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The maximum likelihood phylogeny of Calliblepharis saidana comb. nov. derived from combined alignment (rbcL and SSU rDNA). Support values shown on branches are maximum likelihood bootstrap (MLBt) and Bayesian posterior probability from DNA data. MLBt values of <50% are indicated by hyphens (-). Asterisk means the type species of Hypnea and Calliblepharis. To right of tree, diagrams of cross section of the vegetative thallus (1-6) and female cystocarp structures (7-9) for the family Cystocloniaceae. 1. Axial cell with medullary cells in the genus Hypnea. 2. Axial cell surrounded by several periaxial cells and medullary cells in the genus Calliblepharis. 3. Axial cell surrounded by periaxial cells and medullary cells with some rhizoidal filaments in the genus Craspedocarpus. 4. Axial cell with medullary cell in the genus Rhodophyllis. 5. Inconspicuous axial cell and medulla of elongated cells with rhizoids in the genus Stictosporum. 6. Axial cell and medulla of elongated cells with rhizoids in the genus Austroclonium. 7. Single carposporangia and numerous filaments connecting the gonimoblast and the pericarp in the genus Hypnea. 8. Chains of carposporangia and reticulum of gonimoblast filaments in the Calliblepharis species including type species and C. saidana. 9. Chains of carposporangia and parallel columns of gonimoblast filaments in C. fimbriata and C. celatospora.
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The genera Hypnea and Calliblepharis of the family Cystocloniaceae are discriminated by their female reproductive structure, especially in the formation of carposporangia and gonimoblasts. Hypnea saidana, once classified based on obsolete evidence, has not been studied phylogenetically using molecular analysis and detailed reproductive structure th...
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... ML and BI using the combined alignment of SSU and rbcL sequences yielded almost concordant tree, with little difference at node (Fig. 7). The combined tree re- solved monophyletic node of the genus Calliblepharis in- cluding type species, C. ciliata, even though the node was supported by low bootstrap values. C. saidana formed weak sister relationships with C. fimbriata and C. celatos- ...
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... Calliblepharis but it is absent in other genera of Cystocloniaceae as well as most species of Hypnea (Table 1) (Kylin 1930, Tanaka 1941, Díaz-Tapia et al. 2013). Other genera such as Craspedocarpus, Rhodophyllis, Cystoclo- nium, and Stictosporum have an axial cell surrounding by medullary cell and frequently have rhizoidal filaments in medulla (Fig. 7) (Min-Thein and Womersley 1976). The rhizoidal filaments are absent in both genera Callibleph- aris and Hypnea. The genus Craspedocarpus is similar with Calliblepharis in having several periaxial filaments surrounding axial cell, but Craspedocarpus differs by its rhizoids in the medulla and well developed rosette in cor- ...
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... from Hypnea by its female structures (Fig. 7) ( Kylin 1928Kylin , 1930 and by its molecular phylogeny (DíazTapia et al. 2013, D'Archino et al. 2015). The central clus- ters of cystocarps in Calliblepharis congeners either form a reticulum or arrange themselves into parallel columns (Chiovitti et al. 1998, Díaz-Tapia et al. 2013). C. saidana and the type species C. ciliata ...
Citations
... Even with the recent investments in molecular data and greater sampling intensity (Geraldino et al., 2009(Geraldino et al., , 2010Jesus et al., 2016Jesus et al., , 2019aNauer et al., 2014Nauer et al., , 2015Nauer et al., , 2016Nauer et al., , 2019Kang and Nam, 2018;Cabrera et al., 2020;Huisman et al., 2021;Kundu and Bast, 2021), Hypnea remains a source of several taxonomic problems, with challenging species delimitation (Price et al., 1992;Jesus et al., 2016Jesus et al., , 2019a. This has been pronounced not only in the most species-rich genus of Cystocloniaceae, but also in the family as a whole, as molecular efforts have been limited to a few genes so far (Hommersand and Fredericq, 2003;Won et al., 2016;Yang and Kim, 2017;Hoffman et al., 2018;Soares and Fujii, 2020;Tonicelli et al., 2021). Despite its economic and ecological importance, and taxonomic uncertainties, only two species of Cystocloniaceae are currently represented in genomic datasets to date. ...
