Diversity of bladed Bangiales (Rhodophyta) in western Mediterranean: recognition of the genus Themis and descriptions of T. ballesterosii sp. nov., T. iberica sp. nov., and Pyropia parva sp. nov.

The diversity of the bladed species of the red algal order Bangiales from the Iberian Mediterranean shores has been reassessed after a detailed study of this region. Prior to this study, 11 bladed species of Bangiales had been reported from Mediterranean waters: Porphyra atropurpurea, P. cordata, P. coriacea, P. dioica, P. linearis, P. purpurea, P. umbilicalis, Pyropia leucosticta, Pyropia koreana (as P. olivii), Py. elongata (as P. rosengurttii) and Py. suborbiculata. A combined analysis of the nuclear nSSU and the plastid rbcL genes together with detailed morphological studies has confirmed the presence of species within the genera Porphyra and Pyropia and also revealed a third, undescribed genus, Themis gen. nov. Porphyra linearis, Pyropia elongata and the introduced Pyropia koreana had been previously listed for the Mediterranean and were recorded in this study. An additional four species, including the introduced Pyropia suborbiculata and three new species: Pyropia parva sp. nov., Themis ballesterosii sp. nov., and Themis iberica sp. nov. were also observed. Hence, most of the Porphyra species traditionally reported along these shores were not reported in this survey. This new floristic Bangiales composition confirms the importance of the Mediterranean basin as a hotspot for biodiversity, possible endemics of ancient origin and high proportion of introductions. Our data also continue to confirm the extent of Bangiales diversity at regional and worldwide levels.     

A NEW LOOK AT AN ANCIENT ORDER: GENERIC REVISION OF THE BANGIALES (RHODOPHYTA)1

The red algal order Bangiales has been revised as a result of detailed regional studies and the development of expert local knowledge of Bangiales floras, followed by collaborative global analyses based on wide taxon sampling and molecular analyses. Combined analyses of the nuclear SSU rRNA gene and the plastid RUBISCO LSU (rbcL) gene for 157 Bangiales taxa have been conducted. Fifteen genera of Bangiales, seven filamentous and eight foliose, are recognized. This classification includes five newly described and two resurrected genera. This revision constitutes a major change in understanding relationships and evolution in this order. The genus Porphyra is now restricted to five described species and a number of undescribed species. Other foliose taxa previously placed in Porphyra are now recognized to belong to the genera Boreophyllum gen. nov., Clymene gen. nov., Fuscifolium gen. nov., Lysithea gen. nov., Miuraea gen. nov., Pyropia, and Wildemania. Four of the seven filamentous genera recognized in our analyses already have generic names (Bangia, Dione, Minerva, and Pseudobangia), and are all currently monotypic. The unnamed filamentous genera are clearly composed of multiple species, and few of these species have names. Further research is required: the genus to which the marine taxon Bangia fuscopurpurea belongs is not known, and there are also a large number of species previously described as Porphyra for which nuclear SSU ribosomal RNA (nrSSU) or rbcL sequence data should be obtained so that they can be assigned to the appropriate genus.     

Phylogeography of North Atlantic intertidal tardigrades: refugia,cryptic speciation and the history of the Mid‐Atlantic Islands

Aim To analyse the phylogeographical history of intertidal tardigrades in the North Atlantic in order to improve our understanding of geographical differentiation in microscopic organisms, and to understand the potential importance of the Mid‐Atlantic Islands as stepping stones between the American and European coasts of the Atlantic Ocean. Location Twenty‐four localities from the Mid‐Atlantic Islands (Greenland, Iceland and the Faroe Islands) and both sides of the North Atlantic Ocean. Methods A mitochondrial marker (cytochrome c oxidase subunit I) was sequenced from individual tardigrades belonging to the genus Echiniscoides. The existence of cryptic species was detected using generalized mixed Yule coalescence analysis; lineage ages were estimated with relaxed clock methods; and the degree of geographical differentiation was analysed with samova analyses, haplotype networks and Mantel tests. Results Echiniscoides hoepneri, previously known only from Greenland, was recovered throughout the Mid‐Atlantic Islands. The Faroe Islands population was isolated from Greenland and Iceland, but overall genetic variation was low. The morphospecies Echiniscoides sigismundi had high genetic variation and consisted of at least two cryptic species. A northern and a southern species were both recovered on both sides of the Atlantic, but only the northern species was found on the Mid‐Atlantic Islands. The northern species showed signs of long‐term isolation between the Western and Eastern Atlantic, despite the potential of the Mid‐Atlantic islands to act as stepping‐stones. There was no sign of long‐term isolation in the southern species. The Mid‐Atlantic individuals of the northern species were of Eastern Atlantic origin, but Greenland and Iceland showed signs of long‐term isolation. The genetic pattern found in the southern species is not clearly geographical, and can probably be best explained by secondary contact between former isolated populations. Main conclusions North Atlantic intertidal tardigrades from the genus Echiniscoides showed strong geographical differentiation, and the Mid‐Atlantic Islands seemed unimportant as stepping stones across the Atlantic. The geographical variation of the northern species of E. sigismundi suggests post‐glacial recolonization from several refugia.     

