The Florideophyceae is the most abundant and taxonomically diverse class of red algae (Rhodophyta). However, many aspects of the systematics and divergence times of the group remain unresolved. Using a seven-gene concatenated dataset (nuclear EF2, LSU and SSU rRNAs, mitochondrial cox1, and plastid rbcL, psaA and psbA genes), we generated a robust phylogeny of red algae to provide an evolutionary timeline for florideophyte diversification. Our relaxed molecular clock analysis suggests that the Florideophyceae diverged approximately 943 (817–1,049) million years ago (Ma). The major divergences in this class involved the emergence of Hildenbrandiophycidae [ca. 781 (681–879) Ma], Nemaliophycidae [ca. 661 (597–736) Ma], Corallinophycidae [ca. 579 (543–617) Ma], and the split of Ahnfeltiophycidae and Rhodymeniophycidae [ca. 508 (442–580) Ma]. Within these clades, extant diversity reflects largely Phanerozoic diversification. Divergences within Florideophyceae were accompanied by evolutionary changes in the carposporophyte stage, leading to a successful strategy for maximizing spore production from each fertilization event. Our research provides robust estimates for the divergence times of major lineages within the Florideophyceae. This timeline was used to interpret the emergence of key morphological innovations that characterize these multicellular red algae.
Nucleotide sequences of the nuclear, small-subunit (SSU) ribosomal RNAs, as inferred from polymerase chain reaction (PCR)-amplified products, are presented for Areschougia congesta (Turner) J. Agardh (Solieriaceae), Dasyphloea insignis Montagne (Dumontiaceae), Sarcothalia crassifolia (C. Agardh) Edyvane & Womersley (Gigartinaceae), Nizymenia australis Sonder (Nizymeniaceae), Phacelocarpus peperocarpos (Poiret) Wynne, Ardré & Silva (Phacelocarpaceae), Plocamiocolax pulvinata Setchell, Plocamium angustum (J. Agardh) J.D. Hooker, Plocamium cartilagineum (Linnaeus) Dixon (Plocamiaceae), Rhodymenia linearis J. Agardh (Rhodymeniaceae), and Sphaerococcus coronopifolius Stackhouse (Sphaerococcaceae). Phylogenetic analyses of the SSU sequences between the Plocamiaceae and members of the Sphaerococcaceae, Phacelocarpaceae, and Nizymeniaceae, with which the Plocamiaceae has been associated historically, show SSU differences of between 87 and 105 nucleotides and do not indicate a close relationship. A review of anatomical knowledge of the Plocamiaceae and Pseudoanemoniaceae and new information on vegetative and tetrasporangial development in Plocamium and Plocamiocolax are presented to buttress a case for the Plocamiales ord.nov. Representatives of the Nizymeniaceae and Phacelocarpaceae differ from one another by only nine nucleotides, suggesting that these two taxa are very closely related and perhaps not distinct at the family rank. Key words: Gigartinales, PCR, phylogeny, Plocamiales ord.nov., Pseudoanemoniaceae, Rhodophyta, small-subunit rRNA, systematics.
Glacial vicariance is regarded as one of the most prevalent drivers of phylogeographic structure and speciation among high-latitude organisms, but direct links between ice advances and range fragmentation have been more difficult to establish in marine than in terrestrial systems. Here we investigate the evolution of largely disjunct (and potentially reproductively isolated) phylogeographic lineages within the amphi-boreal kelp Saccharina latissima s. l. Using molecular data (COI, microsatellites) we confirm that S. latissima comprises also the NE Pacific S. cichorioides complex and is composed of divergent lineages with limited range overlap and genetic admixture. Only a few genetic hybrids were detected throughout a Canadian Arctic/NW Greenland contact zone. The degree of genetic differentiation and sympatric isolation of phylogroups suggest that S. latissima s. l. represents a complex of incipient species. Phylogroup distributions compared with paleo-environmental reconstructions of the cryosphere further suggest that diversification within S. latissima results from chronic glacial isolation in disjunct persistence areas intercalated with ephemeral interglacial poleward expansions and admixture at high-latitude (Arctic) contact zones. This study thus supports a role for glaciations not just in redistributing pre-existing marine lineages but also as a speciation pump across multi-glacial cycles for marine organisms otherwise exhibiting cosmopolite amphi-boreal distributions.
