. [ABSTRACT FROM AUTHOR]

Studies in ecological and community genetics have advanced our understanding of the role of intraspecific diversity in structuring communities and ecosystems. However, in near‐shore marine communities, these studies have mostly been restricted to seagrasses, marsh plants, and oysters. Yet, macroalgae are critically important ecosystem engineers in these communities. Greater intraspecific diversity in a macroalgal ecosystem engineer should result in higher primary and secondary production and community resilience. The paucity of studies investigating the consequences of macroalgal intraspecific genetic variation might be due, in part, to the complexity of macroalgal life cycles. The majority of macroalgae have seemingly subtle, but in actuality, profoundly different life cycles than the more typical animal and angiosperm models. Here, we develop a novel genetic diversity metric, PHD, that incorporates the ratio of gametophytic to sporophytic thalli in natural populations. This metric scales from 0 to 1 like many common genetic diversity metrics, such as genotypic richness, enabling comparisons among metrics. We discuss PHD and examples from the literature, with specific reference to the widespread, red seaweed Agarophyton vermiculophyllum. We also discuss a sex diversity metric, PFM, which also scales from 0 to 1, but fewer studies have identified males and females in natural populations. Nevertheless, by incorporating these novel metrics into the repertoire of diversity metrics, we can explore the role of genetic diversity in community and ecosystem dynamics with an emphasis on the unique biology of many macroalgae, as well as other haplodiplontic taxa such as ferns, foraminiferans, and some fungi. [ABSTRACT FROM AUTHOR]

Theory predicts that the maintenance of haplodiplontic life cycles requires ecological differences between the haploid gametophytes and diploid sporophytes, yet evidence of such differences remain scarce. The haplodiplontic red seaweed Gracilaria vermiculophylla has invaded the temperate estuaries of the Northern Hemisphere, where it commonly modifies detrital and trophic pathways. In native populations, abundant hard substratum enables spore settlement, and gametophyte:tetrasporophyte ratios are ~40:60. In contrast, many non‐native populations persist in soft‐sediment habitats without abundant hard substratum, and can be 90%–100% tetrasporophytic. To test for ecologically relevant phenotypic differences, we measured thallus morphology, protein content, organic content, “debranching resistance” (i.e., tensile force required to remove a branch from its main axis node), and material properties between male gametophytes, female gametophytes, and tetrasporophytes from a single, nonnative site in Charleston Harbor, South Carolina, USA in 2015 and 2016. Thallus length and surface area to volume ratio differed between years, but were not significantly different between ploidies. Tetrasporophytes had lower protein content than gametophytes, suggesting the latter may be more attractive to consumers. More force was required to pull a branch from the main axis of tetrasporophytes relative to gametophytes. A difference in debranching resistance may help to maintain tetrasporophyte thallus durability relative to gametophytes, providing a potential advantage in free‐floating populations. These data may shed light on the invasion ecology of an important ecosystem engineer, and may advance our understanding of life cycle evolution and the maintenance of life cycle diversity. [ABSTRACT FROM AUTHOR]

Our knowledge of non-native algae in benthic estuarine habitats is relatively poor, especially compared to algal introductions along open shores or on floating structures. Gracilaria vermiculophylla is a widespread macroalgal invader in the temperate estuaries of the Northern Hemisphere, and, here, we expand its documented range within northeastern Ireland and England. Established populations occur within two inlets in the border counties, Carlingford Lough (Counties Louth and Down) and Dundrum Bay (County Down), but G. vermiculophylla is absent from open coasts between these sites. Repeated surveys in Dundrum Bay showed variable abundances, with an increase in biomass between 2013 and 2016. Three populations were discovered in England, where this species had not previously been identified: Christchurch Harbour (Dorset), Brownsea Island in Poole Harbour (Dorset), and Kingsbridge Estuary (Devon). The Irish and English thalli belong to the most common, invasive cox 1 haplotype 6. Using a combination of morphological observations and 10 microsatellite loci, we found that the population at Carlingford Lough included both reproductive haploid gametophytes and diploid tetrasporophytes and genetic signatures of sexual reproduction, but the populations at Christchurch and Brownsea displayed signatures of partial clonality. Genetic diversity was higher along the south coast of England as compared to the Irish population, consistent with patterns of diversity previously described for the European coasts. Finally, we also note the occurrence of a putative G. vermiculophylla population in Wales at Porthmadog, Gwynedd. As the sites in which we have now documented G. vermiculophylla in the British Isles also host shellfish aquaculture activities, our study is further evidence for the role of aquaculture in the spread of invasive species. [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]

