LIFE HISTORIES IN THE PHYLLOPHORACEAE (RHODOPHYTA: GIGARTINALES) FROM THE PACIFIC COAST OF NORTH AMERICA. I. GYMNOGONGRUS LINEARIS AND G. LEPTOPHYLLUS1

Carpospores of Gymnogongrus linearis (C. Ag.) J. Ag. collected from Sonoma Co., California were cultured and gave rise to crustose plants. Tetrasporogenesis could not be induced. However, tetraspores from field-collected crustose tetrasporophytes found near G. linearis from San Mateo Co., California were cultured. These field crusts superficially resemble Petrocelis middendorffii (Ruprecht) Kjellman, but differ in size, color, number of tetrasporangia per filament, and distal dichotomous branching of the perithallial filaments. Tetraspores gave rise to upright plants identical to G. linearis. Gymnogongrus leptophyllus J. Ag. collected from California and Baja California, Mexico were found as narrow and wide forms. Narrow form isolates recycled directly without producing a crustose tetrasporophyte. These are interpreted as apogamous. Carpospores of the wide form grew into crusts resembling Petrocelis (=Erythrodermis) haematis Hollenberg. Tetrasporogenesis was induced in culture by abrasion or dehydration. Tetraspores from field-collected crusts and laboratory cultured tetrasporophytes grew into plants identical to G. leptophyllus, completing a sexual life history with an alternation of heteromorphic generations.     

PLASTID DNA RESTRICTION ANALYSIS LINKS THE HETEROMORPHIC PHASES OF AN APOMICTIC RED ALGAL LIFE HISTORY1

Gymnogongrus sp. (Phyllophoraceae) from Nova Scotia, Canada, identified tentatively as G. devoniensis (Greville) Schotter, grows in association with an Erythrodermis-like that forms chains of tetrasporangia or bisporangia. The crust resembles tetrasporophytic phases of other Gymnogongrus species, but in culture both it and the G. devoniensis gametophytes cycle independently by apomictic reproduction. A method was developed for extracting organelle DNA from this carrageenophyte genus involving purification of nucleic acids by binding to hydroxylapatite. Plastid DNA from G. devoniensis and bisporangial Erythrodermis-like crusts was compared with that of G. devoniensis and G. crenulatus (Turner) J. Agardh from France and of G. furcellatus (C. Agardh) J. Agardh from Chile. Plastid genomes of all Gymnogongrus species and the Erythrodermis-like crust were approximately 175 kb long. A single 3.5-kb plasmid DNA species was found in G. devoniensis and the Erythrodermis-like bisporophyte but not in other samples. Digestion of plasted DNA with several restriction endonucleases produced identical patterns in G. devoniensis and the Erythrodermis-like bisporophyte from the same location, indicating clearly that these entities represent two phases of an uncoupled life history. These results were confirmed with heteologous probes. A restriction fragment length polymorphism was identified between two Nova Scotian G. devoniensis populations. There was no similarity in restriction patterns between G. devoniensis from Nova Scotia, G. devoniensis from France. G. crenulatus or G. furcellatus, suggesting that molecular taxonomic methods could be important in delineating members of this morphologically variable genus. Further study is necessary to determine whether either Nova Scotian G. devoniensis or French G. devoniensis corresponds to type populations of G. devoniensis from Devon, England.     

