The red alga Asparagopsis taxiformis comprises six cryptic mitochondrial lineages (L1–L6) that are genetically distinct but morphologically cryptic. While these lineages occur across tropical and subtropical oceans worldwide, the phylogeographic structure of Korean populations has remained poorly understood. In this study, 39 individuals were collected from eight coastal localities in Korea and analyzed together with global samples. Phylogenetic analyses of concatenated cox2-3 spacer and partial cox1 sequences revealed that all Korean A. taxiformis samples belong to Lineage-2 (L2), a cosmopolitan lineage distributed across multiple ocean basins. Within Lineage-2, the cox2-3 spacer network displayed a consistent global pattern where the H1 haplotype predominated worldwide, including in Korea, whereas private haplotypes were primarily concentrated in Mediterranean populations, especially in Italy. The cox1 network provided higher resolution, resolving three dominant haplotypes (H1-H3) with distinct geographic ranges in East Asia: H1 mainly in the East Sea and Jeju Island; H2 mainly along the south and west coasts of Korea and Jeju Island; and H3 largely in the Ryukyu Islands. Despite this geographic structure, only one to three mutational steps separate these three haplotypes. Organelle genomic analysis of Korean, Italian, and American individuals further demonstrates that Lineage-2 exhibits low to intermediate divergences, a conserved trait despite its broad global distribution compared to other Rhodymeniophycidae species. Demographic and palaeoceanographic reconstructions suggest that the eastern Italian and southern Korean Peninsula served as refugial areas for Lineage-2, although the mechanisms maintaining this conserved, widespread structure remain unknown.

Exploration of life in extreme environments allows the discovery of organisms with extraordinary biotechnological potential. Hot springs, characterised by extreme conditions (pH < 2; Temperature > 40 degrees C; high concentrations of metals), present a formidable challenge for most life forms, yet support a few specialised archaea, bacteria, and eukaryotes, including the unicellular red alga Galdieria. Beyond their ecological relevance, members of the order Galdieriales emerged as promising platforms for biotechnological applications, including metal and rare earth elements (REEs) recovery, bioremediation under extreme conditions, and sustainable biomass production. Previous phylogenetic studies based on plastid and nuclear genes suggested a complex genetic structure within Galdieria, with evidence for multiple diverging clades within the species G. sulphuraria. To resolve these enigmatic relationships, we employed a genome-scale approach, integrating plastid, mitochondrial, and nuclear data from Galdieria strains collected across diverse geographical locations. Our comprehensive phylogenetic analysis revealed divergence of mitochondrial, plastid, and nuclear genomes into at least seven distinct and well-supported clades; alongside i) independent evolutionary trajectories of these genomic lineages, and ii) incongruent interspecific relationships among the three genomes, reflecting complex evolutionary dynamics. Synonymous and non-synonymous substitution analyses further underscored differential evolutionary pressures on the three cellular genomes acting across the Galdieria species, with strong purifying selection in plastid genes and more relaxed selection in mitochondrial and nuclear genomes. In this context, the genetic and evolutionary framework presented here provides a crucial basis for understanding the molecular determinants underlying the exceptional metabolic versatility and stress tolerance that make Galdieriales attractive candidates for applied biotechnology.

The order Ishigeales is an early-diverging lineage of brown algae that exhibits remarkable morphological diversity ranging from simple filamentous forms to robust terete and foliose thalli, which may reflect complex evolutionary histories in their organelle genomes. However, limited organellar genomic data for the Ishigeales, presently confined to Ishige okamurae, have hindered a comprehensive understanding of organelle genome dynamics in that order. To address this, we de novo assembled the complete plastomes and mitogenomes of the four Ishigeales species: Pilinia rimosa, Petroderma maculiforme, I. okamurae, and I. foliacea. Phylogenetic analyses based on plastome and mitogenome datasets using both concatenation and coalescent-based approaches supported the divergence order from the less complex filamentous morphology of Pi. rimosa to the more morphologically complex terete and foliose Ishige species, yet it also revealed a phylogenetic conflict at the branching point of Pe. maculiforme, suggesting an intricate evolutionary history in filamentous species. Both plastomes and mitogenomes exhibited extensive genomic rearrangement. Notably, mitochondrial genome expansion was observed in Pi. rimosa and Pe. maculiforme, and massive cox2 gene duplication was found in Pi. rimosa. These expansions were likely driven by the proliferation of tandem repeats, and most copies are likely under relaxed purifying selection. Analysis of synonymous substitution rate to the nonsynonymous substitution rate revealed the rapid divergence in several plastid genes between I. okamurae and I. foliacea. In particular, the DPOR gene family exhibited a signal of relaxed purifying selection, potentially linked to their distinct morphological changes in Ishige species. Overall, our findings reveal that Ishigeales underwent lineage-specific differentiation and dynamic evolutionary patterns in the organelle genomes along with morphological variation.