The delimitation of species in the Gracilariales is often difficult due to the lack of diagnostic morphological characters. As a result, non-native species are often misidentified without the use of molecular tools. Recently, studies have investigated the agar quality of the dominant gracilarioid species in the Laguna San Ignacio in Baja Califronia Sur, Mexico, including Vergara-Rodarte et al., (2016) published in the May edition of Cryptogamie, Algologie. The species has been reported as Gracilaria pacifica, G. vermiculophylla and Gracilariopsis sp. Using a combination of three mitochondrial and plastid markers, we identified this species as G. parvispora, extending the known distribution of this non-native species in Baja California. Due to the potential of G. parvispora as a source of agar, more detailed studies of on the invasion history and surveys are necessary in the eastern Pacific to determine its current distribution and impacts on the native biodiversity. [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]

Species with sexual and asexual life cycles may exhibit intraspecific differences in reproductive effort. The spatial separation of sexual and asexual lineages, called geographic parthenogenesis, is common in plants, animals, and algae. Mastocarpus papillatus is a well-documented case of geographic parthenogenesis in which sexuals dominate southern populations, asexuals dominate northern popula-tions, whereas mixed populations occur throughout central California. We quantified abundances and reproductive effort of sexual and asexual fronds and tetrasporophytes at eight sites in California to test the hypotheses that (1) reduced sexual reproduction at higher latitudes and tidal heights explains the observed geographic parthenogenesis and (2) reproductive effort (spore production per blade area) declines with increasing latitude. Abundances of all phases varied site-specifically. However, there was no geographic pattern of reproductive effort of fronds. Reproductive effort of fronds was greater in 2006 than in 2007 and correlated with sea surface temperatures. Sexual fronds exhibited greater reproductive effort than did asexual fronds and sexual reproductive effort was also inversely correlated with local upwelling index. Tetrasporophytes showed greater repro-ductive effort in northern sites, but total supply of tetraspores per m2 was greatest in the middle of the sampling range where crusts were more abundant. There was no decline of reproductive effort at higher latitudes. Geographic patterns of fecundity of life stages do not explain geographic parthe-nogenesis in M. papillatus. Site-specific differences in viability among spores or established thalli of different life cycles may explain their respective geographic distributions, as the sexual and asexual life cycles responded differently to environmental variations. [ABSTRACT FROM AUTHOR]

Chondrus crispus Stackh. has been intensely studied, yet no study to date has elucidated its population structure or mating system despite many populations in which there was a haploid bias and lack of male gametophytes. Therefore, 12 nuclear microsatellite loci were identified in this red alga. Microsatellite markers were developed and tested against a panel of specimens collected from two shore levels at two sites in Brittany, France: Pointe de Primel and Pointe de la Jument, Concarneau. Single locus genetic determinism was verified at eight polymorphic loci, as only one band was observed for haploid genotypes, whereas one or two bands were observed for diploids. These markers enabled the detection of unique genotypes within sampled populations, indicating that very few fronds shared the same multilocus genotype. This finding suggests that asexual reproduction was not the prevailing mode of reproduction. In addition, we explored the hierarchical population structure showing that gene flow is restricted at small spatial scales (<50 m) between upper and lower Chondrus-range populations within a shore. Sexual reproduction predominated in the populations of C. crispus studied, but probably due to fine-scale spatial substructuring, inbreeding was also significant. In conclusion, this study reveals that fine-scale genetic variation is of major importance in C. crispus, suggesting that differences between microhabitats should be essential in understanding evolutionary processes in this species. [ABSTRACT FROM AUTHOR]