MORPHOLOGICAL AND MOLECULAR ANALYSIS OF THE ENDANGERED SPECIES PORPHYRA TENERA (BANGIALES,RHODOPHYTA)1

Porphyra tenera Kjellman, widely cultivated in nori farms before the development of artificial seeding, is currently listed as an endangered species in Japan. To confirm whether a wild‐collected gametophytic blade was P. tenera or the closely related species P. yezoensis Ueda, morphological observations and molecular analyses were made on the pure line HGT‐1 isolated from a wild blade. This pure line was identified as P. tenera based on detailed morphological features. Sequences of the nuclear internal transcribed spacer region 1 and the plastid RUBISCO spacer revealed that P. tenera HGT‐1 was clearly different from P. yezoensis f. narawaensis Miura, the main species cultivated in Japan. PCR‐RFLP analysis of the internal transcribed spacer region was found to be a convenient method for rapid discrimination between P. tenera and cultivated P. yezoensis. The restriction patterns generated by the endonucleases Dra I and Hae III were useful for differentiating between both gametophytic and conchocelis stages of P. tenera HGT‐1 and P. yezoensis f. narawaensis strains. Thus, PCR‐RFLP analysis will serve as a valuable tool for rapid species identification of cultivated Porphyra strains, culture collections of Porphyra strains for breeding material and conservation of biodiversity, and, as codominant cleaved amplified polymorphic sequence markers for interspecific hybridization products between P. tenera and P. yezoensis f. narawaensis. Under the same culture conditions, rate of blade length increase and the blade length‐to‐width ratio were lower in P. tenera HGT‐1 than in P. yezoensis f. narawaensis HG‐4. The HGT‐1 became mature more rapidly than HG‐4 and had thinner blades.     

DISTRIBUTION AND SYSTEMATICS OF THE FRESHWATER GENUS SIRODOTIA(BATRACHOSPERMALES,RHODOPHYTA) IN NORTH AMERICA1

Multivariate morphometrics and image analysis were used to determine the number of well-delineated infrageneric taxa of Sirodotia in North America. Three groupings were distinguished from 25 populations examined from Newfoundland and Quebec in the north to central Mexico in the south. These groupings were statistically related to 10 type specimens, and the following species were recognized: Sirodotia huillensis (Welwitsch ex W. et G. S. West) Skuja (syn. S. ateleia Skuja), S. suecica Kylin (syn. S. acuminata Skuja ex Flint and S. fennica Skuja), and S. tenuissima (Collins) Skuja ex Flint. These species are differentiated on the basis of whorl shape and degree of separation at maturity (S. suecica, rounded and appressed; S. huillensis and S. tenuissima, truncated apex and separated), the density of spermatangia (S. huillensis, dense clusters; S. suecica and S. tenuissima, sparsely aggregated), and the mode of germination of the gonimoblast initial (S. suecica and S. tenuissima, from the nonprotuberant side of the fertilized carpogonium; S. huillensis from the protuberant side). Sirodotia huillensis was found only in the desert-chaparral, whereas S. suecica and S. tenuissima occurred from south-temperate to boreal regions in cool (temperature 8–18° C), low ion (specific conductance 10–99 μS · cm^?1), and mildly acidic to neutral (pH 5.7–7.3) waters.     

BIOGEOGRAPHIC ANALYSIS OF THE ARMORED PLANKTONIC DINOFLAGELLATE CERATIUM IN THE NORTH ATLANTIC AND ADJACENT SEAS1

The distribution of the dinoflagellate genus Ceratium Schrank (Dinophyceae) in the North Atlantic and adjacent seas was studied by a combination of new observations on a large number of plankton samples collected from the northeastern Atlantic and North Sea, data from cruises off the east coast of North America and Caribbean Sea, and reports in the literature of the past 90 years. Seventy species were recorded, and their distribution was examined by several methods. Distribution maps were plotted for all species, and from these the ranges of temperature tolerance were derived. The 240 sets of data, which took the form of lists of species present in 5° latitude / longitude blocks obtained from the new work and the published material, were analyzed by clustering and ordination multivariate techniques using the programs Twinspan and Decorana. Analysis of the individual species showed that surface water temperature is the most important factor determining distribution and the number of species in a particular area. Warm water and /or low latitudes have many more species than cold waters and/or high latitudes. For example, at 5°N there are on average 23 species per block, whereas at 60° N there are only 8 species. On the basis of this work, the Ceratium species are divided into Group 1, Arctic-temperate species normally only found in water of less than 15°C; Group 2, cosmopolitan species, which are found virtually everywhere and are the species most likely to form blooms or “brown water”; Group 3, intermediate species, which extend into neither the coldest nor the warmest water; Group 4, temperate-tropical species, which have a lower temperature boundary of 5°–12° C; Group 5, warm-temperate-tropical species with a lower temperature boundary of 14°–15°C; and Group 6, tropical species, which are rarely found in water of less than 20° C. Analysis of the sample sites also confirmed the predominant influence of temperature, and the Atlantic Ocean was divided into four biogeographical zones of which the boundaries follow isotherms of surface water temperature. Zone 1 consists of the Arctic and subarctic area, with the southern boundary closely following the 10° mean annual temperature (MAT) line. Zone 2 is an intermediate or cold-temperate zone, of which the southern boundary follows the winter 10° C MAT isotherm or the similarly placed summer 15° isotherm. Zone 3 is the warm-temperate or subtropical zone, which is very broad. The southern boundary closely follows the 25°C summer isotherm. Zone 4 is the tropical zone, where water temperature is never likely to be much less than 23°C. These findings are discussed in relation to experimental work and environmental observations. We suggest that the genus Ceratium provides an excellent tool for defining ocean currents and temperature changes and may become of value in studies of global change.     

