Max H. Hommersand

SYSTEMATICS OF THE DELESSERIACEAE (CERAMIALES, RHODOPHYTA) BASED ON LARGE SUBUNIT rDNA AND rbcL SEQUENCES, INCLUDING THE PHYCODRYOIDEAE, SUBFAM. NOV.

The present classification of the Delesseriaceae retains the essential features of Kylins system, which recognizes two subfamilies Delesserioideae and Nitophylloideae and a series of “groups” or tribes. In this study we test the Kylin system based on phylogenetic parsimony and distance analyses inferred from two molecular data sets and morphological evidence. A set of 72 delesseriacean and 7 additional taxa in the order Ceramiales was sequenced in the large subunit rDNA and rbcL analyses. Three large clades were identified in both the separate and combined data sets, one of which corresponds to the Delesserioideae, one to a narrowly circumscribed Nitophylloideae, and one to the Phycodryoideae, subfam. nov., comprising the remainder of the Nitophylloideae sensu Kylin. Two additional trees inferred from rbcL sequences are included to provide broader coverage of relationships among some Delesserioideae and Phycodryoideae. Belonging to the Delesserioideae are the Caloglosseae with Caloglossa; an expanded Hemineureae that includes Hemineura, Patulophycus, Marionella, Laingia, Botryocarpa, and Pseudophycodrys; the Delesserieae with Delesseria and Membranoptera; the Apoglosseae with Apoglossum and a group of southern hemisphere species presently placed in Delesseria that belong in Paraglossum; the Hypoglosseae with Hypoglossum, Branchioglossum, Zellera, and Bartoniella; and the Grinnellieae with Grinnellia. The revised Nitophylloideae contains the Nitophylleae with Nitophyllum, Valeriemaya, Polyneuropsis, and Calonitophyllum and the Martensieae with Opephyllum and Martensia. A new subfamily, Phycodryoideae, is proposed to include the Phycodryeae with Phycodrys, Polyneura, Nienburgia, Cladodonta, Heterodoxia, and Womersleya; the Cryptopleureae with Cryptopleura, Hymenena, Acrosorium, and Botryoglossum; the Myriogrammeae with Myriogramme and Haraldiophyllum; and the Schizoserideae with Schizoseris, Neuroglossum, Drachiella, Abroteia, and species from South America placed in Platyclinia. This research promotes the correlation of molecular and morphological phylogenies.

Assessing red algal supraordinal diversity and taxonomy in the context of contemporary systematic data

The wondrously diverse eukaryotes that constitute the red algae have been the focus of numerous recent molecular surveys and remain a rich source of undescribed and little known species for the traditional taxonomist. Molecular studies place the red algae in the kingdom Plantae; however, supraordinal classification has been largely confined to debate on subclass vs. class level status for the two recognized subgroups, one of which is widely acknowledged as paraphyletic. This narrow focus has generally masked the extent to which red algal classification needs modification. We provide a comprehensive review of the literature pertaining to the antiquity, diversity, and systematics of the red algae and propose a contemporary classification based on recent and traditional evidence.

