PHYLOGENETIC RELATIONSHIPS WITHIN THE GENUS HYPNEA (GIGARTINALES,RHODOPHYTA), WITH A DESCRIPTION OF H. CAESPITOSA SP. NOV.1

Species discrimination within the gigartinalean red algal genus Hypnea has been controversial. To help resolve the controversy and explore phylogeny within the genus, we determined rbcL sequences from 30 specimens of 23 species within the genus, cox1 from 22 specimens of 10 species, and psaA from 16 species. We describe H. caespitosa as a new species characterized by a relatively slender main axis; a pulvinate growth habit with entangled, anastomosing, and subulate uppermost branches; and unilaterally borne tetrasporangial sori. The new species occurs in the warm waters of Malaysia, the Philippines, and Singapore. The phylogenetic trees of rbcL, psaA, and cox1 sequences showed a distant relationship of H. caespitosa to H. pannosa J. Agardh from Baja California and the marked differentiation from other similar species. The rbcL + psaA tree supported monophyly of the genus with high bootstrap values and posterior probabilities. The analysis revealed three clades within the genus, corresponding to three sections, namely, Virgatae, Spinuligerae, and Pulvinatae first recognized by J. G. Agardh. Exceptions were H. japonica T. Tanaka in Pulvinatae and H. spinella (C. Agardh) Kütz. in Spinuligerae.     

GLOBAL SYSTEMATIC AND PHYLOGENETIC ANALYSIS OF SARGASSUM IN THE GULF OF MEXICO,CARIBBEAN AND PACIFIC BASIN

Sargassum is one of the most species‐rich genera in the brown algae with over 400 described species worldwide. The bulk of these species occurs in Pacific‐Indian ocean waters with only a small portion found on the Atlantic side of the Isthmus of Panama. Sargassum also has one of the most subdivided and complex taxonomic systems used within the algae. Systematic distinctions within the genus are further complicated by high rates of phenotypic variability in several key morphological characters. Molecular analyses in such systems should allow testing of systematic concepts while providing insights into speciation and evolutionary patterns. Global molecular phylogenetic analyses using both conserved and variable regions of the Rubisco operon (rbcL and rbcL‐IGS‐rbcS) were performed with species from the Gulf of Mexico, Caribbean, and Pacific basin. Results confirm earlier analyses based on rbcL‐IGS‐rbcS from Pacific species at the subgeneric and sectional level while providing additional insights into the systematics and phylogenetics on a global scale. For example, species east of the Isthmus of Panama form a distinct well‐resolved clade within the tropical subgenus. This result in sharp contrast to traditional systematic treatments but provides a window into the evolutionary history of this genus in the Pacific and Atlantic Ocean basins and a possible means to time speciation events.     

NEW INSIGHTS IN THE TAXONOMY OF THE CERAMIUM SINICOLA COMPLEX: RESURRECTION OF CERAMIUM INTERRUPTUM (CERAMIACEAE,RHODOPHYTA)1

Phylogenetic relationships of the Ceramium sinicola complex (C. interruptum and C. sinicola) including C. codicola were studied using nucleotide sequences of rbcL and small subunit rDNA, and the RUBISCO spacer was used for sequence comparison of each species. A reassessment of the taxonomic rank and the evolutionary trend within the complex was inferred from a comparative morphological study and molecular data sets based on 11 samples from eight populations from the Pacific coast of the United States and Mexico. Intraspecific relationships were poorly resolved, but the resurrection of C. interruptum as a distinct species was strongly supported by both morphological and molecular data. Ceramium interruptum is distinguished by the combination of the following features: thalli uncorticated at the first internode above the dichotomy, presence of four corticating filaments, 7–11 segments between branching points, rhizoids digitate, and epiphytic on a variety of hosts. Our molecular analyses show that C. sinicola is the sister group to C. codicola, and C. interruptum is basal to them. These phylogenetic relationships allowed for an assessment of the trend in the evolution of cortication pattern and attachment mode to the host.     

