Circumscription of Fulbrightiella gen. nov. and Sherwoodiella gen. nov., Two Novel Genera in the Calotrichaceae (Nostocales, Cyanobacteria).

Three novel strains in Calotrichaceae from tropical habitats were isolated and characterized with regard to their morphology, phylogenetic placement, and secondary structures of conserved domains in the 16S-23S internal transcribed spacer (ITS). The strains fell into two clades formerly identified as Calothrix from freshwater and brackish habitats. Based on both morphology and ecology, they differed from the type species of Calothrix, C. confervicola, which is marine, has wide trichomes with short cells, and narrows abruptly to a hyaline hair. The first clade grouped species with heteropolar filaments widened at the base and narrowed gradually toward the apex but not ending in a hair, with basal heterocytes that are formed in series as the apically placed heterocytes senesce; this clade is being named Fulbrightiella gen. nov., with two named species, F. bharadwajae sp. nov. and F. oahuensis sp. nov. The second clade was comprised of a single species with isopolar trichomes that are untapering as hormogonia, but which widen midfilament and taper toward both ends following growth. These trichomes develop pairs of heterocyte mid-filament, causing fragmentation into heteropolar trichomes with basal heterocytes and ends that taper, but not to a hair. This clade consists of a single species at present, Sherwoodiella mauiensis. With this action, four clades in the Calotrichaceae have been named: Macrochaete, Dulcicalothrix, Fulbrightiella, and Sherwoodiella. Calothrix sensu stricto is truly marine, morphologically distinct, and unsequenced; finding and sequencing the generitype for Calothrix remains as the most important and unfinished task in the revision of the Calotrichaceae.

When will taxonomic saturation be achieved? A case study in Nunduva and Kyrtuthrix (Rivulariaceae, Cyanobacteria).

A number of heterocytous, mat-forming, tapering cyanobacteria in Rivulariaceae have recently been observed in both the Atlantic and Pacific coasts in the rocky intertidal and supratidal zones. These belong to the genera Nunduva, Kyrtuthrix, and Phyllonema and have been the subject of several recent studies. Herein, two new species of Nunduva (N. komarkovae and N. sanagustinensis) and two new species of Kyrtuthrix (K. munecosensis and K. totonaca) are characterized and described from the coasts of Mexico. Genetic separation based on the 16S-23S ITS region was pronounced (>10% in all comparisons). Morphological differences between all existing species in these two genera were also observed, but the group is morphologically complex, and these taxa are considered pseudocryptic. Nunduva and Kyrtuthrix remain morphologically and phylogenetically separate even with the addition of new species. However, how long will this remain the case? Many new genera and species of cyanobacteria have recently been described. Will the taxonomy of cyanobacteria eventually become saturated? Will we start to see multiple populations for the same cryptic species, or will future taxonomists collapse multiple species into fewer species, or multiple genera into single genera. The description of even more Nunduva and Kyrtuthrix species causes us to pause and evaluate the future of cyanobacterial taxonomy. These same questions are faced by algal taxonomists studying other phyla, and the resolution may ultimately be similar.

Two new species of Phyllonema (Rivulariaceae, Cyanobacteria) with an emendation of the genus.

Two untapered, heterocytous species were observed and collected from the intertidal and supratidal zones of the Mexican coastline of the Pacific Ocean near Oaxaca and from the Gulf of Mexico. These populations were highly similar in morphology to the freshwater taxon Petalonema incrustans in the Scytonemataceae. However, 16S rRNA sequence data and phylogenetic analysis indicated that they were sister taxa to the epiphyllic, Brazilian species Phyllonema aveceniicola in the Rivulariaceae, described from culture material. While genetic identity between the two new species was high, they differed significantly in morphology, 16S rRNA gene sequence identity, and sequence and structure of the 16S-23S ITS region. Their morphology differed markedly from the generitype of the previously monotypic Phyllonema, which has tapered, heteropolar, single-false branched trichomes with very thin or absent sheath. The two new species, Phyllonema ansata and Phyllonema tangolundensis, described from both culture and environmental material, have untapered, isopolar, geminately false branched trichomes with thick, lamellated sheaths, differences so significant that the species would not be placed in Phyllonema without molecular corroboration. The morphological differences are so significant that a formal emendation of the genus is required. These taxa provide a challenge to algal taxonomy because the morphological differences are such that one would logically conclude that they represent different genera, but the phylogenetic evidence for including them all in the same genus is conclusive. This conclusion is counter to the current trend in algal taxonomy in which taxa with minor morphological differences have been repeatedly placed in separate genera based primarily upon DNA sequence evidence.

