Accumulation and bio-oxidation of arsenite mediated by thermoacidophilic Cyanidiales: innate potential biomaterials toward arsenic remediation.

Addressing geogenic and anthropogenic arsenic (As) pollution is critical for environmental health. This study explored arsenite [As(III)] removal using Cyanidiales, particularly Cyanidium caldarium (Cc) and Galdieria partita (Gp), under acidic to neutral pH, and determined As(III) detoxification mechanisms in relation to As speciation and protein secondary structure in Cyanidiales. Regarding As(III) sorption amounts, Cc outperformed Gp, reaching 83.2 mg g-1 of removal at pH 5.0. Wherein, 23.5 % of sorbed As on Cc presented as arsenate [As(V)] complexation with polysaccharides, alongside other predominant species including As(III)-cysteine (41.2 %) and As(III)-polysaccharides (35.3 %) complexes. This suggested that As(III) was directly transported into cells, rather than As(V). Coupled with the formation of As(III)-cysteine complexes within cells, these mechanisms may be key to efficiently accumulating As(III) in Cyanidiales during the 6-h incubation. These results highlight the potential of Cyanidiales for sustainable As(III) remediation and provide new insights into managing As(III) toxicity.

Testing sea urchin and green sea turtle consumption of the allelopathic macroalga Galaxaura divaricata.

Galaxaura divaricata is a partially calcified macroalga that hampers coral recruitment, growth, and recovery via the excretion of allelopathic secondary metabolites. Herbivorous fishes are not major consumers of Galaxaura spp. and there is a need to understand feeding preferences for Galaxaura divaricata in other macroherbivores, like sea urchins and green sea turtles that could act as potential controlling agents. Under certain environmental conditions, G. divaricata can proliferate and overgrow degraded reefs for several years, as documented for several coral patch reefs in the lagoon of Dongsha Atoll, South China Sea. This study aimed to experimentally test the feeding preferences of five species of sea urchin and two individual green sea turtles, Chelonia mydas, for G. divaricata. Specifically, we quantified and compared the consumption rates of the allelopathic G. divaricata with Gracilaria edulis, a nonallelopathic, fleshy red alga, known to be highly favored by herbivores. Results showed that the five urchin species fed on both G. edulis and G. divaricata. However, urchins consumed 2-8 times less wet weight of G. divaricata (range 0.3-3.1 g urchin-1 24 h-1) compared to G. edulis (range 0.6-18 g urchin-1 24 h-1), suggesting that urchin grazing may exert some control on G. divaricata abundance but is likely ineffective for a large-scale removal of the alga. Further, both green sea turtles avoided G. divaricata and selectively fed on G. edulis. More experiments are needed to test the potential role of herbivores in controlling the overgrowth of coral competitive and allelopathic macroalgae, like Galaxaura on coral reefs.

Water depth outweighs reef condition in shaping non-geniculate coralline algae-associated microbial communities in coral reefs: A case study from Thailand.

Red calcified non-geniculate coralline algae (NGCA) provide habitat structures, stabilize reef structures, and foster coral larval settlement and metamorphosis. Moreover, the microbes associated with NGCA are dependent on the NGCA host species and are affected by environmental factors; however, little is known about the influence of reef conditions and depth gradients on the associated microbial communities and NGCA. In this study, we collected NGCA under different reef conditions and depth gradients and characterized the microbial communities using the V3-V4 hypervariable regions of the 16S rRNA gene. Metagenomic analysis revealed 2 domains, 51 phyla, 123 classes, and 210 genera. The NGCA-associated bacterial communities were dominated by Proteobacteria, Bacteroidetes, Chloroflexi, Actinobacteria, and Acidobacteriota. Gammaproteobacteria and Alphaproteobacteria were the most abundant bacterial classes. Differences in microbial diversity and richness were not apparent between reef conditions and depth gradients. However, there was a significant difference in bacterial evenness among the depth gradients. The bacterial abundance associated with NGCA was greater in deep zones than in shallow zones. The shallow zone exhibited a greater relative abundance of all gene functions than the deep zone, indicating differences in the distribution of gene functions. This study showed that the microbial communities associated with red calcified NGCA are diverse, and that the depth gradient affects their abundance and evenness, highlighting the need for further research to understand the functional roles of these microbial communities in coral reef conservation.

