THE MORPHOLOGY AND DEVELOPMENT OF SOME PROMINENTLY STALKED SOUTHERN AUSTRALIAN HALYMENIACEAE (CRYPTONEMIALES,RHODOPHYTA). I. CRYPTONEMIA KALLYMENIOIDES (HARVEY) KRAFT COMB. NOV. AND C. UNDULATA SONDER1

Several southern Australian red algae of the family Halymeniaceae (Cryptonemiales) are differentiated into hard, massive stalks and considerably softer laminar blades or phyllodes. The taxonomy, morphology and pit-connection ultrastructure of one such species, Cryptonemia kallymenioides (Harvey) Kraft comb. nov., are compared to C. undulata Sonder, which lacks massive stalks. In both species there is extensive periodic secondary cortication of the stalks, resulting in the formation of distinct “growth rings.” The blades of C. kallymenioides appear to be seasonal and its stalks perennial, while plants of C. undulata are apparently perennial but shorter lived than C. kallymenioides. As a result, stalks in the latter can reach 2–3 cm in diameter with up to 18 growth rings, compared to the 1–2 mm diameters and up to 6 rings within the stalks of C. undulata. Heavy secondary thickening of cortical cell walls occurs in both species and confers a “woody” texture to the stalks of C. kallymeniodes. Regardless of the large differences in average stalk diameters between the two species, the pit-connection ultrastructure from cortex to medulla shows much the same sequence of morphological modification. Pit-connections are standard red algal structures in the outer cortex, but become increasingly convoluted on the membrane-bound surfaces abutting cytoplasm and develop wider apertures and less dense cores with increasing distance from the stalk surface. In occasional medullary cells of C. kallymenioides, the cytoplasm disintegrates, leaving cell walls and pit-connections to play an apparently structural role which has not been reported in other red algae. It is suggested that the increase in aperture size and surface areas of pit-connections is compatible with their playing a role in the intercellular transport of solutes towards the inner cell layers which may, in C. kallymenioides, lie many millimeters distant.     

SYNARTHROPHYTON,A NEW GENUS OF CORALLINACEAE (CRYPTONEMIALES,RHODOPHYTA) FROM THE SOUTHERN HEMISPHERE1

Synarthrophyton gen. nov. is described based on southern Australian material of Melobesia patena Hooker fils & Harvey, which shows features intermediate between Mesophyllum Lemoine and Lithothamnium Philippi. It has the thin-walled rounded epithallial cells and coaxial hypothallium characteristic of Mesophyllum and the type of procarp and male structures common to Lithothamnium. Synarthrophyton is unique within the subfamily Melobesioideae (J. E. Areschoug) Yendo because it displays secondary pit connections. Details of the vegetative and reproductive morphology of S. patena (Hooker fils. & Harvey) comb. nov. are given and its taxonomic position in the Corallinaceae Lamouroux discussed.     

ACROSYMPHYTON FIRMUM SP. NOV. (RHODOPHYTA,CRYPTONEMIALES), A NEW SUBTIDAL RED ALGA FROM NEW ZEALAND; DEVELOPMENTAL MORPHOLOGY AND DISTRIBUTION OF THE GAMETOPHYTE1

Acrosymphyton firmum sp. nov., is described from the northeastern coast of North 1., New Zealand. Gametophytes are spring–summer annuals which grow subtidally on cobbles. Thalli are uniaxial; each axial cell bears a whorl of four indeterminate and one determinate branchlets. Indeterminate branchlets are alternately arranged giving the thallus a distichous and feather-like appearance. Numerous corticating rhizoidal filaments are produced from the periaxial and lower whorl branchlet cells. These rhizoids entwine and obscure the main axis as the thallus develops until in the mature plant the axes have a firm consistency and lubricous texture. The carpogonial branch bearing short lateral filaments and auxiliary cell branch with terminal auxiliary cell place this new species in the genus Acrosymphyton Sjöstedt. Of the three described species in the genus, A. firmum is most similar to A. taylori. This is the first report of the genus Acrosymphyton and the only confirmed report of the family Dumontiaceae in New Zealand waters.     

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.     