Cystocloniacae is a highly diverse family of Rhodophyta, including species of ecological and economic importance, whose phylogeny remains largely unresolved. Species delimitation is unclear, particularly in the most speciose genus, Hypnea, and cryptic diversity has been revealed by recent molecular assessments, especially in the tropics. Here, we carried out the first phylogenomic investigation of Cystocloniaceae, focused on the genus Hypnea, inferred from chloroplast and mitochondrial genomes including taxa sampled from new and historical collections. In this work, molecular synapomorphies (gene losses, InDels and gene inversions) were identified to better characterize clades in our congruent organellar phylogenies. We also present taxon-rich phylogenies based on plastid and mitochondrial markers. Molecular and morphological comparisons of historic collections with contemporary specimens revealed the need for taxonomic updates in Hypnea, the synonymization of H. marchantae to a later heterotypic synonym of H. cervicornis and the description of three new species: H. davisiana sp. nov., H. djamilae sp. nov. and H. evaristoae sp. nov.
... The family Cystocloniaceae is one the most diverse in the Gigartinales, with 15 genera and 113 species currently recognized (Guiry & Guiry 2022), and is considered economically important as a source of carrageenan (Chiovitti et al. 1998). Morphologically, the family is characterized by uniaxial, terete or flat thalli, with pseudoparenchymatous or filamentous medulla, procarps with 3-or 4-celled carpogonial branches, and zonately divided tetrasporangia (Yang & Kim 2017, Huisman 2018, Guiry & Guiry 2022. The use of molecular markers to investigate phylogenetic relationships in the red algae has become commonplace, improving the accuracy of taxonomic studies, especially in cases where specimens lack reproductive structures for full morphological identification (Saunders 2005, Sherwood et al. 2010, Saunders & Moore 2013. ...
... The use of molecular markers to investigate phylogenetic relationships in the red algae has become commonplace, improving the accuracy of taxonomic studies, especially in cases where specimens lack reproductive structures for full morphological identification (Saunders 2005, Sherwood et al. 2010, Saunders & Moore 2013. Since the emergence of molecular techniques, a number of studies have been conducted on the order Gigartinales, including the genera Calliblepharis Kützing and Hypnea J.V. Lamouroux (Saunders & Kraft 1994, Geraldino et al. 2010, Díaz-Tapia et al. 2013, Yang & Kim 2017, Soares et al. 2019, Soares & Fujii 2020, Campbell et al. 2022. ...
... Mostly found in subtidal zones, Calliblepharis CALLIBLEPHARIS yASUtAkEI AND HyPNEA tSUDAE Phytotaxa 572 (1) © 2022 Magnolia Press • 75 species have a wide distribution encompassing the north-eastern Atlantic from the British Isles to Mauritania and the Mediterranean Sea, South Africa, Brazil, Japan, Korea, New Zealand, Australia and USA (Guiry & Guiry 2022). Over the past decade, several taxonomic studies using morphological and molecular data have investigated taxonomic and phylogenetic relationships within Calliblepharis; however, the genus remains paraphyletic (Díaz-Tapia et al. 2013, Yang & Kim 2017, Soares et al. 2019, Soares & Fujii 2020. ...
