MOLECULAR SYSTEMATICS OF CERAMIUM AND CENTROCERAS (CERAMIACEAE,RHODOPHYTA) FROM BRAZIL1

Morphological investigations identified 11 Ceramium Roth species, of the 18 previously reported from Brazil. Phylogenetic analyses of sequences of the chloroplast‐encoded rbcL gene confirmed the presence of seven of these species. Three other species are reported from Brazil for the first time. Ceramium affine Setchell & Gardner and C. filicula Harvey ex Womersley were previously known only from the Pacific Ocean (Mexico and Australia, respectively). A new species, C. fujianum Barros‐Barreto et Maggs sp. nov., is described here. Its general habit is similar to that of C. strictum sensu Harvey from Europe but it has one less periaxial cell than C. strictum; its cortical filament arrangement is closest to C. deslongchampsii Chauvin ex Duby, also from Europe, but whorled tetrasporangia partially covered by cortical cells differ strikingly from the naked protruding tetrasporangia of C. deslongchampsii. Ceramium species in which each periaxial cell cuts off transversely only a single basipetal cell formed a robust clade. The genus Ceramium as represented in Brazil is not monophyletic with respect to Centroceras Kützing and Corallophila Weber‐van Bosse; Ceramium nitens, which has axial cells completely covered by rounded cortical cells formed by acropetal and basipetal filaments, did not group with any Ceramium clade but was weakly allied to a species of Corallophila. All three Brazilian Centroceras sequences were attributed to a single species, C. clavulatum.     

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.     

CHARACTERIZATION OF SCHIZOSERIS CONDENSATA,SCHIZOSERIDEAE TRIB. NOV. (DELESSERIACEAE,RHODOPHYTA)1

The Myriogramme group of Kylin contains two distinct clusters of genera that merit recognition at the tribal level. We previously established the tribe Myriogrammeae, and in this paper we erect the Schizoserideae based on a study of the type species of Schizoseris, S. laciniata (=S. condensata), from the southern hemisphere. The Schizoserideae is characterized by 1) marginal and diffuse intercalary meristems; 2) nuclei initially arranged in a plate in the median plane in meristematic and mature cells; 3) chloroplasts one to few, lobed or dissected; 4) microscopic veins absent; 5) procarps scattered, formed singly on either side of the blade with cover cells absent and consisting of a one- to two-celled lateral sterile group, a one- to two-celled basal sterile group, and a four-celled carpogonial branch in which the trichogyne passes beneath the lateral sterile group and emerges anterior to it; 6) auxiliary cell diploidized by a connecting cell cut off posteriolaterally from the fertilized carpogonium; 7) gonimoblast initial cut off laterally from one side of the auxiliary cell and giving rise to unilaterally branched gonimoblast filaments bearing carposporangia in branched chains; 8) gonimoblast fusion cell highly branched, candelabra-like, incorporating all but the basalmost cells of the carposporangial chains and radiating through the central cells in the floor of the cystocarp; 9) spermatangial and tetrasporangial sori formed from surface cells in both monostromatic and polystromatic portions on both sides of the blade; and 10) tetrasporangia formed primarily from cortical rather than from central cells. The Schizoserideae presently includes Schizoseris Kylin, Neuroglossum Kützing, Abroteia J. Agardh, and Polycoryne Skottsberg in Kylin and Skottsberg.     

PROPOSAL OF THE EUPTILOTEAE HOMMERSAND ET FREDERICQ,TRIB. NOV. AND TRANSFER OF SOME SOUTHERN HEMISPHERE PTILOTEAE TO THE CALLITHAMNIEAE (CERAMIACEAE,RHODOPHYTA)1