Revised classification of the Cyanidiophyceae based on plastid genome data with descriptions of the Cavernulicolales ord. nov. and Galdieriales ord. nov. (Rhodophyta).

The Cyanidiophyceae, an extremophilic red algal class, is distributed worldwide in extreme environments. Species grow either in acidic hot environments or in dim light conditions (e.g., "cave Cyanidium"). The taxonomy and classification systems are currently based on morphological, eco-physiological, and molecular phylogenetic characters; however, previous phylogenetic results showed hidden diversity of the Cyanidiophyceae and suggested a revision of the classification system. To clarify phylogenetic relationships within this red algal class, we employ a phylogenomic approach based on 15 plastomes (10 new) and 15 mitogenomes (seven new). Our phylogenies show consistent relationships among four lineages (Galdieria, "cave Cyanidium", Cyanidium, and Cyanidioschyzon lineages). Each lineage is distinguished by organellar genome characteristics. The "cave Cyanidium" lineage is a distinct clade that diverged after the Galdieria clade but within a larger monophyletic clade that included the Cyanidium and Cyanidioschyzon lineages. Because the "cave Cyanidium" lineage is a mesophilic lineage that differs substantially from the other three thermoacidophilic lineages, we describe it as a new order (Cavernulicolales). Based on this evidence, we reclassified the Cyanidiophyceae into four orders: Cyanidiales, Cyanidioschyzonales, Cavernulicolales ord. nov., and Galdieriales ord. nov. The genetic distance among these four orders is comparable to, or greater than, the distances found between other red algal orders and subclasses. Three new genera (Cavernulicola, Gronococcus, Sciadococcus), five new species (Galdieria javensis, Galdieria phlegrea, Galdieria yellowstonensis, Gronococcus sybilensis, Sciadococcus taiwanensis), and a new nomenclatural combination (Cavernulicola chilensis) are proposed.

Removal and concurrent reduction of Cr(VI) by thermoacidophilic Cyanidiales: a novel extreme biomaterial enlightened for acidic and neutral conditions.

Thermoacidophilic Cyanidiales maintain a competitive edge in inhabiting extreme environments enriched with metals. Here, species of Cyanidioschyzon merolae (Cm), Cyanidium caldarium (Cc), and Galdieria partita (Gp) were exploited to remove hexavalent chromium [Cr(VI)]. Cm and Gp could remove 168.1 and 93.7 mg g-1 of Cr(VI) at pH 2.0 and 7.0, respectively, wherein 89% and 62% of sorbed Cr on Cm and Gp occurred as trivalent chromium [Cr(III)]. Apart from surface-sorbed Cr(VI), the in vitro Cr(III) bound with polysaccharide and in vivo chromium(III) hydroxide [Cr(OH)3] attested to the reduction capability of Cyanidiales. The distribution of Cr species varied as a function of sorbed Cr amount, yet a relatively consistent proportion of Cr(OH)3, irrespective of Cr sorption capacity, was found only on Cm and Cc at pH 2.0. In conjunction with TXM (transmission X-ray microscopy) images that showed less impaired cell integrity and possible intracellular Cr distribution on Cm and Cc at pH 2.0, the in vivo Cr(OH)3 might be the key to promoting the Cr sorption capacity (≥ 152 mg g-1). Cyanidiales are promising candidates for the green and sustainable remediation of Cr(VI) due to their great removal capacity, the spontaneous reduction under oxic conditions, and in vivo accumulation.

Novel field observations of coral reef fishes feeding on epiphytic and epizoic organisms associated with the allelopathic seaweed Galaxaura divaricata.