Surveys of Hawaiian macroalgae over the past 15 years have yielded numerous specimens representing species new to science. Calliblepharis yasutakei sp. nov. is here described based on a plant collected at a depth of 98 m from Kapou, Pa-pahânaumokuâkea Marine National Monument, Hawaiʻi. Phylogenetic analyses of three molecular markers (COI, rbcL, and SSU) and analyses of morphological features were used to describe the new species in the family Cystocloniaceae. Calliblepharis yasutakei sp. nov. grouped with C. fimbriata, C. rammediorum, C. occidentalis and C. jolyi in a clade with full support for the rbcL analysis, representing a distinct lineage within the genus. Phylogenetic and vegetative morphological comparisons demonstrated that the new Hawaiian species is most closely related to C. rammediorum from Israel (rbcL similarity of 96.3%), although no female reproductive structures were found to allow a more comprehensive comparison. In order to determine whether C. yasutakei represents the first confirmed report of the genus Calliblepharis in the Hawaiian Islands , phylogenetic and morphological analysis of the Hawaiian Hypnea saidana (=Calliblepharis saidana) specimen acces-sioned at the Bernice P. Bishop Museum was performed. These analyses demonstrated that this specimen belongs to a new species in the genus Hypnea, which is here described as H. tsudae sp. nov. C. yasutakei, in addition to being a new species, is also reported as the first confirmed record of the genus Calliblepharis in the Hawaiian archipelago, and the description of H. tsudae brings the number of species for this genus in Hawaiʻi to eight.
... Besides molecular evidence, morphological data also support assigning the Brazilian specimens to C. saidana. This species was described from Japan as Hypnea saidana Holmes (Holmes 1895) and was transferred to Calliblepharis when Yang & Kim (2017) carried out molecular studies on topotype specimens. Calliblepharis saidana is mainly distinguished by its loose branching, acute apices and arcuate branches, tapering to points (Holmes 1895, Tanaka 1941, Millar 1990, N'Yeurt & Payri 2010. ...
... The use of a molecular approach in plants from the type locality was crucial to show the true placement of specimens previously named as Craspedocarpus jolyi within the genus Calliblepharis. As a result of recent phylogenetic studies, new species and new combinations have been added to Calliblepharis (Díaz-Tapia et al. 2013, D'Archino et al. 2015, Yang & Kim 2017, Hoffman et al. 2018. However, the establishment of a genomebased phylogeny is still crucial to address phylogenetic and taxonomic questions unresolved in Calliblepharis sensu lato. ...
Among the ten species currently accepted in the genus Calliblepharis, the only representative in the Western Atlantic is C. occidentalis. The endemic Brazilian species C. jolyi was merged into Craspedocarpus, but its generic placement is unclear. Because some species superficially resemble flat Hypnea, our knowledge about the diversity and distribution of Calliblepharis could be incomplete. Therefore, to reassess species diversity of Calliblepharis in Brazil, we carried out an rbcL-based molecular survey of specimens collected along the Brazilian coast, including the type locality of C. jolyi (State of Espírito Santo). Molecular analysis of topotypes supported the reinstatement of Calliblepharis jolyi. This species was placed within Calliblepharis sensu lato in a fully supported clade with C. fimbriata, C. occidentalis, C. rammediorum and Calliblepharis sp. Calliblepharis jolyi is easily distinguished from its congeners by the presence of conspicuous lenticular thickenings. A new record of C. saidana, a species so far known only from the Indo-Pacific, was also revealed for the Atlantic Ocean. Samples collected in São Paulo were grouped with C. saidana from Japan (type locality) with rbcL intraspecific divergence of 0.8%. Taken together, our results provide a better understanding of flat Cystocloniaceae, focusing on Brazilian members and their affinities with other species. Our study call attention to the importance of a molecular approach in specimens from type localities, using information of the original descriptions to define their taxonomic status.
... Significant proposals on the classification of Hypnea have been made on the basis of molecular analyses using the rbcL and cox1 genes which have proven effective in determining the relationships within Hypnea and discriminating and delimiting its species (Fredericq et al. 1996;Yamagishi and Masuda 2000;Geraldino et al. 2006Geraldino et al. , 2010Geraldino et al. , 2015Manghisi et al. 2011;Wolf et al. 2011;Kim and Boo 2012;Wiriyadamrikul et al. 2013;Bast et al. 2014;Boo et al. 2014;Jesus et al. 2014Jesus et al. , 2016Jesus et al. , 2018Jesus et al. , 2019Nauer et al. 2014Nauer et al. , 2015Nauer et al. , 2016Nauer et al. , 2019D'Archino et al. 2015;Ale et al. 2016;Hun Boo et al. 2016;Nurcahyanti et al. 2016;Vázquez-Delfín et al. 2016;Yang and Kim 2017). ...