MOLECULAR PHYLOGENY OF THE GENUS KUMANOA (BATRACHOSPERMALES,RHODOPHYTA)1

Species belonging to the newly established genus Kumanoa were sampled from locations worldwide. DNA sequence data from the rbcL gene, cox1 barcode region, and universal plastid amplicon (UPA) were collected. The new sequence data for the rbcL were combined with the extensive batrachospermalean rbcL data available in GenBank. Single gene rbcL results showed the genus Kumanoa to be a well‐supported clade, and there was high statistical support for many of the terminal nodes. However, with this gene alone, there was very little support for any of the internal nodes. Analysis of the concatenated data set (rbcL, cox1, and UPA) provided higher statistical support across the tree. The taxa K. vittata and K. amazonensis formed a basal grade, and both were on relatively long branches. Three new species are proposed, K. holtonii, K. gudjewga, and K. novaecaledonensis; K. procarpa var. americana is raised to species level. In addition, the synonymy of K. capensis and K. breviarticulata is proposed, with K. capensis having precedence. Five new combinations are made, bringing the total number of accepted species in Kumanoa to 31. The phylogenetic analyses did not reveal any interpretable biogeographic patterns within the genus (e.g., K. spermatiophora from the tropical oceanic island Maui, Hawaii, was sister to K. faroensis from temperate midcontinental Ohio in North America). Previously hypothesized relationships among groups of species were not substantiated in the phylogenetic analyses, and no intrageneric classification is recommended based on current knowledge.     

A TAXONOMIC STUDY OF THE GENUS PADINA (DICTYOTALES,PHAEOPHYCEAE) INCLUDING THE DESCRIPTIONS OF FOUR NEW SPECIES FROM JAPAN,HAWAII, AND THE ANDAMAN SEA1

A taxonomic study of the genus Padina from Japan, Southeast Asia, and Hawaii based on morphology and gene sequence data (rbcL and cox3) resulted in the recognition of four new species, that is, Padina macrophylla and Padina ishigakiensis from Ryukyu Islands, Japan; Padina maroensis from Hawaii; and Padina usoehtunii from Myanmar and Thailand. All species are bistratose and morphologically different from one another as well as from any known taxa by a combination of characters relating to degree of calcification; the structure, position, and arrangement of hairlines (HLs) and reproductive sori; and the presence or absence of rhizoid‐like groups of hairs and an indusium. Molecular phylogenetic analyses demonstrated a close relationship between P. ishigakiensis, P. macrophylla, P. maroensis, and Padina australis Hauck. The position of P. usoehtunii, however, was not fully resolved, being either sister to a clade comprising the other three new species and P. australis in the rbcL tree or more closely related to a clade comprising several other recently described species in the cox3 tree. The finding of the four new species demonstrates high species diversity particularly in southern Japan. The following characters were first recognized here to be useful for species delimitation: the presence or absence of small rhizoid‐like groups of hairs on the thallus surface, structure and arrangement of HLs on both surfaces either alternate or irregular, and arrangement of the alternating HLs between both surfaces in equal or unequal distance. The evolutionary trajectory of these and six other morphological characters used in species delineation was traced on the phylogenetic tree.     

A NEW BROWN ALGAL ORDER,ISHIGEALES (PHAEOPHYCEAE), ESTABLISHED ON THE BASIS OF PLASTID PROTEIN‐CODING rbcL,psaA, AND psbA REGION COMPARISONS1

The brown algal family Ishigeaceae currently includes a single genus, Ishige Yendo, with two species. The relationship of the family to other brown algal lineages is less studied in terms of their plastid ultrastructure and molecular phylogeny. We determined the sequences of rbcL from four samples of the two Ishige species and nine putative relatives and the psaA and psbA sequences from 37 representatives of the brown algae. Analyses of individual and combined data sets resulted in similar trees; however, the concatenated data gave greater resolution and clade support than each individual gene. In all the phylogenies, the Phaeophyceae was well resolved, the Ectocarpales being placed in a terminal position and the Ishigeaceae ending up in a basal position. From our ultrastructural study, we concluded that the pyrenoid is absent in the Ishigeaceae, despite the presence of a rudimentary pyrenoid in I. okamurae. These results suggest that the Ishigeaceae is an early diverging brown lineage. Our molecular and morphological data, therefore, lead us to exclude the Ishigeaceae from the Ectocarpales s.l., which have an elaborate pyrenoid, and to propose its own order Ishigeales ord. nov. The Ishigeales is distinguished by oligostichous structure of thalli, phaeophycean hairs formed within cryptostomata, unilocular sporangia transformed from terminal cortical cells, and plurilocular sporangia lacking sterile terminal cells. This study is the first to document the utility of the psaA and psbA sequences for brown algae and also the first report on the multigene phylogeny of the Phaeophyceae based on three protein‐coding plastid genes.     