SOLVING TAXONOMIC AND NOMENCLATURAL PROBLEMS IN PACIFIC GIGARTINACEAE (RHODOPHYTA) USING DNA FROM TYPE MATERIAL

Molecular data obtained by a procedure for extracting PCR‐amplifiable nuclear and chloroplast DNA from old and formalin‐fixed red algal herbarium specimens were used to elucidate problems in the systematics of Pacific Gigartinaceae. Correspondence between nucleotide sequences of the internal transcribed spacer 1 region or the RUBISCO spacer from type specimens and modern collections supports the following conclusions. (1) The type of Fucus cordatus Turner, now Iridaea cordata (Turner) Bory, came from the southern hemisphere (probably from Isla de los Estados, Argentina) rather than from Banks Island, B.C., Canada. (2) The type of Iridaea heterocarpa P. et R. [Mazzaella heterocarpa (P. et R.) Fred.] represents the tetrasporangial phase of a species of Chondrus, possibly C. crispus Stackh. (3) The types of Iridaea lilacina P. et R., I. phyllocarpa P. et R., and Iridophycus furcatum S. et G. represent a single species from Alaska, Mazzaella phyllocarpa (P. et R.) Perest., currently but incorrectly called M. heterocarpa. (4) The type of Iridophycus oregonum Doty represents the tetrasporangial phase of the species from southern Alaska to southern California known incorrectly as M. heterocarpa. (5) Mazzaella splendens (S. et G.) Fred. is more closely related to M. linearis (S. et G.) Fred. than it is to M. flaccida (S. et G.) Fred. (6) Iridophycus coriaceum S. et G. is conspecific with M. splendens, whereas Rhodoglossum coriaceum E.Y. Dawson is an independent species: Mazzaella coriacea (E.Y. Dawson) Hughey. (7) Iridaea cornucopiae P. et R. is conspecific with Mazzaella laminarioides (Bory) Fred., and the type probably came from Chile rather than from the North Pacific. (8) Plants attributed to Iridaea cornucopiae in Pacific North America are referable to Mazzaella parksii (S. et G.) comb. nov. (9) Rhodoglossum parvum G. M. Smith et Hollenb. is an independent species: Mazzaella parva (G. M. Smith et Hollenb.) comb. nov. (10) Grateloupia squarrulosa S. et G., Grateloupia johnstonii S. et G., and Gigartina pectinata E.Y. Dawson represent a single species: Chondracanthus squarrulosus (S. et G.) comb. nov.

PROPOSAL OF THE GRACILARIALES ORD. NOV. (RHODOPHYTA) BASED ON AN ANALYSIS OF THE REPRODUCTIVE DEVELOPMENT OF GRACILARIA VERRUCOSA

The mode of division of vegetative cells, formation of spermatangial parent cells, initiation of the carpogonial branch apparatus, and formation of tetrasporangial initials are homologous developmental processes that are documented for the first time in the type species of the economically important family Gracilariaceae, Gracilaria verrucosa (Hudson) Papenfuss from the British Isles. G. verrucosa is characterized by a supporting cell of intercalary origin that bears a 2‐celled carpogonial branch flanked by two sterile branches, direct fusion of cells of sterile branches onto the carpogonium, formation of an extensive carpogonial fusion cell through the incorporation of additional gametophytic cells prior to gonimoblast initiation, gonimoblast initials produced from fusion cell lobes, schizogenous development of the cytocarp cavity, inner gonimoblast cells producing tubular nutritive cells that fuse with cells of the pericarp or floor of the cystocarp, absence of cytologically modified tissue in the floor of the cystocarp, and carposporangial initials produced in clusters or irregular chains. Spermatangial parent cells are generated in flaments from intercalary cortical cells that line an intercellular space forming a ‘pit’ or ‘conceptacle’. Tetrasporangial initials are transformed from terminal cells derived through division of an outer cortical cell. Tetrasporangia are cruciately divided.

Comparative morphology and taxonomic status of Gracilariopsis (Gracilariales, Rhodophyta)

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.

A MOLECULAR PHYLOGENY OF THE GELIDIALES (RHODOPHYTA) BASED ON ANALYSIS OF PLASTID rbcL NUCLEOTIDE SEQUENCES1

Parsimony analyses of rbcL nucleotide. sequences were used to develop hypotheses of relationships among taxa in the taxonomically difficult order Celidiales including species from seven currently recognized genera: Capreolia, Gelidiella, Gelidium, Onikusa, Pterocladia, Ptilophora, and Suhria. Nucleotide. sequences of rbcL from red algae are variable and provide a large number of informative characters for phylogenetic analysis, yet the absence of insertion/deletion mutations allows for the unambiguous alignment of sequences. Species were resolved into 10 well‐.supported major clades representing genera and species complexes. The topological positions of these 10 clades within trees are also well supported and indicate that Gelidium and Pterocladia as currently circumscribed are not monophyletic. These results call for a revision of the classification of the Gelidiales.

New perspectives in the taxonomy of the Gigartinaceae (Gigartinales, Rhodophyta)

A revised description of the Gigartinaceae is provided, together with a key and short diagnosis of each genus and a list of the species examined. New combinations have been proposed where appropriate. Distinguishing cystocarp and tetrasporangial characters useful for separating genera are illustrated, and the distribution of the genera is shown on a world map.