DNA barcoding is becoming a widely applied tool for the rapid and accurate identification of eukaryotic species. In this study we used the DNA barcode for large-scale screening of the brown algal family Laminariaceae in Canada. With the examination of 194 COI-5P (59 end of cytochrome c oxidase 1) sequences (150 newly determined) from representatives of this family, we confirmed the presence of 12 species in Canadian waters (Cymathaere triplicata, Laminaria digitata, L. ephemera, L. setchellii, L. solidungula, L. yezoensis, Macrocystis integrifolia, Nereocystis leutkeana, Postelsia palmaeformis, Saccharina groenlandica, S. latissima and S. sessilis). Saccharina groenlandica, a species with a history of taxonomic confusion, was found in the Pacific, Hudson Bay (subarctic) and Atlantic Canada extending greatly our comprehension of the biogeography of this species. Additionally, COI-5P data from S. latissima, combined with ITS results, provided insights into historical distributional patterns and uncovered a hybridization zone between incipient species in this complex. These discoveries highlight how the growth of a worldwide barcode database for the assignment of individuals to genetic species will uncover new perspectives on biogeography and species diversity on a global scale.
Nucleotide sequences of the nuclear, small-subunit ribosomal RNAs, as inferred from polymerase chain reaction amplified products, are introduced for representatives of the Dumontiaceae, Endocladiaceae, Halymeniaceae, and Kallymeniaceae of the order Cryptonemiales sensu Kylin, the Mychodeaceae, Phyllophoraceae, Schizymeniaceae, and Sebdeniaceae of the order Gigartinales sensu Kylin, and the Lomentariaceae and Rhodymeniaceae of the order Rhodymeniales. The new sequences are included in phylogenetic analyses incorporating previously published sequences from additional families of the orders Ahnfeltiales, Ceramiales, Gigartinales, Gracilariales, Palmariales, Plocamiales, and Rhodymeniales. We used the molecular data to test the validity of the taxonomic merger of the orders Gigartinales and Cryptonemiales that was proposed by G.T. Kraft and P.A. Robins in 1985. With only two exceptions (the families Halymeniaceae and Sebdeniaceae), phylogenetic analyses of the SSU data support a monophyletic origin for a combined Gigartinales–Cryptonemiales. We therefore propose the resurrection of a redefined Cryptonemiales to consist, at this time, of only the Halymeniaceae and Sebdeniaceae. Because virtually no elements of the original or recent definitions of the Cryptonemiales survive in the characterization of this taxon, we followed procedures allowed by the International Code of Botanical Nomenclature to designate it the Halymeniales ord.nov. Analysis of molecular data further indicates that the Rhodymeniales is a monophyletic assemblage distinct from both the Gigartinales and Halymeniales; it should not be merged with the Gigartinales as is occasionally suggested. Keywords: Cryptonemiales, Gigartinales, Halymeniaceae, Halymeniales, phylogeny, Rhodophyta, Sebdeniaceae, small-subunit rRNA, systematics.
Small-subunit ribosomal RNA sequence data are presented for Rhodogorgon carriebowensis Norris et Bucher (Rhodogorgonales, Rhodophyta) and Galaxaura marginata (Ellis et Solander) Lamouroux (Nemaliales, Rhodophyta). Data for these species were included in a matrix consisting of small-subunit ribosomal RNA sequences for 70 taxonomically diverse red algal species. Distance, parsimony, and maximum-likelihood analyses of these data all strongly support (100% of bootstrap replicates for distance and parsimony) a close alliance between the Rhodogorgonales and Corallinales. Our alignment includes 19 species representing six florideophyte orders (Acrochaetiales, Batrachospermales, Corallinales, Nemaliales, Palmariales, Rhodogorgonales) in which pit plugs are characterized by two cap layers. Distance and parsimony analyses resolve a monophyletic lineage including all "two-cap-layer" species. Our parsimony-derived molecular phylogeny was used to test a published hypothesis of pit-plug evolution. Four aspects of Pueschel's hypothesis are consistent with the results of our molecular systematic study: (i) "naked" plugs represent the ancestral type, (ii) outer cap layers are homologous structures, (iii) domed outer caps are ancestral to plate-like outer caps, and (iv) cap membranes are a derived feature within the two-cap-layer lineage. Directions for future research are discussed. Key words: Acrochaetiales, Batrachospermales, Corallinales, molecular systematics, Nemaliales, Palmariales, phylogeny, pit plug, Rhodogorgonales, Rhodophyta.
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