Morphological and molecular studies demonstrate that the tribe Ptiloteae (Ceramiaceae, Ceramiales) is polyphyletic. The Ptiloteae, sensu stricto, occur only in the Northern Hemisphere and all Southern Hemisphere representatives belong in other tribes. Three genera (Euptilota, Seirospora, and Sciurothamnion) are transferred to the Euptiloteae Hommersand et Fredericq, trib. nov., and the Callithamnieae is revised to include three Ptilota‐like genera, Georgiella, Falklandiella, and Diapse, and two new genera. Heteroptilon Hommersand, gen. nov. is erected to receive Euptilota pappeana Kützing 1849 and Aglaothamnion rigidulum De Clerck, Bolton, Anderson et Coppejans 2004 from South Africa, and Aristoptilon Hommersand et W. A. Nelson, gen. nov. is established to receive Euptilota mooreana Lindauer 1949 from New Zealand. The principal difference between the Euptiloteae and the Callithamnieae is seen in the earliest stages after fertilization. The fertilized carpogonium enlarges and forms a pair of tube‐like protuberances directed toward the auxiliary cells that are cut off as connecting cells in the Euptiloteae, whereas in the Callithamnieae the carpogonium usually divides into two cells, each of which cuts off a small connecting cell that fuses with an adjacent enlarging auxiliary cell. Nuclei are terminal in spermatangia of the Euptiloteae, subtended by mucilaginous vesicles, and are medial in the Callithamnieae situated between apical and basal vesicles. The Euptiloteae and Callithamnieae (including the Ptilota‐like members) are each strongly supported in maximum‐likelihood tree topologies resulting from analyses of combined 18S rDNA, 28S rDNA, 16S rDNA, and rbcL data sets. Their sister relationship is also well supported.     

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.     

A revision of the genus Cryptonemia (Halymeniaceae,Rhodophyta) in Bermuda,western Atlantic Ocean,including five new species and C. bermudensis (Collins & M. Howe) comb. nov.

Cryptonemia specimens collected in Bermuda over the past two decades were analysed using gene sequences encoding the large subunit of the nuclear ribosomal DNA and the large subunit of RuBisCO as genetic markers to elucidate their phylogenetic positions. They were additionally subjected to morphological assessment and compared with historical collections from the islands. Six species are presently found in the flora including C. bermudensis comb. nov., based on Halymenia bermudensis, and the following five new species: C. abyssalis, C. antricola, C. atrocostalis, C. lacunicola and C. perparva. Of the eight species known in the western Atlantic flora prior to this study, none is found in Bermuda. Specimens reported in the islands in the 1900s attributed to C. crenulata and C. luxurians are representative of the new species, C. antricola and C. atrocostalis, respectively.     

ITS1 sequences of type specimens of Gigartina and Sarcothalia and their significance for the classification of South African Gigartinaceae (Gigartinales,Rhodophyta)

Uncertainties about the identity of type specimens of red algae have frequently led to taxonomic and nomenclatural confusion. A procedure for extracting PCR-amplifiable DNA from formalin-fixed material and herbarium specimens was used to investigate the taxonomic status of several South African Gigartinaceae. We compared nucleotide sequences in the internal transcribed spacer 1 (ITS1) region in type specimens and other historically important collections presently referred to Gigartina bracteata, G. radula and G. stiriata. The following opinions are supported: (1) Fucus bracteatus S.G. Gmelin, Chondrodictyon capense Kützing and Iridaea clathrata Decaisne represent a single species of South African Gigartina (G. bracteata (S.G. Gmelin) Setchell & Gardner) in which the disintegration of tetrasporangial sori results in a reticulate thallus. (2) Mastocarpus polycarpus Kützing, M. incrassatus Kützing and Iridaea lapathifolia Kützing represent a single species of South African Gigartina (G. polycarpa (Kützing) Setchell & Gardner) that has often, but erroneously, been called G. radula (Esper) J. Agardh. (3) Mastocarpus verrucosus Kützing is a later heterotypic synonym of Iridaea papillosa Bory (Sarcothalia papillosa (Bory) Leister) and was based on material that probably came from southern South America rather than from South Africa, the provenance given by Kützing. (4) Fucus stiriatus Turner and Sphaerococcus burmannii C. Agardh represent a single species of South African Sarcothalia (S. stiriata (Turner) Leister).     