Abstract The evolution of multicellularity has long been linked to reproductive strategies. A long-standing debate concerns whether multicellular organisms are primarily stabilized by small single-cell propagules that minimize genetic heterogeneity or by larger multicellular and multinucleate propagules that may improve developmental success and survival of individuals. the Xanthophyceae provides an excellent model for investigating these questions, exhibiting transitions between unicellular to multicellular filamentous and coenocytic forms together with diverse reproductive modes, including single-cell zoospores and autospores, and multinucleate monospores and akinetes. However, a robust phylogenetic framework and systematic analyses of character evolution have remained lacking in this lineage. Here, we present a phylogenomic framework based on a nuclear dataset of 680 genes from 18 species, including 17 newly generated transcriptomes. Nuclear phylogenies robustly resolve all sampled inter-ordinal and inter-familial relationships with full concordance between concatenation and coalescent analyses, while plastid (141 genes) and mitochondrial (31 genes) datasets from 33 species recover identical topologies. Based on these results, we establish one new order (Pseudopleurochloridales), emend one order (Heterococcales), and propose five new families. Ancestral character reconstruction indicates at least four independent transitions from unicellular ancestors to simple multicellularity. Bayesian analyses of multicellularity and reproductive characters show that these transitions were consistently accompanied by shifts from multiple autospore-type propagules toward single monospore- and akinete-type propagules, whereas reversions to unicellularity were associated with the reappearance of autospore-based reproduction. These results provide a phylogenomic framework for understanding multicellular evolution in Xanthophyceae and shed light on the relationship between reproductive modes and the emergence of simple multicellularity.

The unicellular red algae, Cyanidiophyceae, that diverged early during Archaeplastida (algal and plant) evolution, occupy a variety of extreme habitats that are inhospitable for most other eukaryotes. With the use of modern genomics and genetics methods, Cyanidiophyceae show a remarkable taxonomic diversity, share haplodiplophasic life cycles, and are engaged in complex trophic interactions with microbes that occupy geothermal niches. Amenable to molecular engineering, Cyanidiophyceae are excellent models for understanding evolutionary mechanisms that underpin their extremophilic lifestyles. Their unique growth conditions make these choice red algae of high interest for biotechnological exploitation in environments unsuitable for crops.

Saccharina japonica is a commercially and ecologically important seaweed and shows rapid ecological speciation. This kelp represents an excellent model for understanding the process and genetic mechanism of diversification and speciation of brown seaweeds. Up to now, there was limited research about the ecological speciation of seaweeds from genomic perspective. In this study, we conducted genome resequencing of four varieties of S. japonica and two sister species (S. angustata and S. longissima) to investigate the genetic mechanism of ecological speciation. The demographic history suggests that the lineage of S. angustata and S. longissima is a sister lineage to that of S. japonica, rather than its direct ancestor, with the genetic lineage of S. japonica var. religiosa diverging earliest from the rest of the taxa. Even though there is lower genetic differentiation among these varieties, natural hybridization and gene flow are limited. We detected some heat resistance (e.g. heat shock protein 70), stress response (e.g. ubiquitin-like protein) and growth-related genes (e.g. immediate upright (imm) upregulated 3) were under positive selection during the ecological speciation. Low linkage disequilibrium decay rate and extensive signals of selective sweeps were detected in these varieties, suggesting that adaptive differentiation under natural selection was the driving force for ecological speciation. The main force driving speciation in this species does not appear to be ongoing hybridization but historical admixture and adaptive differentiation due to natural selection. An understanding of the evolutionary history and ecological speciation of S. japonica represents an important prerequisite for effective use of germplasm in breeding and for the conservation of natural resources.