rbcL gene sequences reveal relationships among north-east Pacific species of Porphyra (Bangiales, Rhodophyta) and a new species, P. aestivalis

Phylogenetic analyses of the rbcL (chloroplast Rubisco large subunit) gene from 23 newly sequenced species of Porphyra, primarily from the north‐east Pacific, one Bangia and previously published sequences from both genera resolve relationships among most species of Porphyra and reveal five clades of species with Porphyra‐type morphologies among a number of Bangia lineages: (1) P. papenfussii V. Krishnam; (2) P. mumfordii S. C. Lindstrom et K. M. Cole and P. rediviva Stiller et Waaland together with a group of north Atlantic species, including the type of the genus, P. purpurea (Wahl‐enb.) C. Agardh; (3) P. cuneiformis (Setch. et Hus) V. Krishnam., P. occidentalis Setch. et Hus, P. schizo‐phylla Hollenb., and P. variegata (Kjellm.) Kjellm. and their Atlantic sibling species, all distromatic; (4) P. aestivalis sp. nov. and its north Atlantic sibling, P. birdiae C. D. Neefus et A. C. Mathieson; and (5) a speciose clade containing both Pacific and Atlantic representatives. Close relationships are confirmed between sibling species previously identified by iso‐zymes, morphology and chromosomal features. The morphologically similar dioecious P. pseudolanceolata V. Krishnam., P. conwayae (S. C. Lindstrom et K. M. Cole) stat. nov., and P. lanceolata (Setch. et Hus) G. M. Smith occur in a strongly supported subclade in clade 5 together with the monoecious P. fallax S. C. Lindstrom et K. M. Cole. Results presented here highlight the need for intensive taxon sampling and for examination of different parts of the genome to understand more fully relationships among species and higher level taxa in the Bangiales.     

Genetic and morphological differentiation of Porphyra and Pyropia species (Bangiales,Rhodophyta) coexisting in a rocky intertidal in Central Chile

A recent molecular taxonomic study along the Chilean coast (18° S–53° S) described 18 candidate species of bladed Bangiales of which only two were formally described. Few studies focused on local genetic and morphological diversity of bladed Bangiales and attempted to determine their intertidal distribution in contrasting habitats, and none were performed in Chile. To delimit intertidal distributions of genetic species, 66 samples of bladed Bangiales were collected at Maitencillo (32° S) in four zones: a rocky platform, a rocky wall, and two boulders zones surrounded by sandy and rocky bottoms, respectively. These samples were identified based on sequences of the mitochondrial COI and chloroplast rbcL markers. We also collected 87 specimens for morphological characterization of the most common species, rapidly assessing their putative species identity using newly developed species‐diagnostic (PCR‐RFLP) markers. Eight microscopic and two macroscopic morphological traits were measured. We described and named three of four species that predominate in Maitencillo (including Pyropia orbicularis): Pyropia variabilis Zapata, Meynard, Ramírez, Contreras‐Porcia, sp. nov., Porphyra luchea Meynard, Ramírez, Contreras‐Porcia sp. nov., and Porphyra longissima Meynard, Ramírez, Contreras‐Porcia, sp. nov. With the exception of Po. longissima restricted to boulders surrounded by sandy bottom, and a morphotype of Py. variabilis restricted to rocky walls, the other species/morphotypes have overlapping intertidal distributions. Except for Po. longissima, which is clearly differentiated morphologically (longest and thinnest blades), we conclude that morphology is not sufficient to differentiate bladed Bangiales. Our findings underscore the importance of refining our knowledge of intrinsic and environmental determinants on the distribution of bladed Bangiales.     

