Pterosiphonia tanakae (Rhodomelaceae, Ceramiales), a new red algal species from Japan

The red alga Pterosiphonia tanakae Uwai et Masuda (Rhodomelaceae, Ceramiales) is established as a new species on the basis of material collected in Hiroshima Bay, the Inland Sea, Japan. It can be distinguished from the majority of previously described species of the genus by two key features, seven to 10 periaxial cells and the presence of weakly developed cortical cells. This species is further distinguished from P. pau-cicorticata Dawson by completely alternate-distichously branching and from Pjavanica (Martens) De Toni and P. spinifera (Kützing) Norris et Aken by the production of first-order laterals that bear branches of up to five orders, which results in the thallus having a widely spreading, fan-shaped appearance. The abundant production of vegetative trichoblasts is a further characteristic feature of P. tanakae.     

Taxonomic notes on Laurencia mariannensis (Rhodomelaceae, Rhodophyta)

The marine red alga Laurencia mariannensis Yamada (Rhodomelaceae, Ceramiales) is characterized by pale rose-red, softly fleshy, slender, terete axes (up to 600 μm in diameter), arising from a loosely entangled, stoloniferous basal system among species that have the following combination of features: the presence of longitudinally orientated secondary pit-connections between contiguous superficial cortical cells; the presence of projecting superficial cortical cells at the upper portions of branches; and the presence of lenticular thickenings in the walls of medullary cells, Furthermore, the presence of two or three corps en cerise per superficial cortical cell and one per trichoblast cell may characterize the species.     

Taxonomic notes on Laurencia brongniartii (Rhodomelaceae, Rhodophyta)

The red alga Laurencia brongniartii J. Agardh (Rhodomelaceae, Ceramiales) is characterized by: (i) the production of four periaxial cells from each vegetative axial segment; (ii) the presence of two or three corps en cerise per superficial cortical cell and one per trichoblast cell; (iii) the production of a single tetrasporangium-bearing periaxial (fourth) cell per fertile segment; (iv) a tetrasporangial arrangement that is intermediate between perpendicular and parallel types; (v) procarps produced from the last-formed (fifth) periaxial cell of the terminal segment of a two-celled female trichoblast; and (vi) distally positioned spermatangial nuclei, in addition to known features. The production of a single tet-rasporangium-bearing periaxial cell per fertile segment allies this species to Laurencia similis Nam et Saito.     

Developmental morphology of Odonthalia floccosa (Ceramiales, Rhodophyta) in laboratory culture

Material of the red alga Odonthalia floccosa (Esper) Fal-kenberg (Rhodomelaceae, Ceramiales), collected from California, was cultured in the laboratory and its life-history was completed. Tetraspores grew into bipolar sporelings that differentiated into a colorless rhizoidal portion and a pigmented upright shoot. The sporelings became compressed apically and formed lateral branches in a regularly distichous manner that were congenitally fused with the main axis. These tetraspore germlings grew into diecious gametophytes. Male ga-metophytes produced numerous spermatangia on modified fertile branchlets (male trichoblasts) that possessed three to four monosiphonous, proximal segments. Female gametophytes formed a single pro-carp on the suprabasal segment of unbranched female trichoblasts. Cystocarps developed on the female gametophytes cocultured with male gametophytes and released viable carpospores that developed into fertile te-trasporophytes. Tetrasporangia were produced from the third and fourth periaxial cells in each of 12–45 successive fertile segments and provided three (two lateral and one basal) cover cells. The occurrence of both spermatangia and procarps on fertile trichoblasts in O. floccosa suggests that the alga is the most derived in these two characters among the species of the genus Odonthalia. This species is distributed in cold temperate regions in the North Pacific, and it should be excluded from the North Atlantic marine algal flora.     

Phylogenomic analysis of pseudocryptic diversity reveals the new genus Deltalsia (Rhodomelaceae,Rhodophyta)

Molecular analyses, in combination with morphological studies, provide invaluable tools for delineating red algal taxa. However, molecular datasets are incomplete and taxonomic revisions are often required once additional species or populations are sequenced. The small red alga Conferva parasitica was described from the British Isles in 1762 and then reported from other parts of Europe. Conferva parasitica was traditionally included in the genus Pterosiphonia (type species P. cloiophylla in Schmitz and Falkenberg 1897), based on its morphological characters, and later transferred to Symphyocladia and finally to Symphyocladiella using molecular data from an Iberian specimen. However, although morphological differences have been observed between specimens of Symphyocladiella parasitica from northern and southern Europe they have yet to be investigated in a phylogenetic context. In this study, we collected specimens from both regions, studied their morphology and analyzed rbcL and cox1 DNA sequences. We determined the phylogenetic position of a British specimen using a phylogenomic approach based on mitochondrial and plastid genomes. Northern and southern European populations attributed to S. parasitica represent different species. Symphyocladiella arecina sp. nov. is proposed for specimens from southern Europe, but British specimens were resolved as a distant sister lineage to the morphologically distinctive Amplisiphonia, so we propose the new genus Deltalsia for this species. Our study highlights the relevance of using materials collected close to the type localities for taxonomic reassessments, and showcases the utility of genome-based phylogenies for resolving classification issues in the red algae.     