BIOGEOGRAPHY OF BATRACHOSPERMUM HELMENTOSUM (BATRACHOSPERMALES, RHODOPHYTA) IN NORTH AMERICA USING MOLECULAR AND MORPHOLOGICAL DATA

Over the course of 3 years (1997–1999), 72 stream sites were sampled for epilithic diatom communities. The analysis of these samples has led to the identification of over 325 species of diatoms. In addition to sampling the diatom community, selected physical and chemical parameters were recorded from each stream reach. These parameters included pH, specific conductance, current velocity, SRP, nitrate, silica, and total alkalinity. Canonical Correspondence Analysis (CCA) was used to identify influential environmental parameters and to assess the response of the diatom community to prominent anthropogenic inputs in the region (i.e. coal mine drainage, eutrophication). The initial analyses indicate that pH was the most influential environmental parameter along the first CCA axis. This shift was not unexpected, as acid mine drainage (AMD) in the region leads to a wide range of pH values (2.8–7.93). The highly acidic sites were characterized by species of the genus Eunotia (specifically E. exigua and E. steineckei ), Frustulia rhomboides , and Pinnularia subcapitata. Furthermore, Achnanthidium minutissimum was the most widely distributed of the diatom species encountered, being found at 94% of the sites sampled. Streams that fluctuated between acidic and circumneutral pH (termed "teeter-totter") had greater abundances of Brachysira vitrea than other streams in this survey. Further implications for the use of these diatom communities as biomonitoring tools and the distribution of assemblages within the Western Allegheny Plateau will be discussed.     

CARRAGEENANS BIOSYNTHESIZED BY CARPOSPOROPHYTES OF RED SEAWEEDS GIGARTINA SKOTTSBERGII (GIGARTINACEAE) AND GYMNOGONGRUS TORULOSUS (PHYLLOPHORACEAE)1

Carrageenans biosynthesized by gametophytic and tetrasporic plants of seaweeds belonging to the Gigartinaceae and Phyllophoraceae are different: gametophytes produce carrageenans of the kappa family, whereas lambda‐carrageenans are extracted from tetrasporophytes. For Gigartina skottsbergii Setchell and Gardner and Gymnogongrus torulosus Hooker et Harvey, mature cystocarps were isolated and carrageenans were extracted. Structural determination by methylation analysis, Fourier transform infrared spectroscopy, and ¹³C‐NMR spectroscopy showed that they were kappa/iota‐carrageenans. For the extract obtained from cystocarps of Gigartina skottsbergii with water at room temperature, the ratio kappa:iota was 1:0.30 and at 90° C was 1:0.43; significant amounts of precursors were also present. The extract obtained from cystocarps of Gymnogongrus torulosus at 90° C showed prevalence of iota‐carrageenans (ratio kappa:iota 1:1.21). These extracts are similar to the polysaccharides produced by gametophytes of these seaweeds. For Gigartina skottsbergii, it was possible to separate the pericarpic tissue from the carposporophyte. Thus, they were extracted separately, and the carrageenans isolated were studied as described before, obtaining similar conclusions. These results clearly show that whereas the carposporophytes are located inside the cystocarp, they produce carrageenans of the kappa family despite of being diploid cells.     