Diversity and distribution of foliose Bangiales (Rhodophyta) in West Greenland: a link between the North Atlantic and North Pacific

Greenland is a continental island in the northern part of the North Atlantic where the foliose Bangiales flora is poorly known. It is an important area for the study of algal biogeography because of the region’s glacial history, in which Greenland has been alternately exposed to or isolated from the North Pacific via the Bering Strait. A molecular study using 3′ rbcL + 5′ rbcL–S sequences was undertaken to assess the diversity of foliose Bangiales on the west coast of Greenland and rbcL sequences were used to study the Greenland flora in a larger phylogenetic and floristic context. New and historic collections document seven species in four genera from the west coast of Greenland. All species had a close link to North Pacific species, being either conspecific with them or North Atlantic–North Pacific vicariant counterparts.     

PORPHYRA REDIVIVA SP. NOV. (RHODOPHYTA): A NEW SPECIES FROM NORTHEAST PACIFIC SALT MARSHES1

A new species, Porphyra rediviva (Bangiales, Rhodophyta), is described from the northeast Pacific based on morphological, cytological, reproductive, ecological, and molecular characters. This species occurs at high intertidal levels in salt marshes along the coasts of Washington, Oregon, and northern California and exhibits a growth optimum at reduced salinity. It is further distinguished by a distinct demarcation between male and female sectors of the gametophytic thalli of epilithic specimens. The species is found most commonly in the drift or trapped in Salicornia beds, but these detached blades never have been found with sporangia or gametangia. Molecular analyses using restriction fragment length polymorphism patterns of polymerase chain reaction–amplified ribosomal DNA (rDNA) show that this salt marsh Porphyra is conspecific throughout its range and is distinct from other Pacific Porphyra species with similar reproductive patterns. Based on molecular data, P. rediviva is related most closely to P. purpurea from the North Atlantic. Fixed rDNA polymorphisms between the two taxa, however, support ecological and cytological evidence that they should be considered different species.     

A review of the bladed Bangiales (Rhodophyta) in China: history,culture and taxonomy

In this paper, we review cultural history, mariculture and taxonomic work to date for Porphyra sensu lato (bladed Bangiales) in China. The bladed Bangiales are a red seaweed group with high species biodiversity and economic value. In China, species occur along the length of the coast and are highly integrated into the country’s culture. Chinese people have used the bladed Bangiales as food and pharmaceuticals for about 1700 years with many references to these seaweeds in ancient books. The mariculture of bladed Bangiales in China also has a long history and an industry has been established based on some species, notably Pyropia yezoensis. The scientific study of the taxonomy of the bladed Bangiales in China began in the late 1920s and to date, based on morphological identification, 25 species and five varieties have been recorded for China, of which 12 species are considered to be endemic to the country. The majority of species have distribution data showing evidence of possible changes due to increasing water temperatures along the coast. The global biodiversity of the bladed Bangiales has been revealed using molecular approaches. This points to the need for molecular taxonomy of Chinese material to document species diversity and distribution, particularly as it includes the wild stocks for seaweed cultivation and because coastal habitats are increasingly impacted by the increasing human population and an expanding mariculture industry. There is a considerable body of literature on the bladed Bangiales in China, but much of it is Chinese and in obscure publications, so we review it here for the benefit of readers worldwide.     