Two new species of Gelidium (Rhodophyta, Gelidiales), Gelidium tenuifolium and Gelidium koshikianum, from Japan

Two new marine red algae, Gelidium tenuifolium sp. nov. and Gelidium koshikianum sp. nov. (Gelidiales, Gelidiaceae) are described from Japan. Gelidium tenuifolium with large‐sized thalli (up to 30 cm tall) is distinguished from other species with such thalIi by the production of wide, flattened and thin branches (up to 2 mm wide and 60–80 urn thick), the presence of an apical depression and simple determinate branches. Gelidium koshikianum with middle‐sized thalli (5–8 cm tall) is distinguished from other species with such thalli by having wide axes (up to 2.5 mm wide) and short (2.0–3.2 mm), unbranched, second‐ and third‐order branches issuing at short intervals (0.8–1.4 mm). In phylogenetic analyses of rbcL sequences, four Gelidium species that are chiefly distributed in Japan including G. tenuifolium were clustered together with 99% bootstrap value (Japanese Gelidium‐complex clade). Gelidium linoides Kützing came to the position of the sister group to G. tenuifolium with 99% bootstrap value. There were four substitutions (0.3% divergence) between G. linoides and G. tenuifolium sequences. Gelidium koshikianum and Gelidium allanii Chapman were clustered together with 100% bootstrap value and they came to the position of the sister group to the Japanese GeIidium‐complex clade with 83% bootstrap value. There were six substitutions (0.4% divergence) between G. koshikianum and G. allanii sequences.     

MORPHOLOGY AND DEVELOPMENT OF AHNFELTIA PLICATA (RHODOPHYTA): PROPOSAL OF AHNFELTIALES ORD. NOV.1

Ahnfeltia plicata (Hudson) Fries, the type species of Ahnfeltia Fries, is currently assigned to the Phyllophoraceae (Gigartinales). Several morphological and biochemical characters distance A. plicata from the Phyllophoraceae but, because sexual reproduction has never been demonstrated, an alternative placement has not been possible. A. plicata now is shown to have a heteromorphic sexual life history. Erect branched gametophytes are dioecious. In male sori, spermatangia are cut off transversely from spermatangial mother cells. Female sori form numerous terminal sessile carpogonia. Following fertilization, several zygotes in each sorus fuse facultatively with undifferentiated intercalary cells of the female sorus and cut off gonimoblast initials obliquely outwards. These initials give rise to branching gonimoblast filaments that fuse with apical and intercalary female sorus cells and with each other, then grow radially outward in the compound external carposporophyte and terminate in carposporangia. Carpospores develop in culture into crustose tetrasporophytes identical to Porphyrodiscus simulans Batters. Field-collected P. simulans tetraspores grew into erect A. plicata axes. Tetrasporangia are formed by division and enlargement of crust apical cells followed by sequential enlargement and maturation of tetrasporocytes in an erosive process. Monosporangia are formed in sori on male gametophytes. Pit plugs of both gametophyte and tetrasporophyte phases consist of naked plug cores without cap layers of membranes. Gametophytes exhibit both cell fusions and secondary pit connections whereas tetrasporophytes form cell fusions but lack secondary pit connections. On the basis of the unique female and postfertilization reproductive development and in conjunction with the pit plug structure which is unique among florideophytes, the order Ahnfeltiales, containing the family Ahnfeltiaceae, is proposed.     