BIOGEOGRAPHYOF BATRACHOSPERMUM GELATINOSUM (BATRACHOSPERMALES,RHODOPHYTA) IN NORTH AMERICA BASED ON MOLECULAR AND MORPHOLOGICAL DATA1

Fifteen populations of the widespread fieshwater red alga Batrachospermum gelatinosum (L.) De Candolle were sampled throughout the geographic range in North America from central Alabama, U.S.A. (33° N), to Ellesmere Island, Northwest Territories (NWT), Canada (80° N). Analysis of ribosomal DNA internal transcribed spacer (ITS) 1 and 2 sequences yielded a parsimony tree with a large polytomy consisting of most populations plus a branch with one Nova Scotia and two NWT populations. The nucleotide variation, both within the polytomy and within the branch, was small (< 1%). The sequence divergence between the branch and polytomy was 3%. The lengths of the ITS 1 and 2 sequences of B. gelatinosum, 216–229 and 448–458 base pairs, respectively, fall within the very broad ranges reported for other red algae. The cluster analysis of 11 morphometric characteristics revealed three groupings of populations, partly based on geographic distribution. All tundra, eastern boreal forest, and mid-western hemlock-hardwood populations were in one grouping, whereas the deciduous forest, coastal plain, and eastern hemlock-hardwood populations were in a second. How ever, one deciduous forest population from Rhode Island, U.S.A. was unassociated. There was considerable overlap in morphometric characteristics among the three groupings. Based on this fact and the relatively small nucleotade variation in ITS sequences, we conclude that B. gelatinosum is a morphologically variable and geographically widespread species that is a valid taxonomic entity.     

MOLECULAR AND MORPHOLOGICAL CHARACTERIZATION OF TEN POLAR AND NEAR‐POLAR STRAINS WITHIN THE OSCILLATORIALES (CYANOBACTERIA)1

An approximately 1400‐bp region of the 16S rRNA gene was sequenced for 10 polar or near‐polar strains putatively placed in the Oscillatorialean genera Oscillatoria, Phormidium, and Lyngbya obtained from the University of Toronto Culture Collection to assess phylogenetic relationships. The strains were also examined for thylakoid structure and cell division type with TEM as well as traditional morphology with LM. Phylogenetic trees constructed using parsimony, distance, and maximum likelihood methods were similar in topology. If the original epithets applied to the sequenced strains (both polar and those from GenBank) were used, it was clear that taxa were not monophyletic. However, using the revised taxonomic system of Anagnostidis and Komárek, we were able to reassign these strains to their current correct taxa (species, genus, and family). When these assignments were made, it was determined that the molecular sequence data analyses were congruent with morphology and ultrastructure. Nine of the polar strains were found to be new species, and eight were described as such: Arthronema gygaxiana Casamatta et Johansen sp. nov., Pseudanabaena tremula Johansen et Casamatta sp. nov., Leptolyngbya angustata Casamatta et Johansen sp. nov., Phormidium lumbricale Johansen et Casamatta sp. nov., Microcoleus glaciei Johansen et Casamatta sp. nov., Microcoleus rushforthii Johansen et Casamatta sp. nov., Microcoleus antarcticus Casamatta et Johansen sp. nov., Microcoleus acremannii Casamatta et Johansen sp. nov. Some genera (Leptolyngbya and Microcoleus) were clearly not monophyletic and require future revision.     