Chlainomonas (Chlamydomonadales, Chlorophyta) is one of the four genera of snow algae known to produce annual pink or red blooms in alpine snow. No Chlainomonas species have been successfully cultured in the laboratory, but diverse cell types have been observed from many field‐collected samples, from multiple species. The diversity of morphologies suggests these algae have complex life cycles with changes in ploidy. Over 7 years (2017–2023), we observed seasonal blooms dominated by a Chlainomonas species from late spring through the summer months on a snow‐on‐lake habitat in an alpine basin in the North Cascade Mountains of Washington, USA. The Bagley Lake Chlainomonas is distinct from previously reported species based on morphology and sequence data. We observed a similar collection of cell types observed in other Chlainomonas species, with the addition of swarming biflagellate cells that emerged from sporangia. We present a life cycle hypothesis for this species that links cell morphologies observed in the field to seasonally available habitat. The progression of cell types suggests cells are undergoing both meiosis and fertilization in the life cycle. Since the life cycle is the most fundamental biological feature of an organism, with direct consequences for evolutionary processes, it is critical to understand how snow algal life cycles will influence their responses to changes in their habitat driven by climate warming. For microbial taxa that live in extreme environments and are difficult to culture, temporal field studies, such as we report here, may be key to creating testable hypotheses for life cycles. [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]

Complex life cycles, in which discrete life stages of the same organism differ in form or function and often occupy different ecological niches, are common in nature. Because stages share the same genome, selective effects on one stage may have cascading consequences through the entire life cycle. Theoretical and empirical studies have not yet generated clear predictions about how life cycle complexity will influence patterns of adaptation in response to rapidly changing environments or tested theoretical predictions for fitness trade-offs (or lack thereof) across life stages. We discuss complex life cycle evolution and outline three hypotheses--ontogenetic decoupling, antagonistic ontogenetic pleiotropy and synergistic ontogenetic pleiotropy--for how selection may operate on organisms with complex life cycles. We suggest a within-generation experimental design that promises significant insight into composite selection across life cycle stages. As part of this design, we conducted simulations to determine the power needed to detect selection across a life cycle using a population genetic framework. This analysis demonstrated that recently published studies reporting withingeneration selection were underpowered to detect small allele frequency changes (approx. 0.1). The power analysis indicates challenging but attainable sampling requirements for many systems, though plants and marine invertebrates with high fecundity are excellent systems for exploring how organisms with complex life cycles may adapt to climate change. [ABSTRACT FROM AUTHOR]

Theoretical ecological models, such as succession and facilitation, were defined in terrestrial habitats, and subsequently applied to marine and freshwater habitats in intertidal and then subtidal realms. One such model is the soil seed bank, defined as all viable seeds (or fruits) found near the soil surface that facilitate community restoration/recovery. "Banks of microscopic forms" have been hypothesized in aquatic habitats and recent work from aquaculture has highlighted dormancy in algal life cycle stages. To reinvigorate the discussions about these algal banks, we discuss differences in life cycles, dispersal, and summarize research on banks of macroalgal stages in aquatic ecosystems that may be easier to explore with modern advances in molecular technology. With focus on seminal work in global kelp forest ecosystems, we present a pilot study in northern California as proof of concept that Nereocystis luetkeana and Alaria marginata stages can be detected within kelp forests in the biofilm of rocks and bedrock using targeted primers long after zoospore release. Considering the increased interest in algae as an economic resource, [blue] carbon sink, and as ecosystem engineers, the potential for "banking" macroalgal forms could be a mechanism of resilience and recovery in aquatic populations that have complex life cycles and environmental cues for reproduction. Molecular barcoding is becoming an important tool for identifying banks of macroalgal forms in marine communities. Understanding banks of macroalgal stages, especially in deforested habitats with intense disturbance and grazer pressure, will allow researchers and marine resource managers to facilitate this natural process in recovery of the aquatic system. [ABSTRACT FROM AUTHOR]