CRYPTIC SPECIES OF SCENEDESMUS (CHLOROPHYTA) FROM DESERT SOIL COMMUNITIES OF WESTERN NORTH AMERICA1

Nine isolates of unicellular green algae were obtained from six geographically separate desert microbiotic crust communities in western North America. Microscopically, eight isolates strongly resembled unicellular forms of Scenedesmus obliquus (Turpin) Kützing. They are oval or crescent shaped, often flattened on one side, with knobby cell apices. SEM indicated a lack of wall ornamentation. Fine filaments connecting cells pole to pole were observed in some isolates, as previously documented in Scenedesmus (Dactylococcus) dissociatus and S. obliquus. The ninth isolate was spherical, without knobby apices or connections between cells, and was similar to unicellular forms that were originally classified as species of Chlorella (Scenedesmus vacuolatus and S. rubescens). None of the isolates formed coenobia in liquid culture. Phylogenetic analysis of the 18S rRNA gene placed all desert isolates in the genus Scenedesmus, separating them into two or three weakly resolved groups along with published sequences of other Scenedesmus isolates. Phylogenetic analyses of the internal transcribed spacer region revealed well‐supported lineages of desert algae that were unsupported with 18S data alone. The eight S. obliquus‐like desert strains formed two distinct clades that excluded the S. obliquus strains from geographically widespread nondesert habitats. The ninth strain was outside of the S. obliquus group, associated with S. raciborskii and S. pectinatus. These results demonstrate three lineages of Scenedesmus from desert soils and provide robust support for the presence of cryptic species in S. obliquus, a morphospecies that is said to have a cosmopolitan distribution. Three new species of Scenedesmus are described.     

PHYLOGENY AND SYSTEMATICS OF EUGLENA (EUGLENACEAE) SPECIES WITH AXIAL,STELLATE CHLOROPLASTS BASED ON MORPHOLOGICAL AND MOLECULAR DATA—NEW TAXA,EMENDED DIAGNOSES,AND EPITYPIFICATIONS1

Morphological and molecular studies, as well as original literature reexamination, necessitate establishment of five Euglena species with a single axial, stellate chloroplast [Euglena viridis (O. F. Müller) Ehrenberg 1830 , Euglena pseudoviridis  Chadefaud 1937 , Euglena stellata  Mainx 1926 , Euglena pseudostellata sp. nov., and Euglena cantabrica  Pringsheim 1956 ], three species with two chloroplasts (Euglena geniculata Dujardin ex Schmitz 1884 , Euglena chadefaudii  Bourrelly 1951 , and Euglena pseudochadefaudii sp. nov.), and one species with three chloroplasts (Euglena tristella  Chu 1946 ). The primary morphological features, allowing distinction of the considered species are the presence and the shape of mucocysts, as well as the number of chloroplasts. Spherical mucocysts occur in E. cantabrica and E. geniculata, while spindle‐shaped mucocysts are present in E. stellata, E. pseudostellata, E. chadefaudii, E. pseudochadefaudii, and E. tristella. No mucocysts are observed in E. viridis and E. pseudoviridis. Two new species (E. pseudochadefaudii sp. nov. and E. pseudostellata sp. nov.) differ from the respective species, E. chadefaudii and E. stellata, only at the molecular level. Molecular signatures and characteristic sequences are designated for nine distinguished species. Emended diagnoses for all and delimitation of epitypes for seven species (except E. viridis and E. tristella) are proposed.     

NEW GENERA AND SPECIES OF CERAMIACEAE (RHODOPHYTA) FROM THE SOUTHEASTERN UNITED STATES1

Five Ceramiaceae (Rhodophyta) are reported from the offshore waters of the southeastern, warm temperate coast of the United States. These include two new monotypic genera, Calliclavula trifurcate Schneider in the Griffithsieae and Nwynea grandispora Searles in the Sphondylothamnieae, and three new species, Callithamniella silvae Searles, Ptilothamnion occidentale Searles, and Lejolisia exposita Schneider et Searles.     

Neosiphonia flavimarina gen. et sp. nov. with a taxonomic reassessment of the genus Polysiphonia (Rhodomelaceae, Rhodophyta)

Vegetative and reproductive development of Neosiphonia flavimarina gen. et sp. nov. (Rhodomelaceae, Ceramiales) from Bangpo on the western coast of Korea was investigated. This species is superficially similar to Polysiphonia, but differs distinctly from the latter in vegetative and reproductive structures. The plants attach by a solid disk composed of a dense cluster of rhizoids cut off from the pericentral cell wall, and bear erect indeterminate branches producing the lateral-branch initials from successive segments in a spiral arrangement. The procarps have a three-celled carpo-gonial branch. Spermatangial branches are formed on a primary branch of the trichoblasts, terminating in a single or occasionally two large, sterile cells. Tetra-sporangia are produced from the second pericentral cell adjacent to the trichoblast basal cell on indeterminate branches, and arranged spirally. Comparing several taxonomic characters among related genera, Neosiphonia occupies an independent phylogenetic position from Polysiphonia and leads to the conclusion that the genus may have a strong link with Fernandosiphonia which has a unilateral branching system. Relevant nomenclatural changes for several Polysiphonia species are also proposed.     