In degraded coral reef ecosystems, allelopathic macroalgae have received increasing attention from marine ecologists because their secondary metabolites (also known as allelochemicals) kill corals that grow adjacent to them and weaken the recovery of degraded reefs. One well-known coral-killing macroalga is the calcareous red seaweed Galaxaura. However, our knowledge of how coral reef fishes interact with allelopathic algae like Galaxaura is very limited. Here, we documented novel observations of feeding interactions of 17 species of coral reef fishes (herbivorous and carnivorous) with the filamentous Galaxaura divaricata on degraded lagoon patch reefs in Dongsha Atoll (South China Sea). Video analyses showed that territorial farming damselfishes (i.e., Dischistodus perspicillatus, D. prosopotaenia, Hemiglyphidodon plagiometopon, Pomacentrus grammorhynchus, P. adelus, and Neoglyphidodon nigroris) and juvenile parrotfishes (Scarus schlegeli, S. ghobban, S. rivulatus, and Chlorurus spilurus) likely used G. divaricata as a feeding substratum. Further, microscopic analyses revealed that the filamentous surface of G. divaricata harbored a wealth of epiphytic microalgae, such as filamentous cyanobacteria (i.e., Leptolyngbya, Lyngbya, Rivularia, Oscillatoria, and Stigonema), diatoms (i.e., Synedra, Nitzschia, Mastogloia, and Pleurosigma), and filamentous red algae (i.e., Heterosiphonia), suggesting that these fishes targeted the nutrient-rich microscopic epiphytes rather than the nutrient-poor host. Juvenile benthic carnivores (i.e., Labridae, Parupeneus multifasciatus, and Meiacanthus grammistes) form feeding assemblages with roving parrotfishes to feed on small invertebrates (i.e., amphipods, copepods, isopods, gastropods, and polychaetes) associated with G. divaricata. Given that coral reef fishes appear to target the epiphytes associated with Galaxaura rather than the alga itself, these observations thus substantiate the threat posed by the overgrowth of G. divaricata to coral recovery in degraded reef systems due to the lack of natural grazers.

A multifaceted ecological assessment reveals the invasion of the freshwater red macroalga Montagnia macrospora (Batrachospermales, Rhodophyta) in Taiwan.

Invasive freshwater macroalgae are rarely described. Montagnia macrospora is a freshwater red alga introduced from South America to East Asia via the global aquarium trade. The earliest occurrence record of this alga in Taiwan is dated 2005. To determine whether M. macrospora has become invasive in Taiwan and to understand the traits that facilitated its invasion, we took a multifaceted approach that combines examination of ecological background and population genetic analysis. Our island-wide survey showed that M. macrospora is widespread in the field across Taiwan, where the climate greatly differs from that of South America, and can self-sustain for nearly a decade. Our population genetic analysis revealed a lack of genetic diversity of M. macrospora in Taiwan, consistent with the hypothesis that the alga expanded through asexual reproduction. Moreover, during our long-term ecological assessments and field surveys, we observed that M. macrospora is an ecological generalist that can survive in a wide range of temperature, pH, illumination, and nutrient enrichment. Taken together, our data suggest that M. macrospora has successfully invaded the freshwater ecosystems of Taiwan, likely due to its ability to disperse asexually and to grow under broad environmental conditions. We hope that our study brings attention to invasive freshwater algae, which have been overlooked in conservation planning and management.

Geographic distance, sedimentation, and substrate shape cryptic crustose coralline algal assemblages in the world's largest subtropical intertidal algal reef.

Algal reefs, concreted by crustose coralline algae (CCA), are the main biotic reefs in temperate waters but rare in the subtropics and tropics. The world's largest known intertidal algal reef in the subtropics is the Taoyuan Algal Reef (TAR) located in the northwestern coast of Taiwan. The biodiversity and ecology of the TAR are scarcely explored, and now the reef is imperiled by industrialization. Here, we document cryptic species of CCA in Taiwan, particularly the TAR, by sequencing the psbA genes of over 1800 specimens collected across Taiwan. We also examine the ecological background of the TAR by surveying its benthic composition and measuring its environmental parameters. Our data reveal that the TAR harbours a high diversity of cryptic CCA species (27 molecular operational taxonomic units, or mOTUs), many of which are potentially new to science (18 mOTUs) and/or endemic to the TAR (9 mOTUs). Comparing the CCA species inventory of the TAR with the rest of Taiwan shows that the TAR represents a unique hotspot of CCA taxa in the waters of Taiwan. Our analyses show that variation in the CCA assemblages in the TAR is associated with geographic distance, sedimentation, and substrate type (for example, reef vs. hermit crab shell), suggesting that dispersal limitation and contemporary environmental selection shape the CCA assemblages in the TAR. The data from this study can inform the monitoring of human impacts on the health of the TAR and contribute to our understanding of the ecological processes underlying algal reef development.

Evaluating the tradeoffs of a generalist parasitoid fungus, Ophiocordyceps unilateralis, on different sympatric ant hosts.