GAYLIELLA GEN. NOV. IN THE TRIBE CERAMIEAE (CERAMIACEAE,RHODOPHYTA) BASED ON MOLECULAR AND MORPHOLOGICAL EVIDENCE1

On the basis of comparative morphology and phylogenetic analyses of rbcL and LSU rDNA sequence data, a new genus, Gayliella gen. nov., is proposed to accommodate the Ceramium flaccidum complex (C. flaccidum, C. byssoideum, C. gracillimum var. byssoideum, and C. taylorii), C. fimbriatum, and a previously undescribed species from Australia. C. transversale is reinstated and recognized as a distinct species. Through this study, G. flaccida (Kützing) comb. nov., G. transversalis (Collins et Hervey) comb. nov., G. fimbriata (Setchell et N. L. Gardner) comb. nov., G. taylorii comb. nov., G. mazoyerae sp. nov., and G. womersleyi sp. nov. are based on detailed comparative morphology. The species referred to as C. flaccidum and C. dawsonii from Brazil also belong to the new genus. Comparison of Gayliella with Ceramium shows that it differs from the latter by having an alternate branching pattern; three cortical initials per periaxial cell, of which the third is directed basipetally and divides horizontally; and unicellular rhizoids produced from periaxial cells. Our phylogenetic analyses of rbcL and LSU rDNA gene sequence data confirm that Gayliella gen. nov. represents a monophyletic clade distinct from most Ceramium species including the type species, C. virgatum. We also transfer C. recticorticum to the new genus Gayliella.     

COMPARATIVE MORPHOLOGY AND TAXONOMIC STATUS OF GRACILARIOPSIS (GRACILARIALES,RHODOPHYTA)1

The vegetative organization and reproductive development of Gracilariopsis lemaneiformis (Bory) Dawson, Acleto et Foldvik [including Gracilaria sjoestedtii Kylin] were investigated. Our observations on spermatangial development and post-fertilization features establish that Gracilariopsis Dawson is distinct at the generic level from Gracilaria Greville, and ice propose the resurrection of Gracilariopsis Dawson as a result. Spermatangial parent cells of Gracilariopsis are superficial, initiated in pairs or groups of three by concavo-convex longitudinal and transverse divisions. Each spermatangial parent cell cuts off a single, colorless spermatangium distally by a transverse division. The female reproductive apparatus consists of a supporting cell that bears a two-celled carpogonial branch flanked by two sterile branches, as in Gracilaria. Likewise, up to six sterile cells fuse with the carpogonium after fertilization to produce a primary fusion cell that generates the gonimoblasts; however, a secondary fusion cell is absent. Inner gonimoblast cells unite with cytologically modified cells of the inner pericarp by means of secondary pit-connections. Tubular nutritive cells are absent. The gonimoblast consists of a central sterile tissue interconnected throughout by secondary pit-connections surmounted by a fertile layer composed of carposporangia aligned in straight chains. The distribution of Gracilariopsis is extended to Western Europe.     

THE GENUS PIKEA (DUMONTIACEAE,RHODOPHYTA) IN ENGLAND AND THE NORTH PACIFIC: COMPARATIVE MORPHOLOGICAL,LIFE HISTORY,AND MOLECULAR STUDIES1

A Pikea species attributed to Pikea californica Harvey has been established in England since at least 1967. Previously, this species was believed to occur only in Japan and Pacific North America. Comparative morphological studies on field-collected material and cultured isolates from England, California, and Japan and analysis of organellar DNA restriction fragment length polymorphisms, detected using labeled organellar DNA as a non-radioactive probe, showed that English Pikea is conspecific with P. californica from California. Both populations consist of dioecious gametophytes with heteromorphic life histories involving crustose tetrasporophytes; 96% of organellar DNA bands were shared between interoceanic samples. A second dioecious species of Pikea, P. pinnata Setchell in Collins, Holden et Setchell, grows sympatrically with P. californica near San Francisco but can be distinguished by softer texture, more regular branching pattern, and elongate cystocarpic axes. Pikea pinnata and P. californica samples shared 49–50% of organellar DNA bands, consistent with their being distinct species. Herbarium specimens of P. robusta Abbott resemble P. pinnata in some morphological features but axes are much wider; P. robusta may represent a further, strictly sub-tidal species but fertile material is unknown. Pikea thalli from Japan, previously attributed to P. californica and described here as Pikea yoshizakii sp. nov., are monoecious and show a strikingly different type of life history. After fertilization, gonimoblast filaments grow outward through the cortex and form tetrasporangial nemathecia; released tetraspores develop directly into erect thalli. Tetrasporoblastic life histories are characteristic of certain members of the Phyllophoraceae but were previously unknown in the Dumontiaceae. Japanese P. yoshizakii shared 55 and 56% of organellar DNA bands with P. californica and P. pinnata, respectively; phylogenetic analysis indicated equally distant relationships to both species. Pikea yoshizakii or a closely similar species with the same life history occurs in southern California and Mexico.     

