A new crustose red alga Peyssonnelia rumoiana (Rhodophyta, Gigartinales) from Japan

The marine red alga Peyssonnelia rumoiana Kato et Masuda, sp. nov. (Peyssonneliaceae, Gigartinales) is described from warm‐ and cold‐temperate regions in Japan. It is principally characterized by having hypo‐thallial filaments comprising a polyflabelate layer, proximal perithallial cells arising from the whole upper surface of each hypothallial cell (Peyssonnelia rubra‐type anatomy) and closely packed in a firm matrix, the production of two filaments from the proximal perithallial cell, unicellular rhizoids, appressed crust margins and hypobasal calcification. The alga is distinguished from related species by: (i) its conspicuously elevated cystocarpic (100–150 μm high) and tetrasporangial (80–110 μm high) nemathecia; (ii) tetrasporangia with or without a unicellular pedicel; and (iii) large (25–45 μm in diameter by 70–115 μm in length) tetrasporangia and (iv) the production of double chains of spermatangia (Peyssonnelia harveyana‐type spermatangial development).     

New records of Peyssonnelia armorica and Peyssonnelia harveyana (Rhodophyta, Gigartinales) from Japan

Two species of the crustose red algal genus Peyssonnelia (Gigartinales, Peyssonneliaceae) are reported from Japanese waters for the first time. These species share the following combination of vegetative and reproductive features: thalli with appressed margins, perithallial filaments arising from the whole upper surface of each hypothallial cell (the Peyssonnelia rubra‐type anatomy), unicellular rhizoids, hypobasal calcification and spermatangia that are produced in double chains (the Peyssonnelia harveyana‐type spermatangial filament). However, they differ obviously from each other in the hypothallus orientation as seen from below, the perithallus structure in relation to the consistency of the crust, the origin of gonimoblasts and the elevation of the nemathecia. Peyssonnelia armorica is characterized by: (i) hypothallial filaments comprising a polyflabellate layer; (ii) easily separable perithallial filaments in a gelatinous matrix; (iii) gonimoblasts originating exclusively from the auxiliary cell; and (iv) semi‐immersed (slightly elevated) nemathecia. Peyssonnelia harveyana is characterized by: (i) hypothallial filaments arranged in parallel rows; (ii) closely packed perithallial filaments in a firm matrix; (iii) gonimoblsts originating from both the auxiliary cell and the connecting filament; and (iv) conspicuously elevated nemathecia.     

Phylogeography of the freshwater red alga Sirodotia (Batrachospermales,Rhodophyta) in Brazil

Considering the lack of knowledge on genetic variation on members of the freshwater red algal of the order Batrachospermales in tropical regions, phylogeographic patterns in Sirodotia populations were investigated using two mitochondrial regions: the cox2‐3 spacer and partial cox1 gene (barcode). Individuals identified as Sirodotia delicatula were analyzed from 14 stream segments across its distribution in Brazil. Phylogenetic analyses based on the ribulose‐1,5‐bisphosphate carboxylase/oxygenase large sub‐unit gene showed three clades, one representing S. delicatula, from all locations in southeastern Brazil and other regions from Brazil. The remaining samples formed two clades, which were highly divergent and distantly positioned from those of S. delicatula: 2.5–2.7% and 3.4–3.7%. This level of variation would warrant the species split of these taxa from mid‐western Brazil. A total of eight cox2‐3 spacer and nine cox1 haplotypes were observed among the 122 individuals studied. One location had two cox2‐3 haplotypes and three locations had two cox1 haplotypes; all others had a single dominant haplotype each. The existence of high intraspecific genetic variation among individuals of distinct locations (several haplotypes), but little variation within a location seems to be a pattern for the Batrachospermales. Haplotype networks showed low variation among the haplotypes from southeastern Brazil (10 locations with divergence of 0.3–1.1% for cox2‐3, 0.1–0.3% for cox1) and high variation among the haplotypes from the mid‐west region (four locations, 4.0–9.3% for cox2‐3, 6.2–8.4% for cox1). Thus, the present data clearly suggest the existence of cryptic species in Sirodotia in Brazil.     