It is essential for the survival and reproduction of parasitoids to adapt to the fluctuating host resources. Phenotypic plasticity may enable a parasitoid species to successfully achieve its control over a range of host species to maximize fitness in different hosts that may each require dissimilar, possibly conflicting, specific adaptations. However, there is limited information on how the fitness effects of host switching partition into costs due to the novelty of host species, where unfamiliarity with host physiological and morphological changes and its anti-parasite defenses reduces parasitoid growth, survivorship and/or reproductive success. In this study, the parasitoid fungus Ophiocordyceps unilateralis sensu lato was found to sympatrically infect a principal host ant species and other alternative sympatric hosts in the forest of central Taiwan. We herein report that the occurrence of ant infections by O. unilateralis s.l. shows spatial and temporal variation patterns on different host species. Results showed that the height from the ground to the leaf where the infected ants grip on, perithecia-forming ability, and growth rate of the stroma of the parasitoid fungus were dissimilar on different host species. These host range expansions not only related the fitness of O. unilateralis s.l. but also influenced the expression of extended phenotypic traits. Our findings revealed that a generalist parasitoid fungus suffered an evolutionary tradeoff between host breadth expansion and host-use efficiency.

Reappraising plastid markers of the red algae for phylogenetic community ecology in the genomic era.

Selection of appropriate genetic markers to quantify phylogenetic diversity is crucial for community ecology studies. Yet, systematic evaluation of marker genes for this purpose is scarcely done. Recently, the combined effort of phycologists has produced a rich plastid genome resource with taxonomic representation spanning all of the major lineages of the red algae (Rhodophyta). In this proof-of-concept study, we leveraged this resource by developing and applying a phylogenomic strategy to seek candidate plastid markers suitable for phylogenetic community analysis. We ranked the core genes of 107 published plastid genomes based on various sequence-derived properties and their tree distance to plastid genome phylogenies. The resulting ranking revealed that the most widely used marker, rbcL, is not necessarily the optimal marker, while other promising markers might have been overlooked. We designed and tested PCR primers for several candidate marker genes, and successfully amplified one of them, rpoC1, in a taxonomically broad set of red algal specimens. We suggest that our general marker identification methodology and the rpoC1 primers will be useful to the phycological community for investigating the biodiversity and community ecology of the red algae.

Comparative Genome Analysis Reveals Cyanidiococcus gen. nov., A New Extremophilic Red Algal Genus Sister to Cyanidioschyzon (Cyanidioschyzonaceae, Rhodophyta).

The taxonomic placement of strains belonging to the extremophilic red alga Galdieria maxima has been controversial due to the inconsistent phylogenetic position inferred from molecular phylogenetic analyses. Galdieria maxima nom. inval. was classified in this genus based on morphology and molecular data in the early work, but some subsequent molecular phylogenetic analyses have inferred strains of G. maxima to be closely related to the genus Cyanidioschyzon. To address this controversy, an isolated strain identified as G. maxima using the rbcL gene sequence as the genetic barcode was examined using a comprehensive analysis across morphological, physiological, and genomic traits. Herein are reported the chloroplast-, mitochondrion-, and chromosome-level nuclear genome assemblies. Comparative analysis of orthologous gene clusters and genome arrangements suggested that the genome structure of this strain was more similar to that of the generitype of Cyanidioschyzon, C. merolae than to the generitype of Galdieria, G. sulphuraria. While the ability to uptake various forms of organic carbon for growth is an important physiological trait of Galdieria, this strain was identified as an ecologically obligate photoautotroph (i.e., the inability to utilize the natural concentrations of organic carbons) and lacked various gene models predicted as sugar transporters. Based on the genomic, morphological, and physiological traits, we propose this strain to be a new genus and species, Cyanidiococcus yangmingshanensis. Re-evaluation of the 18S rRNA and rbcL gene sequences of the authentic strain of G. maxima, IPPAS-P507, with those of C. yangmingshanensis suggests that the rbcL sequences of "G. maxima" deposited in GenBank correspond to misidentified isolates.

Contrasting patterns of genetic structure and phylogeography in the marine agarophytes Gelidiophycus divaricatus and G. freshwateri (Gelidiales, Rhodophyta) from East Asia.