Primary endosymbiosis, resulting in the establishment of a photosynthetic organelle in eukaryotes, has occurred twice: more than 1.5 billion years ago in the common ancestor of Archaeplastida and 90 to 140 million years ago in the Paulinella lineage. The relatively recent, independent event in phototrophic Paulinella provides opportunities for investigating the earlier stages of primary plastid establishment. In Archaeplastida, plastid origin is so ancient that little can be inferred about the eukaryotic host in which endosymbiosis occurred. By contrast, the genus Paulinella includes nine heterotrophic species that are closely related to the phototrophic lineage, which can be used to understand the impact of primary endosymbiosis on "host" cell evolution. Research on heterotrophic Paulinella is very limited, and analyses addressing the evolutionary impacts of primary endosymbiosis on the ancestral heterotrophic lineage have been lacking. In this study, we used long-read sequencing to determine the mitochondrial genome sequence from three taxa of heterotrophic Paulinella that formed a "bloom" in their native environment. We also determined the mitogenomes of two phototrophic Paulinella species. Along with two available mitogenomes, we conducted a comparative analysis of phototrophic and heterotrophic Paulinella mitogenomes and find that gene order rearrangements occurred more frequently in the phototrophic lineage. We detected signatures of relaxed selection in mitochondrial DNA from the phototrophic Paulinella clade. This pattern likely reflects reduced effective population size (Ne) associated with plastid primary endosymbiosis, in line with the endosymbiotic ratchet hypothesis. We propose that the reduced Ne in phototrophic Paulinella strongly impacted mitogenome evolution in these species.

Summary Iron mineralization has profoundly influenced Earth’s biogeochemical history 1,2 , yet the specific mechanisms underlying banded iron formation (BIF) remain unresolved 3–5 . Here we profile the microbiomes of Holocene sediments beneath the Larsen C Ice Shelf (LCIS), Antarctica 6–8 , through stratigraphic analysis of sedimentary ancient DNA combined with metagenomics. Distinct microbial phases aligned with glacial facies boundaries, with sub-ice shelf communities dominated by chemolithoautotrophs including an uncultured Thermodesulfovibrionia . This bacterium, visualized by fluorescence in situ hybridization and designated ‘ Candidatus Mariimomonas ferrooxydans’ (phylum Nitrospirota ), emerged as a keystone taxon with high network centrality. Its genome encodes Cyc2, a fused porin–cytochrome outer membrane protein implicated in Fe(II) oxidation. Heterologous expression of Cyc2 in Escherichia coli confirmed its ability to catalyze iron oxidation, supporting iron precipitation under dark, anoxic conditions. These pristine LCIS sediments, unaltered since the last glacial maximum, provide a modern analogue for synglacial BIFs deposited during Neoproterozoic Snowball Earth events. Our findings deliver direct genomic and functional evidence for chemolithotrophic iron oxidation, challenge phototroph-centric models of BIF genesis, and highlight microbial iron cycling as a recurring force in Earth’s geochemical evolution. Beyond Earth, these insights inform interpretations of iron deposits on other planetary bodies.

The genus Paulinella represents a rare, independent case of primary endosymbiosis, providing a unique system to study the early stages of organelle evolution. Here, we expand current understanding of primary plastid endosymbiosis through the discovery and characterization of two novel photosynthetic amoebae, Paulinella marae sp. nov. and Paulinella murrayi sp. nov., isolated from a brackish water habitat in North Carolina, United States. Complete chromatophore genomes and mitochondrial data revealed conserved gene content but notable structural variation, including genome rearrangements and inversion events. Phylogenetic analyses uncovered significant discordance between nuclear and organelle datasets, likely driven by substitution saturation, limited taxon sampling, and differing evolutionary signals across loci. Ecological observations over multiple years indicate that both species are in low abundance but consistently present, and when coupled with hobbyist data, support the hypothesis that photosynthetic Paulinella species are globally distributed yet under-sampled. These results increase known species diversity within the clade from four to six and highlight the importance of integrating field-based observations with genomic approaches. Overall, this work advances Paulinella as a model for studying ongoing primary endosymbiosis, lineage divergence, and the ecological strategies of low-abundance microbial eukaryotes.