Potential effects of climate change on plant species in the Faroe Islands

Aim To identify the effect of climate change on selected plant species representative of the main vegetation types in the Faroe Islands. Due to a possible weakening of the North Atlantic Current, it is difficult to predict whether the climate in the Faroe Islands will be warmer or colder as a result of global warming. Therefore, two scenarios are proposed. The first scenario assumes an increase in summer and winter temperature of 2 °C, and the second a decrease in summer and winter temperature of 2 °C. Location Temperate, low alpine and alpine areas in the northern and central part of the Faroe Islands. Methods The responses of 12 different plant species in the Faroe Islands were tested against measured soil temperature, expressed as T_min, T_max, snow cover and growing degree days (GDD), using generalised linear modelling (GLM). Results The tolerance to changes in winter soil temperature (0.3–0.8 °C) was found to be lower than the tolerance to changing summer soil temperature (0.7–1.0 °C), and in both cases lower than the predicted climate changes. Conclusions The species most affected by a warming scenario are those that are found with a limited distribution restricted to the uppermost parts of the mountains, especially Salix herbacea, Racomitrium fasciculare, and Bistorta vivipara. For other species, the effect will mainly be a general upward migration. The most vulnerable species are those with a low tolerance, especially Calluna vulgaris, and also Empetrum nigrum, and Nardus stricta. If the climate in the Faroe Islands should become colder, the most vulnerable species are those at low altitudes. A significantly lower temperature would be expected to produce a serious reduction in the extent of Vaccinium myrtillus and Galium saxatilis. Species like Empetrum nigrum, Nardus stricta, and Calluna vulgaris may also be vulnerable. In any case, these species can be expected to migrate downwards.     

A preliminary investigation of the order Bangiales (Bangiophycidae, Rhodophyta) based on sequences of nuclear small-subunit ribosomal RNA genes

We investigated phylogenetic relationships among red algae of the order Bangiales by analysis of sequences of the nuclear gene encoding cytosolic small-subunit ribosomal RNA in Bangia atropurpurea (Roth) C. Ag. and eight samples representing seven species of Porphyra. The ssu-rDNA range from 1818 to 1845 nucleotides in length, with guanosine plus cytosine ratios between 47.0% and 48.6%. A group IC1 intron occurs in the B atropurpurea ssu-rDNAs at the same position as in P. spiralis var. amplifolia Oliveira Filho et Coll and several other eukaryote ssu-rDNAs. The nine sequences form a stable monophyletic group upon phylogenetic analysis. The ssu-rDNA from B. atropurpurea nests stably within the Porphyra group and is closely related to P. amplissima (Kjellm.) Setchell et Hus in Hus, making the genus Porphyra paraphyletic. No correlation is seen between phylogenetic position and number of cell layers in the Porphyra thallus. We discuss possible taxonomic and evolutionary implications of these observations.     

Confirmation of cultivated Porphyra tenera (Bangiales, Rhodophyta) by polymerase chain reaction restriction fragment length polymorphism analyses of the plastid and nuclear DNA

Polymerase chain reaction restriction fragment length polymorphism (PCR‐RFLP) analysis of the plastid ribulose‐1,5‐bisphosphate carboxylase (RuBisCo) spacer region was developed for a more reliable and rapid species identification of cultivated Porphyra in combination with PCR‐RFLP analysis of the nuclear internal transcribed spacer (ITS) region. From the PCR‐RFLP analyses of the plastid and nuclear DNA, we examined seven strains of conchocelis that were used for cultivation as Porphyra tenera Kjellman but without strict species identification. The PCR‐RFLP analyses suggested that two strains, C‐32 and 90‐02, were cultivated P. tenera and that the other five strains, C‐24, C‐28, C‐29, C‐30 and M‐1, were Porphyra yezoensis f. narawaensis Miura. To identify species more accurately and to reveal additional genetic variation, the two strains C‐32 and 90‐02 were further studied by sequencing their RuBisCo spacer and ITS‐1 regions. Although RuBisCo spacer sequences of the two strains were identical to each other, each of their ITS‐1 sequences showed a single substitution. The sequence data again confirmed that the two strains (C‐32 and 90‐02) were cultivated P. tenera, and suggested that the two strains showed some genetic variation. We concluded that PCR‐RFLP analysis of the plastid and nuclear DNA is a powerful tool for reliable and rapid species identification of many strains of cultivated Porphyra in Japan and for the collection of genetically variable breeding material of Porphyra.     