The evolutionary and population demographic history of marine red algae in East Asia is poorly understood. Here, we reconstructed the phylogeographies of two upper intertidal species endemic to East Asia, Gelidiophycus divaricatus and G. freshwateri. Phylogenetic and phylogeographic inferences of 393 mitochondrial cox1, 128 plastid rbcL, and 342 nuclear ITS2 sequences were complemented with ecological niche models. Gelidiophycus divaricatus, a southern species adapted to warm water, is characterized by a high genetic diversity and a strong geographical population structure, characteristic of stable population sizes and sudden reduction to recent expansion. In contrast, G. freshwateri, a northern species adapted to cold temperate conditions, is genetically relatively homogeneous with a shallow population structure resulting from steady population growth and recent equilibrium. The overlap zone of the two species roughly matches summer and winter isotherms, indicating that surface seawater temperature is a key feature influencing species range. Unidirectional genetic introgression was detected at two sites on Jeju Island where G. divaricatus was rare while G. freshwateri was common, suggesting the occurrence of asymmetric natural hybrids, a rarely reported event for rhodophytes. Our results illustrate that Quaternary climate oscillations have left strong imprints on the current day genetic structure and highlight the importance of seawater temperature and sea level change in driving speciation in upper intertidal seaweed species.

Filamentous calcareous alga provides substrate for coral-competitive macroalgae in the degraded lagoon of Dongsha Atoll, Taiwan.

BACKGROUND: The chemically-rich seaweed Galaxaura is not only highly competitive with corals, but also provides substrate for other macroalgae. Its ecology and associated epiphytes remain largely unexplored. To fill this knowledge gap, we undertook an ecological assessment to explore the spatial variation, temporal dynamics, and diversity of epiphytic macroalgae of Galaxaura divaricata on patch reefs in the lagoon of Dongsha Atoll, a shallow coral reef ecosystem in the northern South China Sea that has been repeatedly impacted by mass coral bleaching events. METHODS: Twelve spatially independent patch reefs in the Dongsha lagoon were first surveyed to assess benthic composition in April 2016, and then revisited to determine G. divaricata cover in September 2017, with one additional Galaxaura-dominated reef (site 9). Four surveys over a period of 17 months were then carried out on a degraded patch reef site to assess the temporal variation in G. divaricata cover. Epiphytic macroalgae associated with G. divaricata were quantified and identified through the aid of DNA barcoding at this degraded site. RESULTS: Patch reefs in the Dongsha lagoon were degraded, exhibiting relatively low coral cover (5-43%), but high proportions of macroalgae (13-58%) and other substrate (rubble and dead corals; 23-69%). The distribution of G. divaricata was heterogeneous across the lagoon, with highest abundance (16-41%) in the southeast area. Temporal surveys showed consistently high covers (mean ± SD = 16.9 ± 1.21%) of G. divaricata for 17 months. Additional photographic evidence suggested that overgrowth of G. divaricata can persist for 3.5 years. Yet, G. divaricata provides substrate to other macroalgae (e.g., Lobophora sp.) that also limit the growth of corals. CONCLUSIONS: Our study demonstrates that an allelopathic seaweed, such as G. divaricata, can overgrow degraded coral reefs for extended periods of time. By providing habitat for other harmful macroalgae, a prolonged Galaxaura overgrowth could further enhance the spread of macroalgae, and strengthen negative feedback loops, decreasing the recovery potential of degraded reefs.

Biosynthesis of Ascorbic Acid as a Glucose-Induced Photoprotective Process in the Extremophilic Red Alga Galdieria partita.

The extremophilic red alga Galdieria partita is a facultative heterotroph that occupies mostly low-light microhabitats. However, the exceptional detection of abundant populations of G. partita in sunlight-exposed soil raises the possibility that exogenous organic carbon sources protect cells from photo-oxidative damage. The present study aimed to identify the photoprotective process activated by exogenous glucose under photo-oxidative stress. We demonstrated that exogenous glucose mitigated the photo-oxidative damage of cells exposed to 300 µmol photons m-2 s-1 photosynthetic active radiation. Photosynthesis carbon assimilation scarcely contributed to the cell growth in the presence of glucose, but the photosynthetic apparatus was nevertheless maintained and protected by glucose in a concentration-dependent manner. Supplementation of glucose increased expression of the L-gulonolactone oxidase gene essential for ascorbic acid biosynthesis, whereas no enhanced expression of the genes involved in carotenoid or tocopherol biosynthesis was observed. Under the photo-oxidative stress condition, the ascorbic acid content was strongly enhanced by exogenous glucose. We propose that the biosynthesis of ascorbic acid is one of the major photoprotective processes induced by exogenous glucose. The elucidation of how ascorbic acid is involved in scavenging reactive oxygen species provides key insights into the photoprotective mechanism in red algae.