THE NEW GENUS YONAGUNIA KAWAGUCHI ET MASUDA (HALYMENIACEAE,RHODOPHYTA), BASED ON Y. TENUIFOLIA KAWAGUCHI ET MASUDA SP. NOV. FROM SOUTHERN JAPAN AND INCLUDING Y. FORMOSANA (OKAMURA) KAWAGUCHI ET MASUDA COMB. NOV. FROM SOUTHEAST ASIA1

Yonagunia Kawaguchi et Masuda, gen. nov. (Halymeniaceae, Rhodophyta) is proposed to accommodate a new species, Yonagunia tenuifolia Kawaguchi et Masuda and the species currently known as Prionitis formosana (Okamura) Kawaguchi et Nguyen. Based on auxiliary cell ampullar features, Yonagunia is included in the group of genera with the simplest type of ampulla (the Grateloupia type) that comprises Dermocorynus, Grateloupia, Kintokiocolax, Phyllymenia, and Zymurgia. However, Yonagunia differs from these genera in the behavior of cells in the ampullar filaments immediately after diploidization, most cells of the primary and secondary filaments simultaneously dividing to form grape‐like clusters of small globular cells that subsequently elongate and produce involucral filaments to laxly surround the maturing carposporophyte. Yonagunia is resolved by our rbcL gene sequence analyses as one of five monophyletic clades within the Halymeniaceae (an Aeodes/Pachymenia, a Polyopes, a Carpopertis/Cryptonemia/Halymenia, a Yonagunia, and a Grateloupia clade) that is positioned as sister to the Grateloupia clade. Carpogonial branch apparatuses are identified as a potential taxonomic significance on the same level as auxiliary cell ampullae.     

AN EXPANDED SURVEY OF THE ULTRASTRUCTURE OF RED ALGAL PIT PLUGS1

The fine structure of pit plugs in 90 species of red algae was examined, bringing the total number of species in the continuing survey to 153. The organization of plug caps was confirmed to be a stable, predictable trait within thalli, between generations in heteromorphic life histories, and within the presently recognized orders, with one exception—the Acrochaetiales. Two forms of the outer cap were found in this group, a thin plate, as in the Nemaliales and Palmariales, and a dome, as in Batrachospermales and Corallinales. Variation of pit plug structure indicates that the Acrochaetiales are a heterogeneous assemblage and that pit plugs will be useful in reappraising their systematics. The systematic affinities of several species of uncertain affinities are clarified. Schmitziella endophloea Bornet et Batters is excluded from both orders, Corallinales and Acrochaetiales, with which it previously was allied. Although other ordinal attributions are not precluded by pit plug structure alone, pit plug structure is consistent with placement of Apophlaea sinclairii Harvey and Hildenbrandia rivularis (Liebman) J. Agardh in the Hildenbrandiales, Plagiospora gracilis Kuckuck, Schmitziella endophloea, and Wurdemannia miniata (Duby) J. Feldmann et Hamel in the Gigartinales, and Pseudorhododiscus nipponicus Masuda in the Palmariales.     

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.     

A COMPARATIVE STUDY OF THE RED ALGA GRATELOUPIA FILICINA (HALYMENIACEAE) FROM THE NORTHWESTERN PACIFIC AND MEDITERRANEAN WITH THE DESCRIPTION OF GRATELOUPIA ASIATICA, SP. NOV.

Morphological observations and molecular analyses of the red alga Grateloupia filicina (Halymeniaceae) from two geographically distant regions, eastern Asia (Japan and northern China) in the northwestern Pacific and Italy in the Mediterranean, reveal the presence of two distinct entities. Morphologically, the eastern Asian entity differs substantially from the Italian entity in the following ways: 1) thin and soft thalli with wider axes, 2) denser medullary filaments, 3) scattered reproductive structures over the entire thallus, and 4) a mature auxiliary cell that is oval and slightly larger than other ampullary cells. Phylogenetic analysis based on the ribulose-1,5-bisphosphate carboxylase/oxygenase gene ( rbc L) sequences revealed that the eastern Asian and Italian entities are phylogenetically far apart, strongly supporting the differentiation of these two entities at the species level. The eastern Asian entity is therefore described as a new species, Grateloupia asiatica. This species can be distinguished from most known species of Grateloupia that have widely flattened thalli by its compressed to narrowly flattened axes with numerous pinnate proliferations and from a few species with similar thalli by a particular combination of features, including a gelatinous texture, mostly simple and narrower axes, a thinner cortex, and the absence of catenate proliferations.     