Taxonomic notes on Hawaiian Polysiphonia, with transfer to Neosiphonia (Rhodomelaceae, Rhodophyta)

After detailed observations of type material and other collections, five Hawaiian species of Polysiphonia Greville, nom. cons. are recognized to be species of Neosiphonia M. S. Kim et I. K. Lee; namely, Neosiphonia apiculata (Hollenberg) Masuda et Kogame, Neosiphonia beaudettei (Hollenberg) M. S. Kim et Abbott, comb. nov., Neosiphonia hawaiiensis (Hollenberg) M. S. Kim et Abbott, comb. nov., Neosiphonia profunda (Hollenberg) M. S. Kim et Abbott, comb. nov., and Neosiphonia rubrorhiza (Hollenberg) M. S. Kim et Abbott, comb. nov. These five species are ecorticate, having lateral branch initials and trichoblasts produced on successive segments, rhizoids separated from pericentral cells by a cross wall, three‐celled carpogonial branches (not seen in N. beaudettei and N. rubrorhiza), spermatangial branches arising on a primary branch of the trichoblasts, and tetrasporangia in a spiral series. Although certain characters were not available for some species, all other characters occur in a combination that is unique for members of Neosiphonia.     

Neorhodomela enomotoi sp. nov. (Ceramiales, Rhodophyta) from Japan

The red alga Neorhodomela enomotoi Masuda et Kogame (Rhodomelaceae, Ceramiales) is described as a new species from Japan. It is characterized by the following combination of features: (i) thalli reddish-brown and slightly rigid; (ii) first-order branches mostly indeterminately branched; (iii) adventitious branches infrequent, determinately branched and formed chiefly in the axils of lateral branches; (iv) vegetative trichoblasts abundant; (v) tetrasporangia produced on paniculate ultimate and penultimate branches; and (vi) cystocarps flask-shaped (urceolate). This species has been found growing only in the upper subtidal zone in the warm temperate waters of Japan.     

The anatomy of Neotenophycus ichthyosteus gen. et sp. nov. (Rhodomelaceae,Ceramiales), a bizarre red algal parasite from the central Pacific

A minute parasite of Neosiphonia poko (Hollenberg) Abbott from a shallow lagoon on the central-Pacific Johnston Atoll is described as Neotenophycus ichthyosteus Kraft et Abbott, gen. et sp. nov. The infective parasite cell first connects to a central-axial cell of the host, then emerges from between host pericentral cells at a node before dividing into a three- or four-celled primary axis. Epibasal cells of the parasite divide to form three pericentral cells whose derivatives produce a globular head on the basal cell and on which reproductive structures differentiate almost immediately. Trichoblasts on any life-history stage are completely lacking. Spermatangia are borne on mother cells across the whole thallus surface. Procarps consist of four pericentral cells that encircle a subapical fertile-axial cell in an ampullar configuration, one of the pericentral cells serving as the supporting cell and bearing a four-celled carpogonial branch and a single sterile cell. Diploidization results in a longitudinal/concave division of the auxiliary cell and formation of an arching linear series of inner gonimoblast cells, each dividing toward the thallus surface into gonimoblast filaments of very narrow, horizontally aligned cells terminated by initially monopodial, later by sympodial, carposporangia, the whole of the mature female gametophyte consisting of an amalgam of several cystocarps within a lax jacket of sterile gametophytic tissue. Tetrasporophytes are composed of lobes of pericentral-cell-derived filaments, each axial cell of which is ringed by three pericentral cells producing tetrahedral tetrasporangia enclosed by two pre-sporangial cover cells. Affinities of the new genus are discussed and comparison is made particularly to the enigmatic parasite Episporium centroceratis Möbius. It is concluded that relationships with any previously described tribe are so remote or obscure that the new tribe Neotenophyceae should be proposed for it.     