A phylogenetic analysis of representatives of the red algal genus Hypnea in the Cuban archipelago was carried out utilizing two genetic markers: plastid rbcL and the mitochondrial cox1. Percent of base pair differ- ences and Bayesian analyses indicate the presence of three species of Hypnea in Cuba: H. musciformis, H. spinella, and H. cryptica. The species H. cryptica, first described from Brazil, is confirmed here for the island of Cuba. This first record for the island of Cuba expands the distribution of this species into the Caribbean Sea
... Therefore, the synonymy proposed by Guimarães & Pereira (1993) is not supported by molecular data, but some authors also did not follow the synonimisation (e.g. Hoffman et al. 2018;Wynne 2011;Yang & Kim 2017). The two species also differ in habit in that C. fimbriata exhibits broad blades reaching up to 20 mm wide (Anderson et al. 2016;Kützing 1868;Thrainsson 1986) and the syntype of C. fimbriata (see Guimarães & Pereira 1993, fig. ...
... A molecular account of South African Rhodophyta based on rbcL phylogenies by Hommersand & Fredericq (2003) reported problems for resolving Calliblepharis as a monophyletic group. Similar problems were raised in subsequent papers; for example, Díaz-Tapia, Boo et al. (2013), D'Archino et al. (2015, Kim (2017), andHoffman et al. (2018). At present, we treat the genus as Calliblepharis sensu lato, composed of at least five morphological groups defined by a combination of vegetative and reproductive characters. ...
... Group IV consists of Calliblepharis saidana, which formed a single lineage with unresolved relationships and placed as sister to group III. Yang & Kim (2017) proposed the new combination C. saidana to accommodate specimens previously identified as Hypnea saidana Holmes in Korea and Japan. Calliblepharis saidana has incipient rosettes and nonostiolate cystocarps, like C. celatospora, C. fimbriata and C. occidentalis, but differs from these by the axial filament surrounded by filaments and reticulate placentation. ...
Calliblepharis occidentalis was originally described from northeastern Brazil and later reduced to synonymy with C. fimbriata from South Africa. It was later reinstated, but phylogenetic relationships with other congeners and several morphological characters remain poorly understood. In the present article, topotype specimens of C. occidentalis were reinvestigated using molecular, morphological and anatomical data, and its phylogenetic position was inferred on the basis of rbcL and SSU rRNA. Calliblepharis occidentalis was characterised mainly by the axial filament surrounded by medullary cells, weakly developed rosettes in the outer cortex and carposporophytes with central columns. Our molecular phylogeny showed that C. occidentalis is distinct from C. fimbriata (rbcL divergence of 4.5%). Relationships among the 10 species currently assigned to Calliblepharis remain unresolved and whether they form a monophyletic clade is uncertain. Our results suggest that this genus is composed of five morphological groups, here called Calliblepharis sensu lato, based on vegetative and reproductive features such as type of placentation in the carposporophyte, axial filament arrangement and formation of rosettes. The morphological groups might represent different genera, but this hypothesis needs to be tested with better resolved phylogenies based on more comprehensive gene datasets.
... Although molecular analyses using the rbcL gene have discovered many unrecognized taxa and contributed to red algal phylogenetic reappraisal (Boo et al., 2016;Díaz-Tapia et al., 2017;Rousseau et al., 2017), detailed morphological studies of female reproductive structure anatomy should still be recognized as important (Gurgel & Fredericq, 2004;Kang & Kim, 2016;Yang & Kim, 2017). Systematics of the Halymeniaceae have undergone various revisions. ...