A phylogenetic re‐appraisal of the family Liagoraceae sensu lato (Nemaliales,Rhodophyta) based on sequence analyses of two plastid genes and postfertilization development

The marine red algal family Liagoraceae sensu lato is shown to be polyphyletic based on analyses of a combined rbcL and psaA data set and the pattern of carposporophyte development. Fifteen of eighteen genera analyzed formed a monophyletic lineage that included the genus Liagora. Nemalion did not cluster with Liagoraceae sensu stricto, and Nemaliaceae is reinstated, characterized morphologically by the formation of the primary gonimolobes by longitudinal divisions of the gonimoblast initial. Yamadaella and Liagoropsis, previously placed in the Dermonemataceae, are shown to be independent lineages and are recognized as two new families Yamadaellaceae and Liagoropsidaceae. Yamadaellaceae is characterized by two gonimoblast initials cut off bilaterally from the fertilized carpogonium and diffusely spreading gonimoblast filaments. Liagoropsidaceae is characterized by at least three gonimoblast initials cut off by longitudinal septa from the fertilized carpogonium. In contrast, Liagoraceae sensu stricto is characterized by a single gonimoblast initial cut off transversely or diagonally from the fertilized carpogonium. Reproductive features, such as diffuse gonimoblasts and unfused carpogonial branches following postfertilization, appear to have evolved on more than one occasion in the Nemaliales and are therefore not taxonomically diagnostic at the family level, although they may be useful in recognizing genera.     

Two new species in the Chaetoceros socialis complex (Bacillariophyta): C. sporotruncatus and C. dichatoensis,and characterization of its relatives,C. radicans and C. cinctus

The diatom genus Chaetoceros is one of the most abundant and diverse phytoplankton in marine and brackish waters worldwide. Within this genus, Chaetoceros socialis has been cited as one of the most common species. However, recent studies from different geographic areas have shown the presence of pseudo‐cryptic diversity within the C. socialis complex. Members of this complex are characterized by curved chains (primary colonies) aggregating into globular clusters, where one of the four setae of each cell curves toward the center of the cluster and the other three orient outwards. New light and electron microscopy observations as well as molecular data on marine planktonic diatoms from the coastal waters off Chile revealed the presence of two new species, Chaetoceros sporotruncatus sp. nov. and C. dichatoensis. sp. nov. belonging to the C. socialis complex. The two new species are similar to other members of the complex (i.e., C. socialis and C. gelidus) in the primary and secondary structure of the colony, the orientation pattern of the setae, and the valve ultrastructure. The only morphological characters that can be used to differentiate the species of this complex are aspects related to resting spore morphology. The two newly described species are closely related to each other and form a sister clade to C. gelidus in molecular phylogenies. We also provide a phylogenetic status along with the morphological characterization of C. radicans and C. cintus, which are genetically related to the C. socialis complex.     

THE GENUS HYPOGLOSSUM KÜTZING (DELESSERIACEAE,RHODOPHYTA) IN THE TROPICAL WESTERN ATLANTIC,INCLUDING H. ANOMALUM SP. NOV.1

Observations are made on the occurrence and distribution of the red algal genus Hypoglossum Kützing (Delesseriaceae, Ceramiales) in the tropical western Atlantic. In addition to the type of the genus, H. hypoglossoides (Stackh.) Coll. & Herv., three other species are reported: H. anomalum sp. nov., H. involvens (Harv.) J. Ag., and H. tenuifolium (Harv.) J. Ag. A key is presented to distinguish these four species. The newly described species, H. anomalum, is like other species in the genus in that its branches arise endogenously from the primary axial row but it is unique in that the branches emerge from the parent blade at some point between the midline and the margin of the blade. The new species is reported from Puerto Rico and Florida.     

