Meridionella gen. nov., a New Genus of Cystocloniaceae (Gigartinales, Rhodophyta) from the Southern Hemisphere, Including M. obtusangula comb. nov. and M. antarctica sp. nov.

The classification of Cystoclonium obtusangulum has been questioned since the species was first described by Hooker and Harvey as Gracilaria? obtusangula. The objective of this study was to provide the first comprehensive taxonomic analysis of Cystoclonium obtusangulum, based on DNA sequences coupled with morphological observations made on syntype specimens and new collections. Sequence divergences of rbcL, UPA, and COI-5P, and maximum-likelihood phylogenies for rbcL and 18S demonstrated that specimens identified as Cystoclonium obtusangulum represent a clade of two distinct species that are distantly related to the generitype Cystoclonium purpureum. A new genus, Meridionella gen. nov., is proposed for this clade. The two species placed in this new genus were morphologically indistinguishable cryptic species, but have disjunct distributions, with Meridionella obtusangula comb. nov. found from temperate to cold coasts of South America and the Falkland Islands and Meridionella antarctica sp. nov., occurring in Antarctic waters. Vegetative and reproductive characters of Meridionella gen. nov. are described, and implications of our results for the biogeography of the family Cystocloniaceae are discussed.

Analysis of the complete plastomes of three species of Membranoptera (Ceramiales, Rhodophyta) from Pacific North America.

Next generation sequence data were generated and used to assemble the complete plastomes of the holotype of Membranoptera weeksiae, the neotype (designated here) of M. tenuis, and a specimen examined by Kylin in making the new combination M. platyphylla. The three plastomes were similar in gene content and length and showed high gene synteny to Calliarthron, Grateloupia, Sporolithon, and Vertebrata. Sequence variation in the plastome coding regions were 0.89% between M. weeksiae and M. tenuis, 5.14% between M. weeksiae and M. platyphylla, and 5.18% between M. tenuis and M. platyphylla. We were unable to decipher the complete mitogenomes of the three species due to low coverage and structural problems; however, we assembled and analyzed, the cytochrome oxidase I, II, and III loci and found that M. weeksiae and M. tenuis differed in sequence by 1.3%, M. weeksiae and M. platyphylla by 8.4%, and M. tenuis and M. platyphylla by 8.1%. Evaluation of standard marker genes indicated that sequences from the rbcL, RuBisCO spacer, and CO1 genes closely approximated the pair-wise genetic distances observed between the plastomes of the three species of Membranoptera. A phylogenetic tree based on rbcL sequences showed that M. tenuis and M. weeksiae were sister taxa. Short rbcL sequences were obtained from type specimens of M. dimorpha, M. multiramosa, and M. edentata and confirmed their conspecificity with M. platyphylla. The data support the recognition of three species of Membranoptera occurring south of Alaska: M. platyphylla, M. tenuis, and M. weeksiae.

A NEW METHOD OF CYSTOCARP DEVELOPMENT IN THE RED ALGAL GENUS CALLOPHYLLIS (KALLYMENIACEAE) FROM CHILE(1).

Traditional studies suggest that the Kallymeniaceae can be divided into two major groups, a nonprocarpic Kallymenia group, in which carposporophyte formation involves an auxiliary cell branch system separate from the carpogonial branch system, and a procarpic Callophyllis group, in which the carpogonial branch system gives rise to the carposporophyte directly after fertilization. Based on our phylogenetic studies and unpublished observations, the two groups each contain both procarpic and nonprocarpic genera. Here, we describe a new method of reproductive development in Callophyllis concepcionensis Arakaki, Alveal et Ramírez from Chile. The carpogonial branch system consists of a supporting cell bearing both a three-celled carpogonial branch with trichogyne and two-lobed "subsidiary" cells. After fertilization, large numbers of secondary subcortical and medullary cells are produced. Lobes of the carpogonial branch system cut off connecting cells containing enlarged, presumably diploid nuclei that fuse with these secondary vegetative cells and deposit their nuclei. Derivative enlarged nuclei are transferred from one vegetative cell to another, which ultimately cut off gonimoblast initials that form filaments that surround the central primary medullary cells and produce carposporangia. The repeated involvement of vegetative cells in gonimoblast formation is a new observation, not only in Callophyllis, but in red algae generally. These results call for a revised classification of the Kallymeniaceae based on new morphological and molecular studies.

GRACILARIA HUMMII SP. NOV. (GRACILARIALES, RHODOPHYTA), A NEW NAME FOR THE AGAROPHYTE "GRACILARIA CONFERVOIDES" HARVESTED IN NORTH CAROLINA DURING WORLD WAR II(1).

Gracilaria hummii Hommers. et Freshwater is proposed as a new name for the inshore cylindrical species found in North Carolina that was treated as Gracilaria confervoides (L.) Grev. during World War II, and more recently as G. verrucosa (Huds.) Papenf. Molecular evidence places G. hummii in the Gracilis-group in Gracilaria together with G. gracilis (Stackh.) Steentoft, L. M. Irvine et Farnham, the name currently applied to specimens formerly identified as G. confervoides and G. verrucosa. G. hummii differs from G. gracilis in possessing shallower male conceptacles in which the spermatangial filament originates from a surface cortical cell rather than from a subcortical cell. The cystocarps are similar, except that the gonimoblasts of G. hummii are attached to the base of the pericarp by numerous, prominent thickened terminal tubular cells and because terminal tubular cells are absent above the midregion of the cystocarp or in the vicinity of the ostiole. The gonimoblasts are subtended by a bundle of longitudinally oriented, thick-walled secondary filaments of a type that has not been described before in Gracilaria but that may be characteristic of some other species. G. hummii occupies a basal position in the Gracilis-group and is distinct from all other cylindrical North Carolina Gracilaria species, according to the molecular and morphological evidence.