MORPHOLOGY AND TAXONOMY OF MERISTIELLA GEN. NOV. (SOLIERIACEAE,RHODOPHYTA) 1

Eucheuma acanthocladum (Harvey) J. Agardh, E. gelidium (J Agardh) J. Agardh, E. echinocarpum Areschoug and E. schrammii(P. et H. Crouan) J. Agardh from the tropical and warm temperate waters of the western Atlantic Ocean and Caribbean Sea are transferred to a new genus, Meristiella. Meristiella exhibits the following Unique combination of characters among genera in the Solieriaceae: (1) rotated periaxial cells, (2) a loosely filamentous medulla. (3) an auxiliary cell complex, (4) Single and twin connecting filaments and (5) spinose cystocarps composed of a central, small-celled placentum, based on its reproductive features, Meristiella. is assigned to the tribe Agardhielleae. Culture experiments and herbarium studies provide evidence that E, gelidium and E. acantghocladum are conspecific. Lectotypes are designated for the included species.     

SYSTEMATIC SIGNIFICANCE OF THE ABSENCE OF PIT-PLUG CAP MEMBRANES IN THE BATRACHOSPERMALES (RHODOPHYTA)1

Four species of the Batrachospermales were examined by transmission electron microscopy to determine whether or not cap membranes, a typical structural component of pit plugs in several orders of red algae, were present. Routine specimen preservation methods used in past studies led to contradictory reports, so Batrachospermum keratophytum Bory, B. sirodotii Skuja ex Reis, Sirodotia suecica Kylin, and S. tenuissima (Collins) Skuja ex Flint were prepared by secondary fixation in potassium permanganate or a combination of potassium ferrocyanide-osmium tetroxide to enhance membrane contrast. These fixation procedures produced clear, well-contrasted images in which cap membranes were absent. The absence of cap membranes in S. suecica and S. tenuissima and the presence of cap membranes in two members of the Nemaliales was confirmed by freeze-substitution methods. Absence of cap membranes in representatives of the Batrachospermales further distinguishes the Batrachospermales from the Nemaliales, the order in which they long resided, and demonstrates the value of this character in elucidating ordinal alliances among the Rhodophyta.     

ULTRASTRUCTURE OF MEIOSIS IN DASYA BAILLOUVIANA (RHODOPHYTA). II. PROMETAPHASE I—TELOPHASE II AND POST-DIVISION NUCLEAR BEHAVIOR1

This is the second of two papers which together are the first comprehensive ultrastructural report of meiosis in a red alga. Many details of the meiotic process in Dasya baillouviana (Gmelin) Montagne are the same as those reported previously for mitotic cells in ceramialian red algae, but several characteristics seem unique to meiotic cells. The nucleus and nucleolus of meiotic cells are larger than those of mitotic cells and large accumulations of smooth ER are often found at the division poles during meiosis 1. The function of the ER accumulations is unknown. Importantly, both interkinesis and a simultaneous division of two separate nuclei during meiosis II was demonstrated. These new observations fail to support earlier speculation on higher red algae for a “uninuclear” meiosis (both nuclear divisions within the same nuclear envelope). However, following meiosis II the four nuclei migrate centripetally and possibly fuse in the center of the tetrasporangium. This post-division nuclear maneuvering is not understood, but our interpretation accounts for the earlier and erroneous impression of “uninuclear” meiosis. Perhaps the most important aspect of meiosis observed in Dasya is its basic adherence to the pattern commonly seen in higher plants and animals. This conservatism of the meiotic process lends further skepticism to the belief that red algae are extremely “primitive” organisms, although they undoubtedly represent a very “ancient” group of eukaryotic plants.     

AN EXAMINATION OF PACHYMENIA AND AEODES (HALYMENIACEAE,RHODOPHYTA) IN NEW ZEALAND AND THE TRANSFER OF TWO SPECIES OF AEODES IN SOUTH AFRICA TO PACHYMENIA1

A phylogenetic study was conducted of species of Halymeniaceae from New Zealand presently placed in Aeodes or Pachymenia, based on maximum‐likelihood (ML), maximum‐parsimony (MP), and Bayesian analyses of rbcL and nuclear internal transcribed spacer (ITS) rDNA sequences. We used molecular and morphological data in combination with exhaustive sampling of herbarium collections to clarify the taxonomy and distributions of New Zealand members of Pachymenia and Aeodes. Our study confirms the presence of three erect species of Pachymenia on the New Zealand mainland, and we resurrect the name Pachymenia dichotoma J. Agardh for the widely distributed, southernmost species. Species of Aeodes from South Africa are shown to be closely related to Pachymenia carnosa (J. Agardh) J. Agardh, the type species of Pachymenia, and are accordingly transferred to Pachymenia.     