MORPHOLOGY,MOLECULAR PHYLOGENY AND TAXONOMY OF TWO NEW SPECIES OF PLEODORINA (VOLVOCEAE,CHLOROPHYCEAE)1

The volvocacean genus Pleodorina has been morphologically characterized as having small somatic cells in spheroidal colonies and anisogamous sexual reproduction with sperm packets. In this study we examined two new species that can be assigned to the genus Pleodorina based on morphology: P. starrii H. Nozaki et al. sp. nov. and P. thompsonii F. D. Ott et al. sp. nov. P. starrii was collected from Japan and had 32‐ or 64‐celled colonies with anterior somatic cells and spheroidal individual cellular sheaths that were weakly attached to each other within the colonial envelope. P. thompsonii from Texas (USA) exhibited four or 12 somatic cells in the anterior pole of 16‐ or 32‐celled colonies, respectively, and had a single large pyrenoid in the chloroplast of mature reproductive cells. The chloroplast multigene phylogeny placed P. starrii and P. indica (Iyenger) H. Nozaki in a clade that was robustly separated from the type species P. californica Shaw and P. japonica H. Nozaki. Pleodorina thompsonii was resolved as a basal branch within a large monophyletic group (Eudorina group) composed of Eudorina, Pleodorina and Volvox (excluding section Volvox). Thus, Pleodorina was found among three separate lineages within the Eudorina group in which Eudorina and Volvox were also resolved as nonmonophyletic. The DNA sequences from additional species/strains as well as recognition of morphological attributes that characterize the monophyletic groups within the Eudorina group are needed to construct a natural generic classification within these members of the Volvocaceae.     

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.     

NEW ALGAL SPECIES RECORD FOR COLORADO1

Colonies of the alga Nostoc parmeloides Kützing 1843, containing midge larvae, were collected from Bellows Creek in the southwestern mountains of Colorado near Creede, Mineral County, on February 27, 1970. G. W. Prescott verified the algal identification and confirmed it as a new species record for Colorado. The Nostoc-midge combination has been definitely identified in 8 other Colorado mountain streams. The streams are located in various sections of Colorado above 2333 m; distribution may be statewide.     

SYSTEMATIC REVISION OF THE GENERA LIAGORA AND IZZIELLA (LIAGORACEAE,RHODOPHYTA) FROM TAIWAN BASED ON MOLECULAR ANALYSES AND CARPOSPOROPHYTE DEVELOPMENT,WITH THE DESCRIPTION OF TWO NEW SPECIES1

Some Liagora and Izziella distributed in Taiwan display a wide range of morphological variation and can be difficult to distinguish. To clarify species concepts, we applied DNA sequence analyses and examined carposporophyte development in detail. These studies revealed two new species, which are described herein as Izziella hommersandii sp. nov. and Izziella kuroshioensis sp. nov. I. kuroshioensis superficially resembles Izziella formosana and Izziella orientalis in that its involucral filaments subtend rather than surround the lower portion of the gonimoblast mass (= Izziella type) and a fusion cell is formed from cells of the carpogonial branch, but it can be separated by differences in the cell numbers and branching pattern of the involucral filaments, as well as thallus morphology. In contrast to other species that also bear short lateral branchlets, I. hommersandii is unique in possessing a mixture of short and long involucral filaments, a phenomenon not reported before. The length of the involucral filaments is species specific among species of Izziella and contrasts to the behavior of the involucral filaments after fertilization in species such as “Liagora”setchellii [= Titanophycus setchellii comb. nov.], in which the filaments completely envelop the gonimoblast. In addition, the cells of the carpogonial branch in Titanophycus do not fuse after fertilization to form a fusion cell. Thus, a combination of characters with respect to the behavior of the carpogonial branch and the involucral filaments after fertilization is very useful for delineating species boundaries in Izziella and for separating Titanophycus from Izziella and Liagora.