Snow algal blooms frequently occur throughout alpine and polar environments during spring and summer months; however, our understanding of bloom dynamics is limited. We tracked a recurrent bloom of Chlainomonas sp. on Upper Bagley Lake in the North Cascade Mountains, USA, to assess the spatiotemporal dynamics in bloom color intensity, community photophysiology, and community composition over eight weeks. We found that the algae biomass had a dynamic patchy distribution over space and time, which was decoupled from changes in community composition and life-cycle progress averaged across the bloom. The proportional representation of Chlainomonas sp. remained consistent throughout the study while the overall community composition shows a progression through the bloom. We found that community photophysiology, measured by the maximum quantum yield of PSII (Fv/Fm), decreased on average throughout the bloom. These findings suggest that the Chlainomonas sp. community on Bagley Lake is not simply an algal bloom with rapid increase in biomass followed by a population crash, as is often seen in aquatic systems, though there is a physiological trajectory and sensitivity to environmental stress. These results contribute to our understanding of the biology of Chlainomonas sp. and its response to environmental stress, specifically an extreme warming event. [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]

Ecological and evolutionary processes differ depending on how genetic diversity is organized in space. For clonal organisms, the organization of both genetic and genotypic diversity can influence the fitness effects of competition, the mating system, and reproductive mode, which are key drivers of life cycle evolution. Understanding how individual reproductive behavior contributes to population genetic structure is essential for disentangling these forces, particularly in species with complex and plastic life cycles. The widespread sea anemone, Diadumene lineata , exhibits temperature-dependent fission, which contributes to predictable variation in clonal rate along the Atlantic coast of the United States, part of its non-native range. Because warmer conditions lead to higher rates of clonality, we expected to find lower genotypic and genetic diversity in lower versus higher latitude populations. We developed primers for 11 microsatellite loci and genotyped 207 anemones collected from 8 sites ranging from Florida to Massachusetts. We found clonal influence at all sites, and as predicted, the largest clones were found at lower latitude sites. We also found genetic signatures of sex in the parts of the range where gametogenesis is most common. Evidence of sex outside the native range is novel for this species and provides insights into the dynamics of this successful invader. Our findings also illustrate challenges that partially clonal taxa pose for eco-evolutionary studies, such as difficulty sampling statistically robust numbers of genets and interpretating common population genetic metrics. For example, we found high among-locus variation in F IS , which makes the meaning of mean multilocus F IS unclear. [ABSTRACT FROM AUTHOR]

Generalist consumers potentially limit introduced prey populations when they are willing and able to consume those prey, but because generalist consumers vary in feeding preference, they are not equally effective in resisting invasions. Previous work suggested that during the introduction of the red seaweed Gracilaria vermiculophylla to North American and European shorelines, populations evolved enhanced levels of constitutive and activated prostaglandin-based defenses against Littorina snails and the isopod Idotea balthica. However, when we offered tissue from a total of 700 G. vermiculophylla thalli from 14 native Japanese and 25 non-native sites to a North American population of the generalist amphipod Ampithoe valida, native and non-native populations were consumed at statistically indistinguishable rates. In contrast to previous results using Idotea and Littorina, we found no evidence that mechanical wounding of G. vermiculophylla altered palatability of thalli to A. valida. These results indicate either the strength of anti-herbivore defense varies across the geographic range in the seaweed’s native range in the Northwest Pacific, that the effectiveness of G. vermiculophylla to deter herbivores varies across generalist herbivore species, or both. In either case, we question the conclusion that rapid phenotypic shift in anti-herbivory traits to generalist herbivores occurred in this species. Future studies should focus on whether shifts in anti-herbivore defenses are adaptive by experimentally assessing species-specific herbivore impacts in native and non-native ranges. [ABSTRACT FROM AUTHOR]

Chondrus crispus is a marine red alga with sulfated galactans, called carrageenans, in its extracellular matrix. Chondrus has a complex haplodiplontic life cycle, alternating between male and female gametophytes (n) and tetrasporophytes (2n). The Chondrus life cycle stages are isomorphic; however, a major phenotypic difference is that carrageenan composition varies significantly between the tetrasporophytes (mainly lambda-carrageenan) and the gametophytes (mainly kappa/iota-carrageenans). The disparity in carrageenan structures, which confer different chemical properties, strongly suggests differential regulation of carrageenan-active genes between the phases of the Chondrus life cycles. We used a combination of taxonomy, biochemistry and molecular biology to characterize the tetrasporophytes and male and female gametophytes from Chondrus individuals isolated from the rocky seashore off the northern coast of France. Transcriptomic analyses reveal differential gene expression of genes encoding several galactose-sulfurylases, carbohydrate-sulfotransferases, glycosyltransferases, and one family 16 glycoside hydrolase. Differential expression of carrageenan-related genes was found primarily between gametophytes and tetrasporophytes, but also between the male and female gametophytes. The differential expression of these multigenic genes provides a rare glimpse into cell wall biosynthesis in algae. Furthermore, it strongly supports that carrageenan metabolism holds an important role in the physiological differentiation between the isomorphic life cycle stages of Chondrus. [ABSTRACT FROM AUTHOR]