New Systematic Insights in the Ceramium Sinicolacomplex: Resurrection of C. Interruptum S. & G. (Ceramiaceae,Rhodophyta)

Setchell & Gardner (1924) provided a taxonomic treatment for the genus Ceramium from lower California and the Gulf of California, Mexico, in which they described several new species, including C. sinicola S. & G. described from Ensenada, BCN and C. interruptum S. & G. described from the vicinity of La Paz, BCS. The latter was later reduced to variety rank in C. sinicolaby Dawson (1950), a taxonomic decision that has subsequently been widely adopted. Phylogenetic analyses inferred from three molecular markers (chloroplast‐encoded rbcL, the RUBISCO spacer, and nuclear encoded SSU rDNA) from recent collections from the Pacific North‐west (California, Oregon) and the Gulf of California reveal a well supported assemblage of three corticated taxa: C. codicolaJ. Ag. 1894, C. sinicola and C. sinicola var. interruptum. Sequence divergence values among the three taxa are sufficient to warrant separate species ranking; hence, we reinstate C. interruptum as a widely distributed epiphyte for the region. C. sinicolais more closely related to C. codicolathan to C. interruptum, with the former two restricted to the host Codium. The molecule‐based relationships are congruent with evolutionary trends in cortication pattern and attachment mode.     

Species Diversity,Phylogeny and Large Scale Biogeographic Patterns of the Genus Padina (Phaeophyceae,Dictyotales)

The brown algal genus Padina (Dictyotales, Phaeophyceae) is distributed worldwide in tropical and temperate seas. Global species diversity and distribution ranges, however, remain largely unknown. Species‐level diversity was reassessed using DNA‐based, algorithmic species delineation techniques based on cox3 and rbcL sequence data from 221 specimens collected worldwide. This resulted in estimates ranging from 39 to 61 putative species (ESUs), depending on the technique as well as the locus. We discuss the merits, potential pitfalls, and evolutionary and biogeographic significance of algorithmic species delineation. We unveil patterns whereby ESUs are in all but one case restricted to either the Atlantic or Indo‐Pacific Ocean. Within ocean basins we find evidence for the vast majority of ESUs to be confined to a single marine realm. Exceptions, whereby ESUs span up to three realms, are located in the Indo‐Pacific Ocean. Patterns of range‐restricted species likely arise by repeated founder events and subsequent peripatric speciation, hypothesized to dominate speciation mechanisms for coastal marine organisms in the Indo‐Pacific. Using a three‐gene (cox3, psaA and rbcL), relaxed molecular clock phylogenetic analysis we estimated divergence times, providing a historical framework to interpret biogeographic patterns.     

THE RECLASSIFICATION OF LESSONIA LAMINARIOIDES (LAMINARIALES,PHAEOPHYCEAE): PSEUDOLESSONIA GEN. NOV.1

A new genus, Pseudolessonia, is proposed for the kelp Lessonia laminarioides Postels et Ruprecht (Laminariales, Phaeophyceae), which occurs on the northwest side of the Sea of Okhotsk, in the northwest Pacific Ocean. Pseudolessonia is monotypic and differs from Lessonia in its short primary stipes and its corrugated, unilaterally arranged blades with entire margins. This species is transferred on the basis of morphology and plastid gene sequence comparisons. We determined psaA and rbcL gene sequences from 17 taxa of Pseudolessonia, Lessonia, and putative relatives. Analyses of individual and combined data sets resulted in congruent trees showing a clear separation of Pseudolessonia laminarioides from Lessonia, but suggesting its sister relationships with the clade of Nereocystis, Macrocystis, Pelagophycus, and Postelsia in the North Pacific Ocean. On the other hand, Lessonia species from the South Pacific Ocean formed a strongly supported clade. The results indicate that the basal splitting of the blade, which has been considered a diagnostic character for the family Lessoniaceae, is a result of convergent evolution.     