The molecular systematics of some agar- and carrageenan-containing marine red algae based on rbcL sequence analysis

A hypothesis of phylogenetic relationships inferred by parsimony analysis of plastid-encoded rbcL sequences is presented for red algae containing agar- and carrageenan-like phycocolloids; rbcL encodes the large subunit of ribulose 1,5 bisphosphate carboxylase/oxygenase. Previous studies have shown that Floridean families that contain sulfated galactans as cell wall components are resolved as a monophyletic clade sister to the agarophyte order Ahnfeltiales. Families that have been identified as containing kappa-type carrageenans, often in addition to lambda-type carrageenans, are resolved in three clades: (1) a complex containing the families Solieriaceae, Cystocloniaceae, Hypneaceae, Caulacanthaceae, Tichocarpaceae, Furcellariaceae, and the genera Turnerella and Opuntiella; (2) the Gigartinaceae and Phyllophoraceae, and (3) the genus Endocladia. Except for Tichocarpus and Endocladia, these are all members of the Gigartinales sensu Kylin (1956). Most of the families previously placed in the Cryptonemiales by Kylin appear to contain only lambda-type carrageenans. These fall into two groups, one that clusters with typical carrageenophyte- and the other with typical agarophyte-taxa. The first of these includes the families Polyideaceae, Kallymeniaceae, Dumontiaceae, and Rhizophyllidaceae. The second includes the type family of the Cryptonemiales, the Halymeniaceae, which is divisible into two well-supported clades, one of which possesses special lambda-like carrageenans, the aeodans, and the Schizymeniaceae, a recently created family containing two former gigartinalean genera. The agarophyte orders Gelidiales, Gracilariales and Ceramiales are well resolved as monophyletic clades, but their topological positions are poorly resolved. The Gelidiales and Ceramiales are associated, but the Gracilariales is included in a clade that contains families belonging to the Rhodymeniales, a possible agarophyte order. Finally, two groups that may contain lambda-like carrageenans, the Plocamiales and the Sarcodiaceae, are resolved as separate clades.

Systematics of red algae (Rhodophyta)

Red algal systematics is in a state of flux unparalleled since the early decades of this century. Ultrastructural and biochemical studies are providing features for a revised classification at ordinal and supraordinal ranks, whereas characters of vegetative and reproductive morphology continue to establish discontinuities among taxa at lower ranks. Cytological, biosystematic, and chemotaxonomic contributions are viewed as useful in defining relationships at specific and infraspecific ranks. Chemotaxonomic studies, depending on the properties of the taxa that are being studied, are applicable at all taxonomic levels. The taxonomic importance of life histories and themes of vegetative morphology, including spore germination, heterotrichy, initiation and differentiation of thalli are discussed at length. Homologies of auxiliary cells, carpogonial branches, and connecting filaments are examined in the context of relationships among Corallinales, Cryptonemiales, Gigartinales, and Rhodymeniales, and their inclu...

THE ATLANTIC SPECIES OF SOLIERIA (GIGARTINALES, RHODOPHYTA): THEIR MORPHOLOGY, DISTRIBUTION AND AFFINITIES1

Solieria chordalis (C. Agardh) J. Agardh and S. tenera (J. Agardh) Wynne et Taylor exhibit multiaxial growth from a cluster of four to eight obconical apical cells. A single periaxial cell is cut off from each axial cell and successive periaxial cells are rotated 120° in a zig‐zag pattern along each axial filament. Periaxial cells produce branched, laterally diverging filaments which form the cortex. The medulla is composed of axial cells, elongate cells of lateral filaments, stretched interconnecting cells, and secondary rhizoids. The two species are nonprocarpic. Carpogonial branches are 3‐celled, inwardly directed, with a reflexed trichogyne. The auxiliary cell together with associated darkly‐staining inner cortical cells form an association, the auxiliary cell complex, that is recognizable prior to diploidization. A single, unbranched, non‐septate connecting filament issues from the fertilized carpogonium and fuses with the inner, lateral side of an auxiliary cell. Production of an involucre from surrounding vegetative cells is stimulated and a gonimoblast initial is cut off toward the interior of the thallus which divides to form a compact cluster of gonimoblast cells. A fusion cell is produced through fusion of inner gonimoblast cells with the auxiliary cell that, in turn, fuses progressively with cells of the lateral file bearing the auxiliary cell. Mature cystocarps have terminal carposporangia cut off from gonimoblast cells at the periphery of the fusion cell and are surrounded by an involucre with a distinct ostiole. Tetrasporangia are cut off laterally from surface cortical cells which then cut off one or two additional derivatives toward the outside.