MORPHOLOGY AND SYSTEMATICS OF RHODOCALLIS ELEGANS, RHODOCALLIDEAE, TRIB. NOV. (CERAMIACEAE, RHODOPHYTA), FROM SOUTHEASTERN AUSTRALIA

Recent collections of Rhodocallis elegans Kützing from southeastern Australia have permitted detailed observations of vegetative and reproductive structures that reveal features not exhibited by any existing tribe of Ceramiaceae. As a consequence, we establish the new tribe Rhodocallideae based on the unispecific genus Rhodocallis. Defining characters include: 1) four periaxial cells cut off in an alternating (rhodomelaceous) sequence; 2) determinate branchlets of two types: a) persistent lateral branchlets produced from the first-formed periaxial cells, and b) deciduous transverse branchlets produced from the second and third periaxial cells, with cortical filaments issuing from all four periaxial cells; 3) first- and second-order determinate branchlets terminated by thick-walled spines; 4) indeterminate branches formed at the tips of directly converted determinate branchlets; 5) axial cells of indeterminate branches heavily corticated by a cylinder of descending rhizoidal filaments; 6) spermatangial parent cells borne directly on unmodified outer cortical cells; 7) carpogonial branches borne in series on second and third periaxial cells of modified indeterminate axes; 8) procarps lacking sterile-cell groups; 9) a single derivative of the zygote nucleus transferred from the carpogonium to the auxiliary cell directly through a tube rather than by means of a connecting cell; 10) gonimoblasts surrounded by a network of rhizoidal filaments through which the gonimolobes protrude, the carposporophyte subtended by an investment of determinate branchlets; and 11) tetrasporangia tetrahedrally divided, borne on surface cortical cells of special determinate branchlets and protruding outside the cuticular layer.     

ULTRASTRUCTURE OF MEIOSIS IN DASYA BAILLOUVIANA (RHODOPHYTA). I. PROPHASE I1

This first of two papers on ultrastructural observations of meiosis in the red alga Dasya baillouviana (Gmelin) Montague describes stages of prophase I of meiosis. Although the five stages of prophase were originally derived from light microscopic studies, the same stages were utilized for this study based on the developmental sequence of the synaptonemal complex, which has the same morphology and mode of development as those reported for other red algae. The cytoplasm in early prophase sporocytes was typically less electron dense than either vegetative cells or sporocytes in later stages of meiosis. The reduction in density suggests clearing of ribosomes and other cytoplasmic components prior to conversion from sporophyte to gametophyte control. Leptotene cells often had an amorphous, chromatin-free area, function unknown, which was not obviously associated with any specific nuclear region. Diplotene cells were characterized by nuclei containing prominent ring-shaped nucleoli composed of a dark staining ring of material surrounding an electron-translucent “vacuole.” Packets of electron-dense, fibrillar material were often noted in the cytoplasm of late prophase cells. These packets are thought to he “nuage,” a term applied to large cytoplasmic aggregations of RNA in germ cells of several other phyla. It is suggested that nuage may represent a new infusion of ribosomal and messenger RNA for post-meiotic development. The division pales are established by late prophase and a single polar ring is found within each large “exclusion zone” in close association with a pore-free area of nuclear envelope. Both annulate lamellae and small, numerous vesicles are located in the exclusion zones. The significance of the various aspects of prophase I is discussed with the overall observation that this phase of meiosis in red algae is very similar to the process in higher plant and animal cells.     