The systematics of Lobophora (Dictyotales,Phaeophyceae) in the western Atlantic and eastern Pacific oceans: eight new species

Lobophora is a common tropical to temperate genus of brown algae found in a plethora of habitats including shallow and deep‐water coral reefs, rocky shores, mangroves, seagrass beds, and rhodoliths beds. Recent molecular studies have revealed that Lobophora species diversity has been severely underestimated. Current estimates of the species numbers range from 100 to 140 species with a suggested center of diversity in the Central Indo‐Pacific. This study used three molecular markers (cox3, rbcL, psbA), different single‐marker species delimitation methods (GMYC, ABGD, PTP), and morphological evidence to evaluate Lobophora species diversity in the Western Atlantic and the Eastern Pacific oceans. Cox3 provided the greatest number of primary species hypotheses(PSH), followed by rbcL and then psbA. GMYC species delimitation analysis was the most conservative across all three markers, followed by PTP, and then ABGD. The most informative diagnostic morphological characters were thallus thickness and number of cell layers in both the medulla and the dorsal/ventral cortices. Following a consensus approach, 14 distinct Lobophora species were identified in the Western Atlantic and five in the Eastern Pacific. Eight new species from these two oceans were herein described: L. adpressa sp. nov., L. cocoensis sp. nov., L. colombiana sp. nov., L. crispata sp. nov., L. delicata sp. nov., L. dispersa sp. nov., L. panamensis sp. nov., and L. tortugensis sp. nov. This study showed that the best approach to confidently identify Lobophora species is to analyze DNA sequences (preferably cox3 and rbcL) followed by comparative morphological and geographical assessment.     

57 Systematics of hydropuntia montagne (gracilariaceae,gracilariales, rhodophyta) based on comparative morphology and rbcL sequence analysis

The taxonomic validity of the genus Hydropuntia Montagne (1843) (including Polycavernosa) within the Gracilariaceae (Gracilariales, Rhodophyta) is controversial. Morphological characters that define species of Hydropuntia are said to be variable and to overlap with those of Gracilaria. Here we present a global phylogenetic study of the family based on a Bayesian analysis of a large rbcL DNA sequence dataset indicating that the genus Hydropuntia forms a well supported monophyletic clade within the family, and recognize Hydropuntia as a genus distinct from Gracilaria. We also conducted smaller phylogenetic analyses in which thirty four Hydropuntia rbcL sequences resulted in two major clades within the genus, comprising a Caribbean clade and an Indo‐Pacific clade. Diagnostic reproductive stages that separate these two clades will be illustrated.     

GRACILARIOPSIS MCLACHLANII SP. NOV. AND GRACILARIOPSIS PERSICA SP. NOV. OF THE GRACILARIACEAE (GRACILARIALES,RHODOPHYCEAE) FROM THE INDIAN OCEAN1

Two new species of Gracilariopsis from the Indian Ocean are proposed—Gracilariopsis (Gp.) mclachlanii Buriyo, Bellorin et M. C. Oliveira sp. nov. from Tanzania and Gracilariopsis persica Bellorin, Sohrabipour et E. C. Oliveira sp. nov. from Iran—based on morphology and DNA sequence data (rbcL gene and SSU rDNA). Both species fit the typical features of Gracilariopsis: axes cylindrical throughout, freely and loosely ramified up to four orders, with an abrupt transition in cell size from medulla to cortex, cystocarps lacking tubular nutritive cells and superficial spermatangia. Nucleotide sequence comparisons of rbcL and SSU rDNA placed both species into the Gracilariopsis clade as distinct species from all the accepted species for this genus, forming a deeply divergent lineage together with some species from the Pacific. The new species are very difficult to distinguish on morphological grounds from other species of Gracilariopsis, stressing the importance of homologous molecular marker comparisons for the species recognition in this character‐poor genus.     