Communities are shaped by scale dependent processes. To study the diversity and variation of microbial communities across scales, the invasive and widespread seaweed Agarophyton vermiculophyllum presents a unique opportunity. We characterized pro‐ and eukaryotic communities associated with this holobiont across its known distribution range, which stretches over the northern hemisphere. Our data reveal that community composition and diversity in the holobiont vary at local but also larger geographic scales. While processes acting at the local scale (i.e., within population) are the main structuring drivers of associated microbial communities, changes in community composition also depend on processes acting at larger geographic scales. Interestingly, the largest analysed scale (i.e., native and non‐native ranges) explained variation in the prevalence of predicted functional groups, which could suggest a functional shift in microbiota occurred over the course of the invasion process. While high variability in microbiota at the local scale supports A. vermiculophyllum to be a generalist host, we also identified a number of core taxa. These geographically independent holobiont members imply that cointroduction of specific microbiota may have additionally promoted the invasion process. [ABSTRACT FROM AUTHOR]

Kelp restoration practices have renewed urgency given worldwide kelp declines (e.g., Eger, Layton, et al., [5]; Eger, Marzinelli, et al., [6]). Abbreviations MCP major capsid protein ONT Oxford Nanopore Technologies NGS Next Generation Sequencing OPC Ocean Protection Council PCR polymerase chain reaction Kelp forests are ecologically and economically valuable marine ecosystems located worldwide. Indeed, we lack information on the role viruses might have in shaping kelp ecology and evolution, including on short timescales during kelp population loss and recovery. [Extracted from the article]

Meiosis and syngamy generate an alternation between two ploidy stages, but the timing of these two processes varies widely across taxa, thereby generating life cycle diversity. One hypothesis suggests that life cycles with long‐lived haploid stages are correlated with selfing, asexual reproduction, or both. Though mostly studied in angiosperms, selfing and asexual reproduction are often associated with marginal habitats. Yet, in haploid‐diploid macroalgae, these two reproductive modes have subtle but unique consequences whereby predictions from angiosperms may not apply. Along the western Antarctic Peninsula, there is a thriving macroalgal community, providing an opportunity to explore reproductive system variation in haploid‐diploid macroalgae at high latitudes where endemism is common. Plocamium sp. is a widespread and abundant red macroalga observed within this ecosystem. We sampled 12 sites during the 2017 and 2018 field seasons and used 10 microsatellite loci to describe the reproductive system. Overall genotypic richness and evenness were high, suggesting sexual reproduction. Eight sites were dominated by tetrasporophytes, but there was strong heterozygote deficiency, suggesting intergametophytic selfing. We observed slight differences in the prevailing reproductive mode among sites, possibly due to local conditions (e.g., disturbance) that may contribute to site‐specific variation. It remains to be determined whether high levels of selfing are characteristic of macroalgae more generally at high latitudes, due to the haploid‐diploid life cycle, or both. Further investigations of algal life cycles will likely reveal the processes underlying the maintenance of sexual reproduction more broadly across eukaryotes, but more studies of natural populations are required. [ABSTRACT FROM AUTHOR]

The analysis of biological and ecological traits has a long history in evolutionary and ecological research. However, trait data are often scattered and standardised terminology that transcends taxonomic and biogeographical context are generally missing. As part of the development of a global trait database of marine species, we collated trait information for European seaweeds and structured the data within the standardised framework of the World Register of Marine Species (WoRMS). We collected 45 175 trait records for 21 biologically and ecologically relevant traits of seaweeds. This resulted in a trait database for 1745 European seaweed species of which more than half (56 %) of the records were documented at the species level, while the remaining 44 % were documented at a higher taxonomic level and subsequently inherited at lower levels. The trait database for European seaweeds will serve as a foundation for future research on diversity and evolution of seaweeds and their responses to global changes. The data will contribute to developing detailed trait-based ecosystem models and will be an important tool to inform marine conservation policies. The data are publicly accessible through the AlgaeTraits portal, 10.14284/574 (AlgaeTraits, 2022). [ABSTRACT FROM AUTHOR]