Abstract The red seaweed genus Ahnfeltia is an ancient lineage that has persisted for over 500 million years with remarkably limited diversification despite a global distribution in cold-temperate intertidal habitats. Compared to the highly diverse sister lineage, Rhodymeniophycidae, Ahnfeltia provides a unique system for investigating long-term evolutionary persistence in marine macroalgae. Here, we generated chromosome-scale genomes from five populations across three species and combined population genomics with paleogeographic niche modelling. Our results reveal remarkable genomic conservation, strong geographic isolation with limited gene flow, high homozygosity, and evidence of purifying selection. Niche projections indicate long-term stability and spatial connectivity of suitable cold-temperate habitats. These findings suggest that Ahnfeltia ’s persistence and limited diversification are linked to genomic constraints and stable ecological niches over geological timescales. This study provides new insights into the genomic basis of evolutionary stasis in ancient marine lineages and highlights potential vulnerability to ongoing climate change affecting cold-water coastal ecosystems.

Plants are confronted with a myriad of challenges that they must mitigate without being able to move to new environments. Systemic acquired resistance (SAR) is a secondary immune response in plants to pathogen infection. Little is known about SAR in red algae. The red alga Dasysiphonia japonica can survive for more than six years without lethal damage when infected with the pathogen Olpidiopsis heterosiphoniae but dies within a month when infected with another pathogen, Olpidiopsis dasysiphoniae. During this process, D. japonica infected with O. heterosiphoniae accumulated reactive oxygen species (ROS, DCFH-DA staining) in the host cell membrane, whereas this was not observed with O. dasysiphoniae infection. Transcriptome analysis identified four NADPH-oxidase genes (Djrboh) that generate ROS in the cell membrane of D. japonica. Real-time PCR of four NADPH oxidase genes (Djrboh) showed that two were upregulated (Djrboh1, 2) at the onset of Olpidiopsis heterosiphoniae infection, but not during infection with O. dasysiphoniae in early infection that does not induce SAR. The successful SAR of D. japonica was mediated by low concentrations of ROS signaling and regulation of NADPH-oxidase genes.

Olpidiopsis is one of the major pathogens of commercially valuable red algae. Despite being such an important pathogen, the infection process has been studied and documented mainly in the Olpidiopsis species infecting Pyropia spp., but less so in other red algae. We collected an Olpidiopsis -like pathogen infecting Dasysiphonia japonica in Japan, and it is different from any previous described Olpidiopsis species. Molecular phylogeny inferred from 18S rRNA gene, cox 1, and cox 2 sequences and species delimitation analyses (i.e., Assemble Species by Automatic Partitioning, single-threshold General Mixed Yule Coalescent, Bayesian Poisson Tree Processes) showed that the collected Olpidiopsis sp. formed a supported clade with other Rhodophyta-infecting Olpidiopsis species, and was considered a distinct species by all species-delimitation methods. Furthermore, we tested the host range of this pathogen and studied its morphology through microscopic analysis. This new pathogen was distinguished from other species by the presence of only one sporangium per host cell and, during development, the sporangium became irregularly shaped, almost filling the entire host cell and produced up to 14 discharge tubes to release zoospores. The host range tests found that it only infected species of Dasysiphonia spp. Therefore, we propose the name, Olpidiopsis dasysiphoniae sp. nov., for this new species.

The recognition of new species is fundamental to understanding biodiversity and biogeography. Through an integrative taxonomic approach, combining morphological and multilocus molecular data, a new red algal species, Rhodogorgon truncata sp. nov. is described from the west coast of Thailand. The new species differs from the two previously described species in the genus, i.e. R. ramosissima and R. flagellifera, by unique morphological features including compressed and stocky thalli with determinate short branchlets along the upper portion, giving it a truncated and verrucose appearance. Our morphological observations revealed the smaller size, compressed shape, and truncated apices of R. truncata in contrast to the cylindrical, irregularly branched thalli of R. ramosissima and R. flagellifera. Phylogenetic analyses suggest that R. truncata is more closely related to undescribed Rhodogorgon specimens from the Indo-Pacific than to the Caribbean R. ramosissima. The discovery of this new species expands the known diversity within the genus Rhodogorgon and suggests greater, as yet undiscovered, species diversity among poorly studied red algae in the Indo-Pacific Ocean.