Female reproductive structures, especially ampullae, play a significant role in defining generic boundaries in the family Halymeniaceae, despite recent advances in phylogenetic analyses aided by molecular data. Surveys of red algae from the Korean subtidal zone have revealed new evolutionary lineages within this family. We here propose a new genus in the Halymeniaceae, Nesoia H.W.Lee & M.S.Kim, gen. nov., with a description of N. delicatula sp. nov., N. pulchella sp. nov. and N. latifolia (P. Crouan & H. Crouan ex Kützing) comb. nov., based on Halymenia latifolia. Nesoia, which constitutes a distinct lineage of the Halymeniaceae in rbcL phylogenetic trees, has a foliose thallus with a single-layered cortex and medulla of anticlinal filaments. Carpogonial ampullae bear two-celled carpogonial branches and auxiliary cell ampullae have multiple orders of branching, mostly up to three or four. The auxiliary cell ampullary system of Nesoia is a unique type previously undescribed in the Halymeniaceae: each cell of the first-order ampullary filament produces bilateral second-order filaments, and the auxiliary cell is the basal cell of one of the two second-order filaments borne on the second or third cell of the first-order filament. Discovery of the new genus Nesoia suggests the need for more careful and systematic exploration of species diversity in this group of red algae.
Abbreviations: CoI-5P, 5' end of the cytochrome c oxidase 1 gene; Ml, maximum likelihood; NJ, neighbor joining; rbcl, gene encoding for the large subunit of the enzyme ribulose-1, 5-bisphosphate carboxylase-oxygenase. Abstract the genus Hypnea is well known to be rich in cryptic species, with many new species being recognized recently based on molecular data. therefore, we used mitochondrial (CoI-5P) and plastid (rbcl) molecular markers, in addition to aspects of morphology, to investigate the diversity of Hypnea on the coast of South Florida. Molecular divergences and phylogenetic inferences indicated that four species were present: H. caraibica and H. cryptica, both new occurrences on the Florida coast; H. musciformis, already previously reported and H. spiniformis sp. nov. described in this work. our data reinforce the importance of studies with an integrative taxonomy approach to delimit species and correctly identify them.
Complete and/or partial DNA sequences for the plastid-encoded rbc L gene and the 5′ end of the mitochondrial cytochrome oxidase I (COI-5P) gene were used to re-examine the systematics of Hypnea species (Cystocloniaceae, Rhodophyta) from North Carolina, USA. These data, combined with light microscopic observations, indicate that two species ( Hypnea cryptica and H . musciformis sensu stricto ) are present in nearshore waters of coastal North Carolina. Molecular and morphological analyses with topotype material of Hypnea volubilis from North Carolina offshore waters revealed that it and Calliblepharis saidana are conspecific. Hypnea volubilis is proposed as a heterotypic synonym of C. saidana . This is the first report of Calliblepharis from the United States Atlantic coast and only the second report from the western hemisphere.
The classification of Cystoclonium obtusangulum has been questioned since the species was first described by Hooker and Harvey as Gracilaria? obtusangula. The objective of this study was to provide the first comprehensive taxonomic analysis of Cystoclonium obtusangulum, based on DNA sequences coupled with morphological observations made on syntype specimens and new collections. Sequence divergences of rbcL, UPA and COI‐5P, and maximum‐likelihood phylogenies for rbcL and 18S demonstrated that specimens identified as Cystoclonium obtusangulum represent a clade of two distinct species that is distantly related to the generitype Cystoclonium purpureum. A new genus, Meridionella gen. nov. is proposed for this clade. The two species placed in this new genus were morphologically indistinguishable cryptic species, but have disjunct distributions, with Meridionella obtusangula comb. nov. found from temperate to cold coasts of South America and the Falkland Islands and Meridionella antarctica sp. nov., occurring in Antarctic waters. Vegetative and reproductive characters of Meridionella gen. nov. are described, and implications of our results for the biogeography of the family Cystocloniaceae are discussed.