The morphology and taxonomy of Womersleya monanthos, an endemic species and genus of Delesseriaceae (Ceramiales, Rhodophyta) from southeastern Australia

Womersleya monanthos (J. Agardh) Papenfuss is typically an epiphyte of larger brown and red algae that are common in drift along the southeastern coasts of Australia. A hitherto little-known member of the Phycodrys group of the Nitophylloideae, its reproductive features have been studied in detail and its taxonomic position clarified. Blades are polystromatic throughout and lack veins or nerves, with blades originating from apical cells of primary and second-order cell rows. Intercalary cell divisions take place in primary cell rows and all other branch orders, with third-order laterals arising both abaxially and adaxially on cells of second-order rows. Fertile central cells bear procarps on pericentral cells on both sides of the blade, the procarps consisting of two 4-celled carpogonial branches and a single central group of sterile cells that enlarge and persist at the distal end of a bicampanulate fusion cell at maturity. Spermatangia and tetrasporangia form in circular subapical sori on both sides of the blade or in marginal lobes or proliferations. After comparing it to other members of the Phycodrys group, we conclude that Womersleya is a monotypic genus well distinguished from other genera and with probable closest affinities to the Northern Hemisphere Polyneura, Erythroglossum and Sorella, as well as the Australian endemic, Crassilingua.     

First record of Palisada maris-rubri (Ceramiales, Rhodophyta) from the Mediterranean Sea along with three proposed transfers to the genus Palisada

The first occurrence of Palisada maris-rubri (K.W. Nam et Saito) K.W. Nam (Ceramiales, Rhodomelaceae) from the Mediterranean Sea, is reported. To date the species was known only from tetrasporic specimens from the type locality (Ras Muhammed, Sinai, Egypt, Red Sea). Mediterranean thalli share nearly all vegetative and reproductive features with Red Sea specimens showing more robust thalli with axes to 3 mm broad and ultimate branchlets to 1000 µm broad, absence of intercellular spaces between medullary cells and epidermal cells in transverse section with a palisade arrangement. Male and cystocarpic thalli are recorded for the first time. Moreover, the analysis of characters of three species of Chondrophycus previously known from the Mediterranean Sea ( C. patentirameus (Montagne) K.W. Nam, C. tenerrimus (Cremades) G. Furnari et al. and C. thuyoides (Kützing) G. Furnari) led us to conclude that they belong to the genus Palisada . The following new combinations are formally proposed: P. patentiramea (Montagne) Serio et al., P. thuyoides (Kützing) Serio et al., P. tenerrima (Cremades) Serio et al.     

Molecular relationships within Bostrychia tenuissima (Rhodomelaceae, Rhodophyta)

Bostrychia tenuissima King and Puttock is restricted to southern Australia and New Zealand. Previous studies in Australia have revealed two distinct patterns in the presence of osmotically active polyols. Southern populations only have D-sorbitol whereas northern populations have both D-sorbitol and D-dulcitol. These polyol patterns lead to speculation on the ecotypic differentiation of these two population types. Using single-stranded confirmation polymorphism to rapidly score plastid haplotypes, plus DNA sequencing, a 100% congruence was found between polyol patterns and plastid haplotype. Analysis of 33 B. tenuissima isolates shows that the plastid relationships closely follow biogeo-graphic regions, with south-eastern Australia (southern NSW and Victoria) and South Australia and Tasmania having one haplotype and central and northern NSW having two alternate haplotypes. There is an overlap at the haplotype boundary just south of Sydney, New South Wales.     

Australian Laurencia majuscula (Rhodophyta,Rhodomelaceae) and the Brazilian Laurencia dendroidea are conspecific

Morphological and molecular studies have been undertaken on two species of the red algal genus Laurencia J.V.Lamouroux: Laurencia majuscula (Harvey) A.H.S. Lucas and Laurencia dendroidea J.Agardh, both from their type localities. The phylogenetic position of these species was inferred by analysis of the chloroplast‐encoded rbcL gene sequences from 24 taxa. In all phylogenetic analyses, the Australian Laurencia majuscula and the Brazilian L. dendroidea formed a well‐supported monophyletic clade within the Laurencia sensu stricto. This clade was divided into two subclades corresponding to each geographical region; however, the genetic divergence between Australian L. majuscula and Brazilian L. dendroidea was only 0–1.35%. Examination of the type specimens and sequences of freshly collected samples of both Laurencia majuscula and L. dendroidea show the two to be conspecific despite their disjunct type localities.     

Sesquiterpenoids of Laurencia majuscula (Ceramiales, Rhodophyta) from the Ryukyu Islands, Japan

Two populations of the red alga Laurencia majuscula (Harvey) Lucas (Rhodomelaceae, Ceramiales) from Taketomi Island and Hateruma Island, the Ryukyu Islands, Japan, have been characterized on the basis of both morphological features and halogenated secondary metabolite content. These populations have smaller and more slender thalli than those of other regions. Furthermore, the populations contain two chamigrane-type Sesquiterpenoids, (2R, 3R, 5S)-5-acetoxy-2-bromo-3-chlorochamigra-7(14),9-dien-8-one and (2R, 3R)-2-bromo-3-chlorochamigra-7(14), 9-dien-8-one, and a laurane-type sesquiterpenoid, debromoisolaurinterol, as secondary metabolites which are different from those previously reported from other populations. These results are consistent with the concept of ‘chemical races’ within a single species of Laurencia.     