The influence of tourism-based provisioning on fish behavior and benthic composition.

Distribution of non-natural food (provisioning) to attract fish, though popular in coral reef tourism, has often been discouraged due to its assumed adverse effects on fish health and behavior. However, the effects of provisioning on community structure, anti-predator, and foraging behavior of teleost fishes, as well as their potential to indirectly affect benthic organisms, are not yet clear. Here, we compared fish composition, wariness, foraging behavior of herbivorous fishes, and the benthic cover between provisioned and control sites. We found significant differences in fish abundance, species number, and composition at some locations, but not all. Although most provisioned herbivorous fish did not reduce their biting rates of benthic algae, provisioned sites still had higher coverage of green macroalgae. Our results dispute widely held presumptions on the effects of tourism-based provisioning on the ecology and behavior of teleost fishes, as well as the benthic cover. These findings suggest that while regulation of provisioning is necessary to manage and mitigate any deleterious outcomes, when moderated and monitored, it could still provide an educationally beneficial tool for coral reef ecotourism.

The effects of contemporary selection and dispersal limitation on the community assembly of acidophilic microalgae.

Extremophilic microalgae are primary producers in acidic habitats, such as volcanic sites and acid mine drainages, and play a central role in biogeochemical cycles. Yet, basic knowledge about their species composition and community assembly is lacking. Here, we begin to fill this knowledge gap by performing the first large-scale survey of microalgal diversity in acidic geothermal sites across the West Pacific Island Chain. We collected 72 environmental samples in 12 geothermal sites, measured temperature and pH, and performed rbcL amplicon-based 454 pyrosequencing. Using these data, we estimated the diversity of microalgal species, and then examined the relative contribution of contemporary selection (i.e., local environmental variables) and dispersal limitation on the assembly of these communities. A species delimitation analysis uncovered seven major microalgae (four red, two green, and one diatom) and higher species diversity than previously appreciated. A distance-based redundancy analysis with variation partitioning revealed that dispersal limitation has a greater influence on the community assembly of microalgae than contemporary selection. Consistent with this finding, community similarity among the sampled sites decayed more quickly over geographical distance than differences in environmental factors. Our work paves the way for future studies to understand the ecology and biogeography of microalgae in extreme habitats.

Complete mitochondrial genomes of six species of the freshwater red algal order Batrachospermales (Rhodophyta).

Only two mitochondrial (mt) genomes had been reported in members of the red algal order Batrachospermales, which are confined to freshwater habitats. Additional mt genomes of six representative members (Batrachospermum macrosporum, Kumanoa ambigua, K. mahlacensis, Paralemanea sp., Sheathia arcuata, and Sirodotia delicatula) were sequenced aiming to gain insights on the evolution of their mt genomes from a comparative analysis with other red algal groups. Mt genomes sequenced had the following characteristics: lengths ranging between 24,864 nt and 29,785 nt, 22 to 26 protein-coding genes, G + C contents of 21.3 to 30.7%, number of tRNA of 16 to 37, non-coding DNA from 3.8% to 14.8%. Comparative analysis revealed that mt genomes in Batrachospermales are highly conserved in terms of genome size and gene content and synteny. Phylogenetic analyses based on COI nucleotide data revealed high bootstrap support only for the genera usually recovered in the phylogenetic analyses but no support for supra-generic groups. The insertion of a group II intron carrying an ORF coding for the corresponding intron maturase interrupting the COI gene was observed in Paralamenea sp. and accounted for its larger genome in comparison to the other Batrachospermales mt genomes.

Organization of plastid genomes in the freshwater red algal order Batrachospermales (Rhodophyta).