Asparagopsis taxiformis has high potential to mitigate enteric methane emissions from livestock due to its high bromoform content. However, the current supply of gametophytes is limited due to insufficient biomass. Year‐round production of gametophyte can address insufficient biomass through a continuous supply of initial biomass from tetrasporophyte. In this study, we evaluated the effects of temperature (10, 20, and 30°C) and photoperiod (8:16, 12:12, and 16:8 h light:dark cycles), as well as irradiance (10, 20, 40, 80, 160 μmol photons · m −2 · s −1 ) and nutrient conditions (high nutrient: 500 μM nitrate and 30 μM phosphate; low nutrient: 50 μM nitrate and 3 μM phosphate) on the growth and reproduction of tetrasporophyte of A. taxiformis. Temperature was a key factor in both growth and reproduction, whereas photoperiod was a key factor in reproduction. Growth of tetrasporophyte was inhibited by 10°C regardless of photoperiod. The development of tetrasporangia was only observed at 20°C with an 8:16 h light:dark cycle. At 20°C with an 8:16 cycle, irradiance affected the development of tetrasporangia at high nutrient concentration. The development of tetrasporangia was observed at 20 and 40 μmol photons · m −2 · s −1 , with the highest growth rate observed at 160 μmol photons · m −2 · s −1 without the development of tetrasporangia. These results suggest that controlling irradiance at 20°C on an 8:16 h light:dark cycle under high nutrient concentration can regulate the growth and reproduction of A. taxiformis tetrasporophyte during cultivation.

Sargassum fusiforme and Sargassum thunbergii are ecologically and commercially important seaweeds that thrive in intertidal zones and are frequently exposed to extreme variation in environmental stress. Despite their importance, limited genomic information exists for these species, which hinders a comprehensive understanding of the evolution and adaptation of the genus Sargassum to marine coastal habitats. Two Sargassum genomes were generated in this study. The genome sizes of S. fusiforme and S. thunbergii were 438 and 376 Mbp, respectively, which are larger than the published genomes of the brown seaweed group, Ectocarpales. Expansion of the Sargassum genomes was significantly explained by the spread of transposable elements (TEs). Additionally, extensive gene duplications and their diversification occurred particularly through tandem, proximal, and dispersed duplications, which likely played an important role in response to environmental stress. Differentially expressed gene analysis under ambient and desiccation stress conditions confirmed that some duplicated genes respond to stress. We identified enhanced disease susceptibility 1 (EDS1) genes that promote salicylic acid (SA) biosynthesis, and their expansion is likely linked to TEs. We also confirmed the potential role of EDS1 by analyzing its subcellular localization (in Arabidopsis thaliana) and quantified the increased SA levels under desiccation conditions. This study demonstrates that the genomic evolution has played a critical role in allowing S. fusiforme and S. thunbergii to adapt to harsh intertidal conditions. The genomic resources of Sargassum species provided here will be instrumental in advancing future research, aiding in the understanding of adaptive evolution in brown algae.

Bifacial cambium, which produces xylem and phloem, and monopodial architecture, characterized by apical dominance and lateral branching from axillary buds, are key developmental features of seed plants, consisting of angiosperms and gymnosperms. These allow seed plants to adapt to diverse environments by optimizing resource allocation and structural integrity. In seed plants, SUPPRESSOR OF MAX2-LIKE (SMXL) family members function in phloem development and strigolactone-induced inhibition of axillary bud outgrowth. Although strigolactone signaling regulates most SMXL family members, the only known regulator of SMXL4 and SMXL5 is the RNA-binding protein JULGI. We demonstrate that in angiosperms, by directly regulating SMXL4/5 expression, JULGI uncouples SMXL4/5 activity from strigolactone signaling. JULGI and ancestral SMXLs from seedless vascular plants or SMXL4/5 from seed plants are coexpressed in the phloem tissues of vascular plants, from lycophytes to angiosperms. Core angiosperm SMXL4/5 mRNAs contain a G-rich element in the 5' untranslated region (UTR) that serves as a target sequence for JULGI to negatively regulate SMXL4/5 expression. Heterologous expression of JULGIs from various angiosperms rescued the Arabidopsis jul1 jul2 mutant. Expressing SMXL4/5s from seed plants and ancestral SMXLs rescued Arabidopsis smxl4 smxl5. Angiosperm SMXL4/5s lack an RGKT motif for proteasomal degradation. Indeed, treatment with the synthetic strigolactone analog rac-GR24 induced proteasomal degradation of SMXL from ferns and SMXL5a from gymnosperms, but not SMXL4/5 from angiosperms. These findings suggest that in ancestral angiosperms, the 5' UTR of SMXL4/5 gained G-rich elements, creating a regulatory module with JULGI that allows the phloem development pathway to act independently of strigolactone signaling.