LIFE HISTORY AND HYBRIDIZATION STUDIES ON GIGARTINA STELLATA AND PETROCELIS CRUENTA (RHODOPHYTA) IN THE NORTH ATLANTIC1

Fourteen isolates of the crustose marine red alga Petrocelis cruenta J. Agardh from various localities in the British Isles, France (including the type locality), Spain and Portugal gave rise in culture to dioecious foliose plants identifiable as Gigartina stellata (Stackhouse) Batters although two isolates formed only sterile foliose blades. A total of 145 isolates of Gigartina stellata were also grown in culture from various localities in the U.S.A. (Maine), the British Isles, Iceland, Denmark, France, Spain and Portugal using both carpospores and vegetative blade apices. Two basic types of life history were found among these isolates: a direct-type life history involving the formation of further foliose plants from carpospores, some isolates of which also form spermatangia on the same papillae as the cystocarps; and a heteromorphic-type in which only crustose plants resembling Petrocelis cruenta are formed from carpospores. Only heteromorphic-type life histories were found from Spain and Portugal. Both life history types were found in plants from the U.S.A., the British Isles and northern France. Only direct-type life histories were found in plants from Iceland and Denmark. Some Petrocelis-like crusts derived from field collected G. stellata carpospores and Petrocelis crusts of hybrid progeny formed tetrasporangia in 8:16 h LD, 10° C but not in 8:16 h LD, 15° C; 16:8 h LD 10° C or 15° C; and 10:6.5:1: 6.5 h LDLD, 10° C. The spores thus formed were viable and produced normal dioecious male and female gametophytes. Short day and low temperature conditions appear necessary for tetrasporogenesis. The results from crossing experiments with 32 male and 27 female isolates of the heteromorphic-type derived from both G. stellata and P. cruenta showed that two virtually non-interbreeding populations with a high degree of geographical separation exist in the north-eastern Atlantic. Morphological differences between plants from each population are described. On the basis of culture and crossing results, Petrocelis cruenta J. Agardh is placed in synonymy with Gigartina stellata (Stackhouse in Withering) Batters.     

INFRAGENERIC TAXONOMY OF AHNFELTIA (AHNFELTIALES,RHODOPHYTA)1

Ahnfeltia plicata (Hudson) Fries, the type species of Ahnfeltia Fries, is reported to be a widespread alga which is an important source of agar. However, after characterizing type material and representative populations of A. plicata, we found that several taxa usually considered to be synonyms of A. plicata instead represent separate species with differing geographic distributions. Ahnfeltia plicata sensu stricto is characterized by the development of external carposporophytes on female sori that are present only on mature axes. Pseudocarposporophytes bearing monosporangia and cortical monosporangial sori occur on both young and old axes. Characteristic vegetative features include long medullary cells and cortical growth rings. Ahnfeltia plicata s.s. is circumpolar in both hemispheres and is harvested for agar in the White Sea. Ahnfeltia setacea (Kútzing) Schmitz from the Falkland Islands, Gymnogongrus comosus Kützing from Chile and Gymnogongrus filiformis Kützing from southern Argentina appear to be synonymous with A. plicata. Ahnfeltia fastigiata (Postels et Ruprecht) Makienko, from the North Pacific, differs from A. plicata principally in the development of female sori and carposporophytes only near apices; medullary cells are shorter and cortical growth rings, cortical monosporangia and pseudocarposporophytes are not formed. It is dioecious and its life history involves the development from carpospores of a crustose Porphyrodiscus tetrasporophyte. We suggest that A. plicata var. tobuchiensis Kanno et Matsubara, the principal form of Ahnfeltia utilized for agar production in the North Pacific, represents an unattached ecad of A. fastigiata. Ahnfeltia elongata Montagne, known only from Chile and with carposporophytes restricted to main axes, is more closely related to A. plicata than to A. fastigiata. Thalli are large and regularly dichotomous, with long medullary cells, and do not form pseudocarposporophytes or monosporangia.     