Little is known about genome organization in members of the order Batrachospermales, and the infra-ordinal relationship remains unresolved. Plastid (cp) genomes of seven members of the freshwater red algal order Batrachospermales were sequenced, with the following aims: (i) to describe the characteristics of cp genomes and compare these with other red algal groups; (ii) to infer the phylogenetic relationships among these members to better understand the infra-ordinal classification. Cp genomes of Batrachospermales are large, with several cases of gene loss, they are gene-dense (high gene content for the genome size and short intergenic regions) and have highly conserved gene order. Phylogenetic analyses based on concatenated nucleotide genome data roughly supports the current taxonomic system for the order. Comparative analyses confirm data for members of the class Florideophyceae that cp genomes in Batrachospermales is highly conserved, with little variation in gene composition. However, relevant new features were revealed in our study: genome sizes in members of Batrachospermales are close to the lowest values reported for Florideophyceae; differences in cp genome size within the order are large in comparison with other orders (Ceramiales, Gelidiales, Gracilariales, Hildenbrandiales, and Nemaliales); and members of Batrachospermales have the lowest number of protein-coding genes among the Florideophyceae. In terms of gene loss, apcF, which encodes the allophycocyanin beta subunit, is absent in all sequenced taxa of Batrachospermales. We reinforce that the interordinal relationships between the freshwater orders Batrachospermales and Thoreales within the Nemaliophycidae is not well resolved due to limited taxon sampling.

Concerted Divergence after Gene Duplication in Polycomb Repressive Complexes.

Duplicated genes are a major contributor to genome evolution and phenotypic novelty. There are multiple possible evolutionary fates of duplicated genes. Here, we provide an example of concerted divergence of simultaneously duplicated genes whose products function in the same complex. We studied POLYCOMB REPRESSIVE COMPLEX2 (PRC2) in Brassicaceae. The VERNALIZATION (VRN)-PRC2 complex contains VRN2 and SWINGER (SWN), and both genes were duplicated during a whole-genome duplication to generate FERTILIZATION INDEPENDENT SEED2 (FIS2) and MEDEA (MEA), which function in the Brassicaceae-specific FIS-PRC2 complex that regulates seed development. We examined the expression of FIS2, MEA, and their paralogs, compared their cytosine and histone methylation patterns, and analyzed the sequence evolution of the genes. We found that FIS2 and MEA have reproductive-specific expression patterns that are correlated and derived from the broadly expressed VRN2 and SWN in outgroup species. In vegetative tissues of Arabidopsis (Arabidopsis thaliana), repressive methylation marks are enriched in FIS2 and MEA, whereas active marks are associated with their paralogs. We detected comparable accelerated amino acid substitution rates in FIS2 and MEA but not in their paralogs. We also show divergence patterns of the PRC2-associated VERNALIZATION5/VIN3-LIKE2 that are similar to FIS2 and MEA These lines of evidence indicate that FIS2 and MEA have diverged in concert, resulting in functional divergence of the PRC2 complexes in Brassicaceae. This type of concerted divergence is a previously unreported fate of duplicated genes. In addition, the Brassicaceae-specific FIS-PRC2 complex modified the regulatory pathways in female gametophyte and seed development.

Phylogeographic surveys and apomictic genetic connectivity in the North Atlantic red seaweed Mastocarpus stellatus.

The North Atlantic red alga Mastocarpus stellatus is characterized by two life histories (sexual-type and direct-type), which correspond to two geographically isolated breeding groups. These features enable M. stellatus to be an interesting model to investigate how environmental shift and apomictic propagation have influenced its population genetic structure, historical demography and distribution dynamic. To test these ideas, we obtained 456 specimens from 15 locations on both sides of the North Atlantic and sequenced portion of the nuclear internal transcribed spacer (ITS), mitochondrial cox2-3 region (COX) and plastid RuBisCo spacer (RLS). Median-joining networks and ML trees inferred from COX and RLS consistently revealed two gene lineages (mtDNA: CN, CS; cpDNA: RN, RS). The concatenated COX and RLS markers yielded three cytotypes: a northern CN-RN, a southern CS-RS and a mixed cytotype CS-RN, which enabled us to roughly separate samples into D (direct-type life-cycle) and S (sexual-type life-cycle) groups (northern CN-RN and mixed cytotype CS-RN=D; southern CS-RS=S). Pairwise FST analysis of the D group revealed a high level of genetic differentiation both along European coasts and across the Atlantic basin. Bayesian skyline plots (BSPs) and IMa analyses indicated that M. stellatus underwent slight demographic expansion at the late-Pleistocene, with the beginning of divergence between lineages dating to c. 0.189Ma (95%HPD: 0.083-0.385Ma). IMa analyses also revealed asymmetric genetic exchange among European populations and a predominant postglacial trans-Atlantic migration from Norway and Galway Bay to North America. Our study highlights the importance of phylogeographic approaches to discover the imprints of climate change, life histories and gene flow in driving population genetic connectivity and biogeographic distribution of intertidal seaweeds in the North Atlantic.