Macrochaete gen. nov. (Nostocales, Cyanobacteria), a taxon morphologically and molecularly distinct from Calothrix.

Historically, the genus Calothrix included all noncolonial, tapered, heterocytous filaments within the cyanobacteria. However, recent molecular phylogenies show that "Calothrix" defined in this sense represents five distinct clades. The type species of Calothrix is marine, with solitary basal heterocytes, no akinetes, and distal ends tapering abruptly into short hairs. We examined the morphology and phylogeny of 45 tapering cyanobacteria in the Rivulariaceae, including freshwater and marine representatives of both Calothrix (35 strains) and its sister taxon Rivularia (10 strains). The marine Calothrix fall into two lineages, but we lack the generitype and so cannot identify the clade corresponding to the type species. The freshwater and soil Calothrix fall into the C. parietina clade and are characterized by having a basal heterocyte, no akinetes, and gradual tapering-but not into a long hyaline hair. Macrochaete gen. nov. is a freshwater taxon sister to the Calothrix lineages but clearly separated from Rivularia. The species in this genus differ morphologically from Calothrix by their ability to produce two heteromorphic basal heterocytes and specific secondary structures of the 16S-23S ITS. An additional feature present in most species is the presence of a distal, long hyaline hair, but this character has incomplete penetrance due to its expression only under specific environmental conditions (low phosphate), and in one species appears to be lost. We recognize three species: M. psychrophila (type species) from cold environments (high mountains, Antarctica), M. santannae from wet walls of subtropical South America, and M. lichenoides, a phycobiont of lichens from Europe.

Phylogenetic complexities of the members of Rivulariaceae with the re-creation of the family Calotrichaceae and description of Dulcicalothrix necridiiformans gen nov., sp nov., and reclassification of Calothrix desertica.

A freshwater dwelling, tapering, heterocytous cyanobacterium (strain V13) was isolated from an oligotrophic pond in the Shrirampur taluka, Ahmednagar district of Maharashtra in India. Initial morphological examination indicated that strain V13 belonged to the genus Calothrix. Subsequent molecular and phylogenetic assessment based on 16S rRNA gene, led us to describe the freshwater/terrestrial clade of Calothrix strains without terminal hairs as a new genus Dulcicalothrix gen. nov., with the type species Dulcicalothrix necridiiformans sp. nov. (Strain V13) on the basis of the necridia forming ability of the strain. Also, the 16S-23S ITS secondary structure analysis clearly differentiated strain V13 from the other members of the clade. Past studies and the current state of knowledge makes it imperative to separate the groups Calothrix (marine/freshwater Calothrix), Macrochaete and Dulcicalothrix (freshwater/terrestrial Calothrix) into separate genera in accordance with the International Code of Nomenclature for Algae, Fungi and Plants. Robust phylogenetic evidence and previous reports strongly support the re-erection of the family Calotrichaceae distinct from the existing family Rivulariaceae.

CHARACTERIZATION OF A MARINE CYANOBACTERIUM THAT BORES INTO CARBONATES AND THE REDESCRIPTION OF THE GENUS MASTIGOCOLEUS(1).

A marine, filamentous, endolithic cyanobacterium, strain BC008, was obtained in pure culture and characterized using a polyphasic approach. BC008 could bore into calcium carbonate minerals (calcite, aragonite) and, weakly, into strontium carbonate (strontianite), but not into other carbonates, phosphates, sulfates, silicates, or oxides, including those of calcium. We describe procedures for its continued cultivation in an actively boring state. BC008 was developmentally complex: it displayed lateral, terminal, and intercalary heterocysts; true branching; trichome tapering; and motile hormogonia. It also displayed considerable morphological plasticity between boring and nonboring modes. Boring brought about a halving of trichome diameter, a marked decrease in the ratio of heterocysts to vegetative cells, and a significant preference for lateral versus terminal heterocyst development. The cytoplasm of vegetative cells was filled with 20 nm thick, nanocompartment-like structures of polyhedral appearance and of unknown function. BC008 was capable of complementary chromatic adaptation but did not produce sheath pigments. When boring, it conformed well morphologically to Lagerheim's (1886) description of Mastigocoleus testarum, one of the most common and pervasive bioerosive agents of marine carbonates. We propose strain BC008 as type strain for the species. Multigene (16S rRNA, nif  H, rbcL) phylogenies confirm that Mastigocoleus is a distinct, deeply branching genus of cyanobacteria that shares affinities and critical traits with two major taxonomic groups in the heterocystous clade (Nostocales and Stigonematales). We provide a revision of the genus and species descriptions based on our strain and findings.