23 Phylogenetic position of Alaria Fistulosa (laminariales,phaeophyceae) based on nuclear its and plastid coding genes

Alaria is a common kelp genus generally found in the lower intertidal and shallow subtidal regions of rocky shores in the cold waters of the northern Hemisphere. About 16 species are currently recognized worldwide and, of these, A. fistulosa is distinguished by having hollow midrib and large blades with 10–30 m in length and 30–90 cm in width. It is the only canopy‐producing kelp in the northwest Pacific, where it is restricted to the waters of north Hokkaido, Kamchatka, Aleutian Islands, and Alaska. In order to know the phylogenetic position of A. fistulosa, sequences of nr DNA ITS and plastid rbcL including spacer and psaA regions were determined in A. fistulosa and compared with homologous positions of newly sequenced putative relatives and with published sequences of other kelp species. Combined data of ITS and Rubisco spacer show that A. fistulosa was more related to the clade of Lessoniopsis and Pterygophora than to the clade of other species of Alaria, which is supported by the rbcL and psaA sequence data. The topologies from nuclear and plastid DNA sequences lead to phylogenetic independence of A. fistulosa, which is clearly different from the genus Alaria.     

CHEMOTAXONOMY IN MARINE ALGAE: CHEMICAL SEPARATION OF SOME LAURENCIA SPECIES (RHODOPHYTA) FROM THE GULF OF CALIFORNIA1

Species separation in the genus Laurencia (Rhodomelaceae, Rhodophyta) is complicated by the high degree of morphological variation within the species. Chemical investigations on a worldwide basis of over 15 species indicate that 1 or more of the halogenated natural products synthesized by Laurencia are unique to each species. Our chemical investigations of Laurencia pacifica, as presently understood from the Gulf of California, indicate that more than 1 species had been included under this name. Thin layer chromatographic (TLC) comparisons of the halogenated components of 3 recognizable forms of “L. pacifica” were completed. The results revealed 3 distinct forms, with halogenated products unique to each form. In each form the observed chemical characters had been previously isolated and identified and could now be positively assigned to their algal source. Comparisons were also conducted with L. pacifica Kylin (1941) from the type locality of the species, La Jolla, California, and revealed that it contained another halogenated product different from those isolated from the Gulf species. We conclude that 3 species of Laurencia have been elucidated in the Gulf of California and these are separate from L. pacifica Kylin. Each species can be distinguished by its characteristic array of halogenated compounds. Comparative thin layer chromatography of the lipid components of morphologically similar Laurencia species should prove to be a useful new taxonomic aid.     

MOLECULAR PHYLOGENY OF THE GREEN ALGAL ORDER PRASIOLALES (TREBOUXIOPHYCEAE,CHLOROPHYTA)1

The systematics of the Prasiolales was investigated by phylogenetic inference based on analyses of the rbcL and 18S rRNA genes for representatives of all four genera currently attributed to this order (Prasiococcus, Prasiola, Prasiolopsis, Rosenvingiella), including all type species. The rbcL gene had higher sequence divergence than the 18S rRNA gene and was more useful for phylogenetic inference at the ranks of genus and species. In the rbcL gene phylogeny, three main clades were observed, corresponding to Prasiola, Prasiolopsis, and Rosenvingiella. Prasiococcus was nested among species of Prasiola occurring in subaerial and supralittoral habitats. Trichophilus welckeri Weber Bosse, a subaerial alga occurring in the fur of sloths in Amazonia, was closely related to Prasiolopsis ramosa Vischer. The species of Prasiola were grouped into three well‐supported clades comprising (i) marine species, (ii) freshwater and terrestrial species with linear blades, and (iii) terrestrial species with rounded or fan‐shaped blades. Sequence divergence was unexpectedly low in the marine group, which included species with different morphologies. For the 18S rRNA gene, the phylogenetic analyses produced several clades observed for the rbcL gene sequence analysis, but, due to very little sequence variation, it showed considerably lower resolution for inference at the species and genus levels. Due to the low support of some internal branches, the results of the analyses did not allow an unambiguous clarification of the origin and the early evolution of the Prasiolales.     