Invasive species can successfully and rapidly colonize new niches and expand ranges via founder effects and enhanced tolerance towards environmental stresses. However, the underpinning molecular mechanisms (i.e., gene expression changes) facilitating rapid adaptation to harsh environments are still poorly understood. The red seaweed Gracilaria vermiculophylla, which is native to the northwest Pacific but invaded North American and European coastal habitats over the last 100 years, provides an excellent model to examine whether enhanced tolerance at the level of gene expression contributed to its invasion success. We collected G. vermiculophylla from its native range in Japan and from two non‐native regions along the Delmarva Peninsula (Eastern United States) and in Germany. Thalli were reared in a common garden for 4 months at which time we performed comparative transcriptome (mRNA) and microRNA (miRNA) sequencing. MRNA‐expression profiling identified 59 genes that were differently expressed between native and non‐native thalli. Of these genes, most were involved in metabolic pathways, including photosynthesis, abiotic stress, and biosynthesis of products and hormones in all four non‐native sites. MiRNA‐based target‐gene correlation analysis in native/non‐native pairs revealed that some target genes are positively or negatively regulated via epigenetic mechanisms. Importantly, these genes are mostly associated with metabolism and defence capability (e.g., metal transporter Nramp5, senescence‐associated protein, cell wall‐associated hydrolase, ycf68 protein and cytochrome P450‐like TBP). Thus, our gene expression results indicate that resource reallocation to metabolic processes is most likely a predominant mechanism contributing to the range‐wide persistence and adaptation of G. vermiculophylla in the invaded range. This study, therefore, provides molecular insight into the speed and nature of invasion‐mediated rapid adaption. [ABSTRACT FROM AUTHOR]

Seaweeds are colonized by a microbial community, which can be directly linked to their performance. This community is shaped by an interplay of stochastic and deterministic processes, including mechanisms which the holobiont host deploys to manipulate its associated microbiota. The Anna Karenina principle predicts that when a holobiont is exposed to suboptimal or stressful conditions, these host mechanisms may be compromised. This leads to a relative increase of stochastic processes that may potentially result in the succession of a microbial community harmful to the host. Based on this principle, we used the variability in microbial communities (i.e., beta diversity) as a proxy for stability within the invasive holobiont Gracilaria vermiculophylla during a simulated invasion in a common garden experiment. Independent of host range, host performance declined at elevated temperature (22°C) and disease incidence and beta diversity increased. Under thermally stressful conditions, beta diversity increased more in epibiota from native populations, suggesting that epibiota from non‐native holobionts are thermally more stable. This pattern reflects an increase in deterministic processes acting on epibiota associated with non‐native hosts, which in the setting of a common garden can be assumed to originate from the host itself. Therefore, these experimental data suggest that the invasion process may have selected for hosts better able to maintain stable microbiota during stress. Future studies are needed to identify the underlying host mechanisms. [ABSTRACT FROM AUTHOR]

The analysis of biological and ecological traits has a long history in evolutionary and ecological research. However, trait data are often scattered and standardised terminology that transcends taxonomic and biogeographical context are generally missing. As part of the development of a global trait database of marine species, we collated trait information for European seaweeds and structured the data within the standardised framework of the World Register of Marine Species (WoRMS). We collected 45,175 records for 21 biologically and ecologically relevant traits of seaweeds. This resulted in a trait database for 1,745 European seaweed species of which more than half (56%) of the records were documented at the species level, while the remaining 44% was documented at a higher taxonomic level and subsequently inherited at lower levels. The trait database for European seaweeds will serve as a foundation for future research on diversity and evolution of seaweeds, and their responses to global changes. The data will contribute to developing detailed trait-based ecosystem models, and will be an important tool to inform marine conservation policies. The data is publicly accessible through the AlgaeTraits portal, algaetraits.org, doi: https://doi.org/10.14284/574, (AlgaeTraits, 2022). [ABSTRACT FROM AUTHOR]