Research Note: Simple molecular discrimination of cultivated Porphyra species (Porphyra yezoensis and Porphyra tenera) and related wild species (Bangiales,Rhodophyta)

To discriminate between cultivated Porphyra species (Porphyra yezoensis and Porphyra tenera) and closely related wild Porphyra species, we developed a polymerase chain reaction‐restriction fragment length polymorphism (PCR‐RFLP) analysis of the rbcL gene using five restriction enzymes. Although our previous PCR‐RFLP analyses of internal transcribed spacer (ITS) rDNA and plastid RuBisCO spacer regions could not always discriminate wild P. yezoensis, wild P. tenera, and closely related wild species, the PCR‐RFLP profiles of the rbcL gene were useful in discriminating samples collected from natural habitats. Therefore, PCR‐RFLP analysis of the rbcL gene will help in the simple identification of a large number of samples, not only for the establishment of reliable cultures as breeding material, but also for the taxonomic investigations of species that are closely related to cultivated Porphyra.     

An evaluation of species relationships in the Porphyra perforata complex (Bangiales,Rhodophyta) using starch gel electrophoresis

Traditional morphological features have formed the basis for distinguishing species of Porphyra. Among these features are number of cell layers, number of chloroplasts per cell, arrangement of reproductive structures on the thallus, and overall morphology. Chromosome number and chromosome morphology have helped corroborate some species identities. A survey of northeast Pacific species of Porphyra using starch gel electrophoresis of 15 soluble proteins has shown that electrophoretic banding patterns provide a reliable diagnostic tool for species identification. Data from starch gel electrophoresis are presented to confirm the identities of species formerly associated with the Porphyra perforata species-complex in British Columbia and northern Washington. Porphyra abbottae, P. fallax, P. kanakaensis, and P. torta are recognized as distinct species, and Porphyra sanjuanensis is synonymized with P. perforata.     

Characterizing the microbial culprit of white spot disease of the conchocelis stage of Porphyra yezoensis (Bangiales, Rhodophyta)

White spot disease is the most frequent and harmful disease affecting the shell-boring conchocelis stage of the economically important red alga Porphyra yezoensis (= Pyropia yezoensis (Ueda) Hwang & Choi) (Bangiales, Rhodophyta). To date, its potential pathogens and pathogenesis remain poorly understood. In this study, we isolated 7 bacteria, 26 fungi, and 10 actinomycetes from sick conchocelis. Re-infection experiments revealed that only one fungus, GF014, caused disease. Morphological characteristics of the colony, mycelium, sporangium, and spore of GF014 suggested that the strain belongs to the order Pleosporales and the genus Phoma. Classification based on internal transcribed spacer (ITS) sequences supported the results of morphological identification. These results indicate that GF014 is a species of Phoma and a specific pathogen for Porphyra. GF014 preferred organic nitrogen sources, and 20–28 °C and low salinity water were its optimal living conditions. Our findings will play an active role in preventing and treating white spot disease in P. yezoensis.     

GENETIC DIVERSITY AND INTROGRESSION IN TWO CULTIVATED SPECIES (PORPHYRA YEZOENSIS AND PORPHYRA TENERA) AND CLOSELY RELATED WILD SPECIES OF PORPHYRA (BANGIALES,RHODOPHYTA)1

We investigated the genetic variations of the samples that were tentatively identified as two cultivated Porphyra species (Porphyra yezoensis Ueda and Porphyra tenera Kjellm.) from various natural populations in Japan using molecular analyses of plastid and nuclear DNA. From PCR‐RFLP analyses using nuclear internal transcribed spacer (ITS) rDNA and plastid RUBISCO spacer regions and phylogenetic analyses using plastid rbcL and nuclear ITS‐1 rDNA sequences, our samples from natural populations of P. yezoensis and P. tenera showed remarkably higher genetic variations than found in strains that are currently used for cultivation. In addition, it is inferred that our samples contain four wild Porphyra species, and that three of the four species, containing Porphyra kinositae, are closely related to cultivated Porphyra species. Furthermore, our PCR‐RFLP and molecular phylogenetic analyses using both the nuclear and plastid DNA demonstrated the occurrence of plastid introgression from P. yezoensis to P. tenera and suggested the possibility of plastid introgression from cultivated P. yezoensis to wild P. yezoensis. These results imply the importance of collecting and establishing more strains of cultivated Porphyra species and related wild species from natural populations as genetic resources for further improvement of cultivated Porphyra strains.     

Origin and dispersal of Potamophylax cingulatus (Trichoptera: Limnephilidae) in Iceland

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