STRUCTURE AND DEVELOPMENT OF THE CARPOSPOROPHYTE OF NEMALION HELMINTHOIDES (NEMALIONALES,RHODOPHYTA)1

Following fertilization, the carposporophyte of Nemalion helminthoides (Velley in With.) Batters differentiates into four distinct regions: the fusion cell, the sterile gonimoblast cells, the carposporangial mother cells and the carposporangia. The gonimoblast is formed by apically dividing, monopodial filaments of limited growth which may later become pseudodichotomous. Upon differentiation of a terminal carposporangium, a gonimoblast filament may continue to grow sympodially. A single carposporangial mother cell may produce carposporangia in several different directions as well as proliferate successive carposporangia within the sporangial walls that remain after carpospore liberation. As the carposporophyte matures, the gonimoblast initial, the stalk cell, the hypogynous and subhypogynous cells fuse. Except for the fusion cell, all cells of the carposporophyte show organelle polarity and contain a distally located, lobed chloroplast and proximal nucleus.     

VEGETATIVE AND REPRODUCTIVE MORPHOLOGY IN RETICULOCAULIS GEN. NOV. AND NACCARIA HAWAIIANA SP. NOV. (RHODOPHYTA,NACCARIACEAE)1

A secondarily formed reticulum of cells that encloses axial strands and a carposporophyte that is almost entirely converted to sporangia are the diagnostic features of a new genus, Reticulocaulis (Naccariaceae, Nemaliales), with R. mucosissimus as the type species. A new species of Naccaria, N. hawaiiana permits close comparison with the new genus and other previously described species of Naccaria. Until now, no Naccariaceae were known in the Pacific, and these new taxa are part of the sketchily known subtidal marine flora of Hawaii.     

SYSTEMATICS OF GRATELOUPIA FILICINA (HALYMENIACEAE,RHODOPHYTA), BASED ON rbcL SEQUENCE ANALYSES AND MORPHOLOGICAL EVIDENCE,INCLUDING THE REINSTATEMENT OF G. MINIMA AND THE DESCRIPTION OF G. CAPENSIS SP. NOV.1

Grateloupia filicina (C. Agardh) Lamouroux, originally described from the Mediterranean Sea, has long been considered a textbook example of a marine red alga with a cosmopolitan distribution. An rbcL‐based molecular phylogeny, encompassing samples covering the entire geographic distribution of the species, revealed a plethora of “cryptic” species, whereby the presence of genuine G. filicina is limited to the Mediterranean basin. The phylogeny revealed a strong biogeographic imprint, with specimens from temperate regions resolved in clades composed of species inhabiting the same geographic region. Presence of widely divergent morphologies in the temperate clades indicated that several lineages have converged independently to a G. filicina‐type morphology. Tropical representatives are resolved in a single clade with very uniform G. filicina‐type morphology and pairwise sequence divergences that are lower than the average divergence observed in temperate lineages. This, combined with a lack of clear geographic structure among the tropical lineages, may indicate a more recent divergence with long‐range dispersal capacities. Violations to the biogeographic signal in temperate lineages seemed to be due to either inadequate taxonomy or recent introductions. Grateloupia minima P. & H. Crouan, a taxon placed in synonymy under G. filicina, is reinstated as a separate species distributed in the northeast Atlantic Ocean. Grateloupia capensis sp. nov. is described to accommodate specimens from South Africa with a G. filicina‐type morphology, and G. filicina var. luxurians is elevated to species status. Morphological and anatomical characters were put forward that support the distinctiveness of these three distinct species.     

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 DEVELOPMENT1

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.     

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.     

MORPHOLOGY AND SYSTEMATICS OF SCIUROTHAMNION STEGENGAE GEN. ET SP. NOV. (CERAMIACEAE,RHODOPHYTA) FROM THE INDO‐PACIFIC1

A new ceramiaceous alga, Sciurothamnion stegengae De Clerck et Kraft, gen. et sp. nov., is described from the western Indian Ocean and the Philippines. Sciurothamnion appears related to the tribe Callithamnieae on the basis of the position and composition of its procarps and by the majority of post‐fertilization events. It differs, however, from all current members of the tribe by the presence of two periaxial cells bearing determinate laterals per axial cell. Additionally, unlike any present representative of the subfamily Callithamnioideae, no intercalary foot cell is formed after diploidization of the paired auxiliary cells. The genus is characterized by a terminal foot cell (“disposal cell”), which segregates the haploid nuclei of the diploidized auxiliary cell from the diploid zygote nucleus. The nature of three types of foot cells reported in the Ceramiaceae (intercalary foot cells containing only haploid nuclei, intercalary foot cells containing haploid nuclei and a diploid nucleus, and terminal foot cells containing only haploid nuclei) is discussed.