Fungal symbionts of terrestrial plants are among the most widespread and well‐studied symbioses, relatively little is known about fungi that are associated with macroalgae. To fill the gap in marine fungal taxonomy, we combined simple culture methods with amplicon sequencing to characterize the fungal communities associated with three brown (Sargassum muticum, Pelvetia canaliculata, and Himanthalia elongata) and two red (Mastocarpus stellatus and Chondrus crispus) macroalgae from one intertidal zone. In addition to characterizing novel fungal diversity, we tested three hypotheses: fungal diversity and community composition vary (i) among species distributed at different tidal heights, (ii) among tissue types (apices, mid‐thallus, and stipe), and (iii) among "isomorphic" C. crispus life cycle stages. Almost 70% of our reads were classified as Ascomycota, 29% as Basidiomycota, and 1% that could not be classified to a phylum. Thirty fungal isolates were obtained, 18 of which were also detected with amplicon sequencing. Fungal communities differed by host and tissue type. Interestingly, P. canaliculata, a fucoid at the extreme high intertidal, did not show differences in fungal diversity across the thallus. As found in filamentous algal endophytes, fungal diversity varied among the three life cycle stages in C. crispus. Female gametophytes were also compositionally more dispersed as compared to the fewer variable tetrasporophytes and male gametophytes. We demonstrate the utility of combining relatively simple cultivation and sequencing approaches to characterize and study macroalgal–fungal associations and highlight the need to understand the role of fungi in near‐shore marine ecosystems. [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]

Aim: Kelp forests worldwide are important marine ecosystems that foster high primary to secondary productivity and multiple ecosystem services. These ecosystems are increasingly under threat from extreme storms, changing ocean temperatures, harvesting, and greater herbivore pressure at regional and global scales, necessitating urgent documentation of their historical to present‐day distributions. Species range shifts to higher latitudes have already been documented in some species that dominate subtidal habitats within Europe. Very little is known about kelp forest ecosystems in Ireland, where rocky coastlines are dominated by Laminaria hyperborea. In order to rectify this substantial knowledge gap, we compiled historical records from an array of sources to present historical distribution, kelp and kelp forest recording effort over time, and present rational for the monitoring of kelp habitats to better understand ecosystem resilience. Location: Ireland (Northern Ireland and Éire). Methods: Herbaria, literature from the Linnaean society dating back to late 1700s, journal articles, government reports, and online databases were scoured for information on L. hyperborea. Information about kelp ecosystems was solicited from dive clubs and citizen science groups that are active along Ireland's coastlines. Results: Data were used to create distribution maps and analyze methodology and technology used to record L. hyperborea presence and kelp ecosystems within Ireland. We discuss the recent surge in studies on Irish kelp ecosystems, fauna associated with kelp ecosystems that may be used as indicators of ecosystem health and suggest methodologies for continued monitoring. Main Conclusions: While there has been a steady increase in recording effort of the dominant subtidal kelp forest species, L. hyperborea, only recently have studies begun to address other important eco‐evolutionary processes at work in kelp forests including connectivity among kelp populations in Ireland. Further monitoring, using suggested methodologies, is required to better understand the resilience of kelp ecosystems in Ireland. [ABSTRACT FROM AUTHOR]

Inbreeding is a potent evolutionary force shaping the distribution of genetic variation within and among populations of plants and animals. Yet, our understanding of the forces shaping the expression and evolution of nonrandom mating in general, and inbreeding in particular, remains remarkably incomplete. Most research on plant mating systems focuses on self‐fertilization and its consequences for automatic selection, inbreeding depression, purging, and reproductive assurance, whereas studies of animal mating systems have often assumed that inbreeding is rare, and that natural selection favors traits that promote outbreeding. Given that many sessile and sedentary marine invertebrates and marine macroalgae share key life history features with seed plants (e.g., low mobility, modular construction, and the release of gametes into the environment), their mating systems may be similar. Here, we show that published estimates of inbreeding coefficients (FIS) for sessile and sedentary marine organisms are similar and at least as high as noted in terrestrial seed plants. We also found that variation in FIS within invertebrates is related to the potential to self‐fertilize, disperse, and choose mates. The similarity of FIS for these organismal groups suggests that inbreeding could play a larger role in the evolution of sessile and sedentary marine organisms than is currently recognized. Specifically, associations between traits of marine invertebrates and FIS suggest that inbreeding could drive evolutionary transitions between hermaphroditism and separate sexes, direct development and multiphasic life cycles, and external and internal fertilization. [ABSTRACT FROM AUTHOR]