Reproductive morphology and taxonomic reappraisal of Exophyllum wentii Weber-van Bosse (Rhodomelaceae, Rhodophyta) from Bali Island, Indonesia

The little‐known and rarely collected alga Exophyllum wentii Weber‐van Bosse is re‐described in detail from the type material, as well as from new collections from Indonesia, which for the first time reveal in detail the structure of cystocarpic and spermatangial plants and the development of tetrasporangial stichidia under culture conditions. New morphological reproductive information confirms placement of the genus Exophyllum within the Rhodomelaceae. Exophyllum is distinguished from other related genera within the Rhodomelaceae by its cartilaginous, non‐trichoblastic decumbent thallus with multiple holdfasts and its discoid spermatangial organs. Some Pacific material earlier attributed to E. wentii was found to be misidentified and re‐assigned to the Dasyaceae.     

A morphological and molecular assessment of the genus Sinotubimorpha (Halymeniaceae, Rhodophyta)

A critical reassessment of the morphological features of two closely related red algal genera,Grateloupia C.Agardh and Sinotubimorpha W.X.Li & Z.F.Ding (Halymeniaceae),pointed out that members of the t...     

Hybridization studies in Bostrychia. 1: B. radicans (Rhodomelaceae, Rhodophyta) from Pacific and Atlantic North America

Bostrychia radicans(Montagne) Montagne is a pantropical/temperate red alga associated with mangroves and saltmarsh plants. Collections were made from a similar north-south geographic distribution along both the Pacific and Atlantic coasts of North America. Hybridization studies were performed with cultured isolates to assess the extent of interfertility and reproductive isolation along these two coastlines. All male and female gametophytes derived from single tetrasporophytes were intercompatible. Almost all isolates extending over 1500 km of coast line from northern Pacific Mexico are compatible, forming cystocarps that released viable carpospores. Even isolates which morphologically would be placed in two species [B. radicans and B. moritziana(Sender ex Kützing) J. Agardh], based on the presence or absence of monosiphonous branches, were capable of hybridizing. Crosses of isolates from the Atlantic USA showed a greater amount of incompatibility. Certain isolates were not compatible with any other isolates including isolates collected in close proximity (North Carolina isolates), while other isolates from the same locality were compatible (South Carolina). An isolate from South Carolina formed tetrasporophytes with isolates from Pacific Mexico but tetraspores were not viable. Certain incompatible crosses formed ‘pseudocystocarps’ but viable carposporophytes did not develop. Generalizations about reproductive isolation within a species must also consider differences between populations from different biogeographic regions that may reflect different paleoclimatological histories, founder effects and unique dispersal events.     

The morphology,taxonomy, and molecular phylogeny of Heterocladia and Trigenea (rhodomelaceae,rhodophyta), with delineation of the little‐known tribe Heterocladieae*

A morphological, anatomical, and molecular study of the two genera (Heterocladia and Trigenea) and three species of the tribe Heterocladieae (Rhodomelaceae, Ceramiales) is presented. First collections of male and female gametophytes of Heterocladia australis Decaisne and Trigenea umbellata J. Agardh and of tetrasporophytes of the type species of Trigenea, T. australis Sonder, have allowed a much clearer assessment of these taxa from a classical morphological standpoint. Reproductive and vegetative characters of the two Trigenea species are shown to be virtually identical to those of Heterocladia, which differs from Trigenea principally in having both flattened and terete lateral branches, as opposed to exclusively terete axes throughout. As a consequence, we propose to transfer the Trigenea species to the earlier‐named genus Heterocladia as H. caudata L. Phillips, H.‐G. Choi, G.W. Saunders et Kraft, nom. nov. and H. umbellata ( J. Agardh) L. Phillips, H.‐G. Choi, G.W. Saunders et Kraft, comb. nov. The close relationship of the three species is supported by molecular data, as nucleotide sequences of the 18S rRNA gene from each are nearly identical. The same sequences from species of eight other rhodomelaceous genera plus those from five outgroup taxa are analyzed to provide grounds for preliminary phylogenetic inferences about the position of the Heterocladieae in the Rhodomelaceae. Both the Heterocladieae and the Rhodomelaceae are monophyletic taxa in our analyses, the Heterocladieae grouping weakly with the Bostrychieae and the problematic Australian endemic genus Sonderella, the latter yet to be assigned to a tribe. Representatives of groups with which Heterocladia has been associated previously, such as the Lophothalieae and Brongniartelleae, appear to be only distantly related, although many more taxa need to be analyzed before the systematic position of the genus becomes clear.