Dictyotopsis propagulifera is an enigmatic species of brown algae, first described by Wilhelm Troll in 1931. In the present study, we successfully established a unialgal culture strain of D. propagulifera collected from mangrove sediments in Perak, Malaysia, directly across the Malacca Strait from the type locality in Indonesia. Our observations over 10 years of culture confirm vegetative propagule formation as originally described by Troll. However, sexual reproduction typical of Dictyotales (e.g. flagellate sperm and non-flagellate eggs) was not confirmed. Detailed ultrastructural analysis revealed that D. propagulifera shares many cellular characteristics with Dictyota, including chloroplasts with the three-layered thylakoid lamellae, well-developed plasmodesmata forming pit fields and the presence of a single apical meristematic cell. In addition, biochemical analysis confirmed the presence of the polyol mannitol as main photosynthetic product and organic osmolyte, which is characteristic for brown algae. Phylogenomic analysis using both plastid and mitochondrial genomes, along with multigene analysis, further supports the monophyly of D. propagulifera and Dictyota, following the earlier divergence of Dictyopteris. These findings not only provide an updated description of D. propagulifera based on modern microscopy techniques, but also propose potential evolutionary pathways responsible for its developmental patterns and asexual reproductive strategy.

We isolated into culture, along with their host cells (Desmidium, Spondylosum and Geminella), Stylococcus aureus and a previously undescribed taxon we name Stylococcus brevis sp. nov. The loricas of S. aureus were attached to the host cell with a long, thin stalk, and cells produced a long pseudopodium that often branched at the tip. Cells divided transversely, followed by daughter cell rotation of 90 degrees, giving the appearance of longitudinal division. One daughter cell was flagellate, escaped the mother lorica and swam to a host cell where it attached, producing a new lorica. The swimming cell had a long immature flagellum and short mature flagellum that were easily observed using a light microscope. Conversely, the loricas of S. brevis were anchored to the host cell with a short, thick stalk, and the cells produced a short pseudopodium. Stylococcus brevis cell division was the same as for S. aureus; however, its swimming cells had an extremely short mature flagellum that was visible by light microscopy only after chemical fixation. Molecular phylogenetic analyses using the 18S rRNA and rbcL genes showed that the two Stylococcus species were distinct species. Surprisingly, Stylococcus branched with Lagynion scherffelii (generitype) and L. ampullaceum, suggesting that these algae are congeneric, which supports the idea that the typical Stylococcus lorica morphology and stalk formation are a response to the host cell. We found no evidence suggesting that Stylococcus and Stylochrysalis are synonyms.

The conserved eukaryotic functions of cell cycle genes have primarily been studied using animal/plant models and unicellular algae. Cell cycle progression and its regulatory components in red (Rhodophyta) seaweeds are poorly understood. We analyzed diurnal gene expression data to investigate the cell cycle in the red seaweed Gracilariopsis chorda. We identified cell cycle progression and transitions in G. chorda which are induced by interactions of key regulators such as E2F/DP, RBR, cyclin-dependent kinases, and cyclins from dusk to dawn. However, several typical CDK inhibitor proteins are absent in red seaweeds. Interestingly, the G1-S transition in G. chorda is controlled by delayed transcription of GINS subunit 3. We propose that the delayed S phase entry in this seaweed may have evolved to minimize DNA damage (e.g., due to UV radiation) during replication. Our results provide important insights into cell cycle-associated physiology and its molecular mechanisms in red seaweeds.

A reclassification of Cyanidium chilense under the new genus Cavernulicola was recently proposed together with a new family (Cavernulicolaceae) and a new order (Cavernulicolales). Unfortunately, due to an error in the required citation of the basionym, the name "Cavernulicola chilensis" was invalid and cannot be accepted as the generitype of Cavernulicola. This means that Cavernulicola, Cavernulicolaceae, and Cavernulicolales are likewise invalid names under the provisions of the International Code of Nomenclature for algae, fungi, and plants (ICN, Shenzhen Code). In this contribution, each of these names is validated.