Circumscription of species in the genus Sirodotia (Batrachospermales,Rhodophyta) based on molecular and morphological data

Species level taxonomy and phylogeographical distribution patterns in the freshwater rhodophyte Sirodotia are resolved through phylogenetic inferences based on rbcL and cox2–3 sequence data. Previous studies focused on the taxonomy of specific Sirodotia species or the distributions across a limited geographical region. Our molecular phylogenies included samples attributable to five recognized Sirodotia species and include collections from Australia, Brazil, Costa Rica, Canada, Finland, Mexico, New Zealand, South Africa and the United States. Both rbcL and cox2–3 phylogenies inferred S. suecica, S. tenuissima and S. goebelii as a monophyletic group with little sequence divergence. This result supports the synonymy of S. tenuissima and S. goebelii with S. suecica (the species name with priority). Within this clade, samples collected from Australia and New Zealand formed a monophyletic group with no other discernible phylogeographical patterns within S. suecica. This result seems to be somewhat unusual in the Batrachospermales, as other species have shown greater genetic variation among geographically distant locations. As in previous studies, S. huillensis and S. delicatula were inferred as a separate species based on the rbcL phylogeny, supporting the current taxonomy. A specimen of S. aff. huillensis from South Africa, may represent a new species but further research is necessary before it can be designated as such.     

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...     

Ochtodes searlesii sp. nov. (Gigartinales,Rhodophyta), from the Pacific tropical coast of Mexico,based on morphological and molecular evidence

Ochtodes searlesii Mendoza‐González, Mateo‐Cid et Sentíes sp. nov. is described from Michoacán, tropical Mexican Pacific, on the basis of comparative morphology and rbcL sequence analysis. It is distinguished from other Ochtodes species by its erect axes arising from an encrusting base, its small terete fronds, regularly dichotomously branched axes, and obliquely divided zonate tetrasporangia. Phylogenetic analyses showed that three Pacific Mexican samples, from Caletilla, Zapote and La Majahuita (Michoacán), were identical and formed a distinctive and well supported Clade segregated from other species of Ochtodes from Brazil, Cuba, Ecuador, Guadeloupe and Mexico. The Mexican entity is morphologically distinct from other Ochtodes species as well. On this basis we propose a new Ochtodes species.     

Reproductive ecology of the freshwater red alga Batrachospermum delicatulum (Batrachospermales, Rhodophyta) in three tropical streams

Batrachospermum delicatulum specimens from three stream segments were analyzed from a tropical region in south‐ eastern Brazil (20°18′– 20°49′S, 49°13′– 49°46′W). Physical and chemical parameters and the spatial placement of thalli were investigated along with the reproductive characteristics of the gametophytic phase. Sequence data of the cox 2‐ 3 spacer region was also utilized to evaluate genetic variation in individuals within and among stream segments. Gametophyte occurred under relatively diverse environmental conditions, whereas thalli abundance was weakly or not correlated to environmental variables within the stream segments. All specimens examined were dioecious. The ratio of male/female plants was relatively low (0.5 to 1.3) and male plants tended to occur as clumps (two or three plants together). High reproductive success was observed, as indicated by the occurrence of 100% fertilized (carposporophytic) female plants. This is similar to previous reports for this and other dioecious species, which is remarkable considering the relatively low proportion of male/female plants. Results support the two hypotheses to explain the high reproductive success in dioecious species. The occurrence of male plants in clumps was evidence for a strict spatial relationship (i.e. male plants located in upstream position of female plants in order to release spermatia, which would be carried by eddies through female plants). In contrast, the occurrence of male and female plants adjacent to each other allowed outcrossing among neighboring plants with intermingled male and female branches, which seemed more applicable to some situations (low turbulence habitats). The cox 2‐ 3 spacer region from the 18 individuals sequenced was 376 bp and the DNA sequence was identical with no base pair substitutions. Likewise, a previous study of another Batrachospermum species showed that the same haplotypes were present in all stream segments from the same drainage basin, even though the stream segments were a considerable distance apart. Short distance dispersal either by small birds or waterway connectivity might explain these findings.     