Pigment variation within and among algal species may have important ecological consequences because small changes in the concentration and composition of pigments can influence the photosynthetic efficiency and rate as well as the spectra of light utilized. Toward the goal of developing a rapid method for comparing pigment composition among algal thalli, we characterized the relationship between visual color information taken from photographs (e.g., red, green, and blue color values) and photopigment composition in the non-native red alga Agarophyton vermiculophyllum (Ohmi) Gurgel, J.N.Norris & Fredericq. We used a set of 19 thalli, collected from across the known native and non-native range in the Northern Hemisphere, which exhibited substantial color variation at the time of field collection, and sustained this variation after being maintained in a common garden. We identified a set of ecologically interesting pigment traits that are readily predicted by color information, including chlorophyll a and phycobilin concentration. Finally, we demonstrated the repeatability of estimating color phenotypes from photographs of thalli taken under a range of light conditions in order to evaluate the utility of this approach for field studies. We suggest this method could be useful for the rapid, high-throughput phenotyping of photopigments in other red algae as well. [ABSTRACT FROM AUTHOR]

Clonal behavior has been hypothesized to provide an escape from allometric metabolic scaling that limits the maximum mass achieved by a single individual. Here, we demonstrate the capacity of a wide-spread, non-native sea anemone to buffer its colony biomass accumulation rate across environments by modulating ramet body size through environmentally dependent growth, fission, and catabolism. In 2015, thermal reaction norms for growth and fission behavior were constructed using clonal lines of the sea anemone Diadumene lineata. In 2018, variation in growth patterns under a factorial cross of temperature level and oxygen availability was examined to test the hypothesis that individual ramet size is regulated by oxygen limitation in accordance with optimal size theory. Across a wide range of temperatures, colonies accumulated a similar amount of biomass despite a radical shift from unitary to clonal growth, supporting fission as a mechanism to buffer growth rates over a range of conditions. Individual body size appears to be regulated by the environment with increased temperature and reduced oxygen modifying fission and mass-specific growth patterns, leading to the production of smaller-bodied ramets in warm conditions. However, whether anemones in common garden conditions reduce individual body size through catabolism or fission depends on the region of origin and may relate to differences in seasonal temperature patterns among coastlines, which influence the energetic benefits of fission rate plasticity. [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 susceptibility of native and non-native populations of the red alga Gracilaria vermiculophylla to fouling was compared in common garden experiments. Native and non-native algae were enclosed into dialysis membrane tubes, and the tubes were exposed to natural fouling. Fouling on the outside of the tubes was mediated by chemical compounds excreted by G. vermiculophylla that diffused through the membranes. Fouling pressure was significantly higher in the Kiel Fjord (non-native range) than in Akkeshi Bay (native range), but, at both sites, tubes containing non-native G. vermiculophylla were less fouled than those with native conspecifics. This is the first in situ evidence that susceptibility to fouling differs between native and non-native populations of an aquatic organism. The technique of enclosing organisms into dialysis tubes represents a simple, efficient and accurate way to test chemical antifouling defenses and could possibly be applied to other organisms. [ABSTRACT FROM AUTHOR]

Chondrus crispus, or Irish moss, is a common edible red seaweed that can be found on rocky shores in the Northern Atlantic. The cell wall contains carrageenan and C. crispus is the original source of this commercially used thickener. Because of the ecological and economic importance of this red alga a relatively important research literature exists and one of the recent achievements in C. crispus research is the sequencing of its genome. In this chapter we review some of the literature with the aim to promote C. crispus as a model organism for florideophyte red seaweeds. We consider subjects like commercial and historical uses, ecology, genetics, population structure, mating systems, physiology, cell wall biology and genomics. [ABSTRACT FROM AUTHOR]