PHYLOGEOGRAPHY OF THE SPOTTED SAND BASS, PARALABRAX MACULATOFASCIATUS: DIVERGENCE OF GULF OF CALIFORNIA AND PACIFIC COAST POPULATIONS

Abstract Have the warm tropical waters and currents of the southern Gulf of California, Mexico (also known as the Sea of Cortez), formed a barrier to gene flow, resulting in disjunct populations in the upper gulf that are isolated from the outer Pacific Coast? Phylogeographic and genetic divergences of the spotted sand bass, Paralabrax maculatofasciatus, from three Gulf of California and two outer Pacific coastal locations were tested using mitochondrial DNA (mtDNA) control region sequences. Sequence data from two congeners that are sympatrically distributed along the outer Pacific Coast, the barred sand bass, P. nebulifer, and the kelp bass, P. clathratus, were used to gauge the levels of genetic divergences. Differences among the three species and between the northern gulf and outer Pacific coastal populations of P. maculatofasciatus also were analyzed using 40 allozymic presumptive gene loci. Allozyme and mtDNA analyses each revealed many fixed differences among the species. Three significant allozymic frequency differences and two fixed mtDNA substitutions differentiated the gulf and outer Pacific coastal populations of P. maculatofasciatus. Three unique mtDNA haplotypes and three unique allozyme alleles were identified from the outer Pacific coastal population. The gulf sites contained four unique mtDNA haplotypes and six unique allozyme alleles. Partitioning of the mtDNA variation revealed that 72% of the variance occurred between the gulf and outer Pacific Coast, 20% between sampling sites in the two regions, and 8% within the sites. There appears to be little gene flow across the waters of the southern Baja Penninsula, producing divergence estimated as 120,000 to 600,000 years between the outer Pacific coastal and the Gulf of California populations. This separation level may date to a hypothesized seaway closure near La Paz, Mexico, during the mid‐Pleistocene, and characterizes other fish populations. A second pattern of deeper allopatric species‐level divergences in some other fishes may date to a Pliocene closure of a mid‐Baja Penninsular seaway. Significant differences also were discerned in P. maculatofasciatus between the San Diego and central Baja California coastal sites and between the upper/central and the lower gulf locations. Variation between locations in the two regions may be indicative of larval retention and low adult migration, which needs to be tested further.     

Toward an inordinate fondness for stars,beetles and Lobophora? Species diversity of the genus Lobophora (Dictyotales,Phaeophyceae) in New Caledonia

Until the recent use of molecular markers, species diversity of Lobophora, an ecologically important brown algal genus with a worldwide distribution in temperate and tropical seas, has been critically underestimated. Using a DNA‐based taxonomic approach, we re‐examined diversity of the genus from New Caledonia in the Southwest Pacific Ocean. First, species were delineated using general mixed Yule coalescent‐based and barcoding gap approaches applied to a mitochondrial cox3 data set. Results were subsequently confirmed using chloroplast psbA and rbcL data sets. Species delimitation analyses agreed well across markers and delimitation algorithms, with the barcoding gap approach being slightly more conservative. Analyses of the cox3 data set resulted in 31–39 molecular operational taxonomic units (MOTUs), four of which are previously described species (L. asiatica, L. crassa, L. nigrescens s.l., L. pachyventera). Of the remaining MOTUs for which we obtained a representative number of sequences and results are corroborated across analyses and genes, we described 10 species de novo: L. abaculusa, L. abscondita, L. densa, L. dimorpha, L. gibbera, L. hederacea, L. monticola, L. petila, L. rosacea, and L. undulata. Our study presents an excellent case of how a traditional morphology‐based taxonomy fails to provide accurate estimates of algal diversity. Furthermore, the level of Lobophora diversity unveiled from a single locality in the Pacific Ocean raises important questions with respect to the global diversity of the genus, the distributions and range sizes of the individual species, as well as the mechanisms facilitating coexistence.     