Phylogeography of the freshwater red alga B atrachospermum viride‐brasiliense (Rhodophyta,Batrachospermales)

Phylogeography of B atrachospermum viride‐brasiliense was investigated using two mitochondrial regions: the cox2‐3 spacer and the barcode region of cox1 gene. Eighty‐seven individuals were analyzed from nine stream segments throughout its distribution in Brazil. Ten cox2‐3 spacer and nine cox1 haplotypes were observed among the individuals studied (87 vs. 43, respectively). Divergences among haplotypes were relatively low (≤2.4% for cox2‐3 and ≤1.8% for cox1). Most locations have a single haplotype, whereas only two locations had two haplotypes for both markers. The haplotype network for cox2‐3 showed a phylogeographic trend from the south towards the southeast with haplotypes from the southeast more closely related. For cox1 a trend from the southeast spreading towards the south and north was revealed, with the southern haplotypes more closely associated. Results clearly indicated that B . viride‐brasiliense represents a single species and the phylogeographic pattern consisted of a closely connected group of haplotypes from southern and southeastern Brazil. Levels of intra‐ and inter‐population variation were similar for the two markers with slightly higher values for cox2‐3. The trend observed in this study is similar to that in other members of Batrachospermales with little variation within a stream segment (one or two haplotypes) and more distant haplotypes showing higher divergences. This pattern could be attributed to the fact that colonization of a site might be rare by a single event with subsequent proliferation of the population. The geographic distribution of B . viride‐brasiliense was interpreted according to the biogeographic models proposed for South America being limited to three morpho‐climatic domains or biogeographic provinces: tropical Atlantic rainforest, sub‐tropical rainforest and cerrado (Brazilian savannah).     

A morphological and molecular assessment of the genus Prionitis J. Agardh (Halymeniaceae, Rhodophyta)

A critical reassessment of the morphological features of two closely related red algal genera, Grateloupia C. Agardh and Prionitis J. Agardh (Halymeniaceae), shows that members of the two genera share very similar reproductive (including the Grateloupia‐type auxiliary‐cell ampullae) and vegetative characters. Diagnostic features hitherto used for distinguishing these two genera, the texture of blades (lubricous to leathery in Grateloupia vs cartilaginous in Prionitis) and the position of reproductive structures (scattered over the entire blade in Grateloupia vs confined to particular portions of the blade in Prionitis), are continuous across some 75 species of both genera, thus making it difficult to draw a clear‐cut distinction between the two genera. In ribulose‐1,5‐bisphosphate carboxylase/oxygenase gene (rbcL) sequence analyses, the species of Grateloupia and Prionitis, including the two generitypes, constitute a large monophyletic clade in the Halymeniaceae. It is therefore proposed that Prionitis be included in the synonymy under Grateloupia and the appropriate combinations are proposed.     

Delineation of Two Sibling Red Algal Species,Gracilaria Gracilis and Gracilaria Dura (Gracilariales,Rhodophyta), Using Multiple DNA Markers: Resurrection of the Species G. Dura Previously Described in the Northern Atlantic 200 Years Ago1

Molecular markers belonging to the three different genomes, mitochondrial (cox2‐cox3 spacer), plastid (rbcL), and nuclear (internal transcribed spacer [ITS] 2 region), were used to compare samples of the two morphologically related species Gracilaria gracilis (Stackh.) Steentoft, L. M. Irvine et Farnham and G. dura (C. Agardh) J. Agardh collected along Atlantic coasts. In northern Europe, the distinction between these two species is ambiguous, and they are currently recognized under the single name of G. gracilis. The low but congruent patterns of genetic divergence observed for markers of the three genomic compartments highly suggest that these two taxa correspond effectively to two different genetic entities as previously described 200 years ago, based on morphological traits. However, thanks to the combination of different DNA markers, occurrence of “incongruent” cytotypes (i.e., mitotypes of G. dura associated with chlorotypes of G. gracilis) in individuals collected from Brittany, suggests interspecific hybridization between the two sibling species studied.     