CHARACTERIZATION OF MARTENSIA (DELESSERIACEAE, RHODOPHYTA) BASED ON A MORPHOLOGICAL AND MOLECULAR STUDY OF THE TYPE SPECIES, M. ELEGANS, AND M. NATALENSIS SP. NOV. FROM SOUTH AFRICA(1).

An examination of a series of collections from the coast of Natal, South Africa, has revealed the presence of two species of Martensia C. Hering nom. cons: M. elegans C. Hering 1841, the type species, and an undescribed species, M. natalensis sp. nov. The two are similar in gross morphology, with both having the network arranged in a single band, and with reproductive thalli of M. elegans usually larger and more robust than those of M. natalensis. Molecular studies based on rbcL sequence analyses place the two in separate, strongly supported clades. The first assemblage occurs primarily in the Indo-West Pacific Ocean, and the second is widely distributed in tropical and warm-temperate waters. Criteria that have been used in the past for separating the two, namely, the number and shape of the blades, the presence of a single- versus a multiple-banded network, and blade margins entire or toothed, were determined to be unreliable. Although the examination of additional species is required, the morphology and position of procarps and cystocarps, whether at or near the corners of the longitudinal lamellae and the cross-connecting strands or along the lobed, membranous edges of the longitudinal lamellae or on the thallus margins, may prove to be diagnostic at the subgenus level. We recognize subg. Martensia, including the type of Martensia: M. elegans and subg. Mesotrema (J. Agardh) De Toni based on Martensia pavonia (J. Agardh) J. Agardh.

THE BIOGEOGRAPHIC ORIGIN OF ARCTIC ENDEMIC SEAWEEDS: A THERMOGEOGRAPHIC VIEW(1).

The Arctic is geologically and biogeographically young, and the origin of its seaweed flora has been widely debated. The Arctic littoral biogeographic region dates from the latest Tertiary and Pleistocene. Following the opening of Bering Strait, about 3.5 mya, the "Great Trans-Arctic Biotic Interchange" populated the Arctic with a fauna strongly dominated by species of North Pacific origin. The Thermogeographic Model (TM) demonstrates why climate and geography continued to support this pattern in the Pleistocene. Thus, Arctic and Atlantic subarctic species of seaweeds are likely to be evolutionarily "based" in the North Pacific, subarctic species are likely to be widespread in the warmer Arctic, and species of Atlantic Boreal or warmer origin are unlikely in the Arctic and Subarctic. Although Arctic seaweeds have been thought to have a greater affinity with the North Atlantic, we have reanalyzed the Arctic endemic algal flora, using the Thermogeographic Model and evolutionary trees based on molecular data, to demonstrate otherwise. There are 35 congeneric species of the six, abundant Arctic Rhodophyta that we treat in this paper; 32 of these species (91%) occur in the North Pacific, two species (6%) occur in the Boreal or warmer Atlantic Ocean, and a single species is panoceanic, but restricted to the Subarctic. Laminaria solidungula J. Agardh, a kelp Arctic "endemic" species, has 18 sister species. While only eleven (61%) occur in the North Pacific, this rapidly dispersing and evolving genus is a terminal member of a diverse family and order (Laminariales) widely accepted to have evolved in the North Pacific. Thus, both the physical/time-based TM and the dominant biogeographic pattern of relatives of Arctic macrophytes suggest strong compliance with the evidence of zoology, geology, and paleoclimatology that the Arctic marine flora is largely of Pacific origin.

TWO TYPES OF AUXILIARY CELL AMPULLAE IN GRATELOUPIA (HALYMENIACEAE, RHODOPHYTA), INCLUDING G. TAIWANENSIS SP. NOV. AND G. ORIENTALIS SP. NOV. FROM TAIWAN BASED ON rbcL GENE SEQUENCE ANALYSIS AND CYSTOCARP DEVELOPMENT(1).

Few species in the genus Grateloupia have been investigated in detail with respect to the development of the auxiliary cell ampullae before or after diploidization. In this study, we document the vegetative and reproductive structures of two new species of Grateloupia, G. taiwanensis S.-M. Lin et H.-Y. Liang sp. nov. and G. orientalis S.-M. Lin et H.-Y. Liang sp. nov., plus a third species, G. ramosissima Okamura, from Taiwan. Two distinct patterns are reported for the development of the auxiliary cell ampullae: (1) ampullae consisting of three orders of unbranched filaments that branch after diploidization of the auxiliary cell and form a pericarp together with the surrounding secondary medullary filaments (G. taiwanensis type), and (2) ampullae composed of only two orders of unbranched filaments in which only a few cells are incorporated into a basal fusion cell after diploization of the auxiliary cell and the pericarp consists almost entirely of secondary medullary filaments (G. orientalis type). G. orientalis is positioned in a large clade based on rbcL gene sequence analysis that includes the type species of Grateloupia C. Agardh 1822, G. filicina. G. taiwanensis clusters with a clade that includes the generitype of Phyllymenia J. Agardh 1848, Ph. belangeri from South Africa; that of Prionitis J. Agardh 1851, Pr. lanceolata from Pacific North America; and that of Pachymeniopsis Y. Yamada ex Kawab. 1954, Pa. lanceolata from Japan. A reexamination of the type species of the genera Grateloupia, Phyllymenia, Prionitis, and Pachymeniopsis is required to clarify the generic and interspecific relationships among the species presently placed in Grateloupia.

GAYLIELLA GEN. NOV. IN THE TRIBE CERAMIEAE (CERAMIACEAE, RHODOPHYTA) BASED ON MOLECULAR AND MORPHOLOGICAL EVIDENCE(1).

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.

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.