Phylogenetic position of the Hawaiian geraniums based on rbcL Sequences

The seven currently recognized species of Geranium endemic to the Hawaiian Islands are unusual in their shrubby or arborescent habit and unlobed, parallel-veined leaves rather than the palmately cleft or lobed leaves and herbaceous habit typical of the genus. Their placement within the genus and their biogeographic source have been obscured by this morphological distictiveness and the limited resolution of relationships on the basis of morphology in the very speciose subgenus Geranium. Phylogenetic analysis of rbcL gene sequences provides strong support for the monophyly of the Hawaiian group, and indicates that the Hawaiian clade is deeply nested within section Geranium rather than comprising a separate section. The continental relatives studied to date with the greatest similarity in sequence to the Hawaiian group are native to the Americas rather than Asia or the Pacific. The Hawaiian species are extremely similar to one another in rbcL sequence, while the tree topology obtained is consistent with a basal position for Geranium arboreum within the group.     

Reinstatement of Myelophycus caespitosus (Scytosiphonaceae,Phaeophyceae) from Japan

Reinstatement of Myelophycus caespitosus Kjellman (Ectocarpales s.l., Phaeophyceae) described from western Japan is proposed based on the comparisons of DNA sequences of Japanese Myelophycus specimens, the type specimen of Chordaria simplex and lectotypified specimen of Myelophycus caespitosus. In the genetic analyses using mitochondrial cox1 and cox3, chloroplast atpB, psbA and rbcL DNA sequences, the specimens morphologically referable to M. simplex formed two distinct clades (clade‐1 and clade‐2) supported by high statistical values. Clade‐1 was distributed on the western coast of Japan (Honshu, Shikoku and Kyushu) from the Kii Peninsula to Tsushima and the Pacific coast of Aomori, northeastern Honshu, and clade‐2 on the Pacific coast of central Honshu from Sanriku to the Kii Peninsula. Based on the cox3 DNA sequences and the location of the type locality at Shimoda, Izu Peninsula, clade‐2 was concluded to correspond to true M. simplex. There were no recognizable differences in the representative morphological features (height and diameter of sporophytes and gametophytes, number of cells comprising cortical, subcortical and medullary layers, thickness of plurilocular gametangia, or length and diameter of unilocular zoidangia) between the specimens included in the two taxa. Analyses of the specimens of the two taxa, including sympatric populations on the Kii Peninsula, using a nuclear genetic marker ocm3 did not suggest any genetic exchanges between the two taxa. On the other hand, cox3 gene sequence of the voucher specimen of Myelophycus caespitosus collected from Goto, Kyushu, western Japan housed in the UPS herbarium was included in clade‐1. In conclusion, we propose the reinstatement of M. caespitosus and to lectotypify the specimen in UPS. Korean Myelophycus specimens reported from Cheju Island and Wando were considered to belong to M. caespitosus based on the reported rbcL sequences.     

BIODIVERSITY OF CORALLINE ALGAE IN THE NORTHEASTERN ATLANTIC INCLUDING CORALLINA CAESPITOSA SP. NOV. (CORALLINOIDEAE,RHODOPHYTA)1

The Corallinoideae (Corallinaceae) is represented in the northeastern Atlantic by Corallina officinalis L.; Corallina elongata J. Ellis et Sol.; Haliptilon squamatum (L.) H. W. Johans., L. M. Irvine et A. M. Webster; and Jania rubens (L.) J. V. Lamour. The delimitation of these geniculate coralline red algae is based primarily on morphological characters. Molecular analysis based on cox1 and 18S rRNA gene phylogenies supported the division of the Corallinoideae into the tribes Janieae and Corallineae. Within the Janieae, a sequence difference of 46–48 bp (8.6%–8.9%) between specimens of H. squamatum and J. rubens in the cox1 phylogeny leads us to conclude that they are congeneric. J. rubens var. rubens and J. rubens var. corniculata (L.) Yendo clustered together in both phylogenies, suggesting that for those genes, there was no genetic basis for the morphological variation. Within the Corallineae, it appears that in some regions, the name C. elongata has been misapplied. C. officinalis samples formed two clusters that differed by 45–54 bp (8.4%–10.0%), indicating species‐level divergence, and morphological differences were sufficient to define two species. One of these clusters was consistent with the morphology of the type specimen of C. officinalis (LINN 1293.9). The other species cluster is therefore described here as Corallina caespitosa sp. nov. This study has demonstrated that there is a clear need for a revision of the genus Corallina to determine the extent of “pseudocryptic” diversity in this group of red algae.