CRYPTIC DIVERSITY AND PHYLOGENETIC RELATIONSHIPS WITHIN THE MASTOCARPUS PAPILLATUS SPECIES COMPLEX (RHODOPHYTA,PHYLLOPHORACEAE)1

Mastocarpus papillatus (C. Agardh) Kütz. is a common intertidal red alga occurring along the west coast of North America from Baja California to Alaska. Sequencing of both the chloroplast‐encoded rbcL gene and the nuclear ribosomal internal transcribed spacer (ITS) regions of ～200 specimens from California to Alaska revealed that M. papillatus is actually a complex of at least five species. All five species have high bootstrap support in phylogenetic analyses of both genetic regions, and in the case of the ITS marker, the species also have distinctive patterns of indels. Three of the species are localized in the mid‐ to upper intertidal, whereas two of the species occur in the low intertidal. The species also have different geographic ranges that overlap in the Vancouver Island area of British Columbia. No distinctive, reliable morphological differences were observed among the species. Although a variety of names are available for species in the complex, it is not yet clear which name goes with which species. As part of the survey, I also sequenced other species of Mastocarpus in the northeast Pacific region, and I provide new distribution records for M. jardinii ( J. Agardh) J. A. West and for a nonpapillate and probably undescribed species of Mastocarpus.     

DISTRIBUTION AND SYSTEMATICS OF BATRACHOSPERMUM (BATRACHOSPERMALES,RHODOPHYTA) IN NORTH AMERICA. 3. SECTION SETACEA1

Six populations of Batirachospermum section Setacea from North America were compared to eight type specimens using multivariate morphometrics and image analysis. From this analysis, four species in this section were distinguished worldwide: B. atrum (Hudson) Hartley [syn. B. gallaei Sirodot]; B. orrthostichum Skuja, B. sertularina (Bory) Bory]; B. diatyches Entwisle; B. androinvolucrum sp. nov.; and B. puiggarianum Grunow in Wittrock et Nordstedt (syn. B. angolense Welwitsch ex West et West, B. nigrescens Welwitsch ex West et West). Two of these species were found in North America: B. atrum in California and Texas and B. androinvolucrum in British Columbia, Washington State, and Alabama. The new species, B. androinvolucrum, is distinguished by having spermatangia restricted to one-celled involucral bracts of the carpogonial branch.     

DISTRIBUTION AND SYSTEMATICS OF BATRACHOSPERMUM(BATRACHOSPERMALES,RHODOPHYTA) IN NORTH AMERICA. 6. SECTION TURFOSA1

Twenty-one populations of Batrachospermum section Turfosa from North America were compared to nine type and two historically important specimens using multivariate morphometrics and image analysis. The protologues of six other infrageneric taxa were also compared. From this analysis, six species are recognized worldwide: B, de-sikacharyi Sankaran, B. gombakense Kumano et Ratnasabapathy, B. keratophytum Bory de Saint-Vincent [syn. B. vagum var. keratophytum (Bory de Saint-Vincent) Sirodot, B. gulbenkianum Reis, and B. suevorum Kützing nom. Meg.], B. sinense Jao, B. turfosum Bory de Saint-Vincent [syn. B. vagum (Roth) C. Agardh and B. vagum var. undulato-pedicellatum Kumano et Watanabe], and B. vogesiacum F. G. Schultz ex Skuja [syn. B. vagum var. flagelliforme Sirodot, B. flagelliforme (Sirodot) Necchi], These species are distinguished on the basis of carposporophyte-bearing branch cortication, secondary fascicle development, monoecy or dioecy, presence of spermatangia on involucral filaments and monosporangia, and dimensions of trichogynes and carposporangia. Peripheral cortication has been previously used to separate species in this section, but we observed that this feature is quite widespread in the section. Presence of indeterminate gonimoblast filaments has been reported for some taxa in section Turfosa, but no such structures were seen in any of the specimens examined. Only B. keratophytum has been collected in North America, ranging from southwestern Greenland (64°N) to Louisiana (30° N).     

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.