Patterns of genetic diversity of the cryptogenic red alga Polysiphonia morrowii (Ceramiales,Rhodophyta) suggest multiple origins of the Atlantic populations

The red alga Polysiphonia morrowii, native to the North Pacific (Northeast Asia), has recently been reported worldwide. To determine the origin of the French and Argentine populations of this introduced species, we compared samples from these two areas with samples collected in Korea and at Hakodate, Japan, the type locality of the species. Combined analyses of chloroplastic (rbcL) and mitochondrial (cox1) DNA revealed that the French and Argentine populations are closely related and differ substantially from the Korean and Japanese populations. The genetic structure of P. morrowii populations from South Atlantic and North Atlantic, which showed high haplotype diversity compared with populations from the North Pacific, suggested the occurrence of multiple introduction events from areas outside of the so‐called native regions. Although similar, the French and Argentine populations are not genetically identical. Thus, the genetic structure of these two introduced areas may have been modified by cryptic and recurrent introduction events directly from Asia or from other introduced areas that act as introduction relays. In addition, the large number of private cytoplasmic types identified in the two introduced regions strongly suggests that local populations of P. morrowii existed before the recent detection of these invasions. Our results suggest that the most likely scenario is that the source population(s) of the French and Argentine populations was not located only in the North Pacific and/or that P. morrowii is a cryptogenic species.     

Delimitating cryptic species in the Gracilaria domingensis complex (Gracilariaceae,Rhodophyta) using molecular and morphological data

Species in the genus Gracilaria that display conspicuously flattened vegetative morphologies are a taxonomically challenging group of marine benthic red algae. This is a result of their species richness, morphological similarity, and broad phenotypic plasticity. Within this group, the Gracilaria domingensis complex is one of the most common, conspicuous, and morphologically variable species along the tropical western Atlantic Ocean. Previous research has identified that members of this complex belong to two distantly related clades. However, despite this increased phylogentic resolution, species delimitations within each of these clades remain unclear. Our study assessed the species diversity within this difficult complex using morphological and molecular data from three genetic markers (cox1, UPA, and rbcL). We additionally applied six single‐marker species delimitation methods (SDM: ABGD, GMYCs, GMYCm, SPN, bPTP, and PTP) to rbcL, which were largely in agreement regarding species delimitation. These results, combined with our analysis of morphology, indicate that the G. domingensis complex includes seven distinct species, each of which are not all most closely related: G. cervicornis; a ressurected G. ferox; G. apiculata subsp. apiculata; a new species, Gracilaria baiana sp. nov.; G. intermedia subsp. intermedia; G. venezuelensis; and G. domingensis sensu stricto, which includes the later heterotypic synonym, G. yoneshigueana. Our study demonstrates the value of multipronged strategies, including the use of both molecular and morphological approaches, to decipher cryptic species of red algae.     

Five Cyanophora (Cyanophorales,Glaucophyta) species delineated based on morphological and molecular data

Cyanophora is an important glaucophyte genus of unicellular biflagellates that may have retained ancestral features of photosynthetic eukaryotes. The nuclear genome of Cyanophora was recently sequenced, but taxonomic studies of more than two strains are lacking for this genus. Furthermore, no study has used molecular methods to taxonomically delineate Cyanophora species. Here, we delimited the species of Cyanophora using light and electron microscopy, combined with molecular data from several globally distributed strains, including one newly established. Using a light microscope, we identified two distinct morphological groups: one with ovoid to ellipsoidal vegetative cells and another with dorsoventrally flattened or broad, bean‐shaped vegetative cells containing duplicated plastids. Our light and scanning electron microscopy clearly distinguished three species with ovoid to ellipsoidal cells (C. paradoxa Korshikov, C. cuspidata Tos.Takah. & Nozaki sp. nov., and C. kugrensii Tos.Takah. & Nozaki sp. nov.) and two species with broad, bean‐shaped cells (C. biloba Kugrens, B.L.Clay, C.J.Mey. & R.E.Lee and C. sudae Tos.Takah. & Nozaki sp. nov.) based on differences in cell shape and surface ornamentations of the vegetative cells under the field‐emission scanning electron microscope. Molecular phylogenetic analyses of P700 chl a apoprotein A2 (psaB) genes and internal transcribed spacer (ITS) regions of nuclear ribosomal DNA (rDNA), as well as a comparison of secondary structures of nuclear rDNA ITS‐2 and genetic distances of psaB genes, supported the delineation of five morphological species of Cyanophora.     

Recent developments in the systematics of the Gigartinaceae (Gigartinales, Rhodophyta) based on rbcL sequence analysis and morphological evidence

The phylogenetic systematics of the Gigartinaceae is discussed for seven genera and three undescribed generic lineages and 65 taxa representing 62 species based on an analysis of rbcL sequences and morphological evidence. An examination of rbcL trees resulting from analyses of these taxa identifies seven lineages: (i) ‘Gigartina’ alveata; (ii) Rhodoglossum/Gigartina; (iii) Chondracanthus; (iv) Ostiophyllum; (v) Sarcothalia; (vi) ‘Gigartina’ skottsbergii; and (vii) a large clade containing Iridaea/‘Sarcothalia’, Mazzaella and Chondrus. These lineages and Chondrus are strongly supported; however, two groups, Iridaea/‘Sarcothalia’ and Mazzaella, receive no bootstrap support. The morphology of the female reproductive system is investigated with the aid of computer-generated, color-coded tracings of photographs of cystocarps seen in cross section at different developmental stages. Seven basic cystocarp types were found which corresponded to species groups seen in rbcL trees. These were: (i) a ‘Gigartina’ alveata group in which the carposporangia-bearing filaments develop apomictically from gametophytic cells; (ii) a Rhodoglossum/Gigartina group in which gonimoblast filaments penetrate the surrounding envelope fusing progessively with envelope cells; (iii) a Chondracanthus group in which gonimoblast filaments penetrate the envelope but fuse with envelope cells only at late developmental stages; (iv) a Sarcothalia group in which the gonimoblast filaments displace an envelope composed mainly of secondary gametophytic filaments and link to envelope cells by terminal tubular gonimoblast cells; (v) an Iridaea group similar to the Sarcothalia group, but with an envelope composed of a mixture of medullary cells and secondary gametophytic filaments; (vi) a Mazzaella group that lacks a true envelope and in which gonimoblast filaments connect to modified gametophytic cells by means of terminal tubular cells; (vii) a Chondrus group in which gonimoblast filaments penetrate the medulla and link to modified medullary cells by means of conjunctor cells forming secondary pit connections. The further separation of these groups into genera is based largely on tetrasporangial characters.     

Stucture and reproduction of Cordylecladia guiryi sp. nov. (Rhodophyta,Rhodymeniales) from the Mediterranean Sea

Abstract

Cordylecladia guiryi sp. nov. is described from the Mediterranean Sea, after a comparison of habit, morphology and reproduction with C. erecta, the only species of the genus. The species is characterized by the small dimension of its axes which arise single from the basal crust and by its habitat, growing epiphytically on Posidonia oceanica Delile leaves. Tetrasporangial and gametangial plants as well as the development of the carposporophyte have been studied in detail. The carpogonial branch is 4-celled and it is formed on a supporting cell that also bears a 3-celled auxiliary branch. Tetrasporangia are cruciately divided. The reproductive details confirm Sparling's view about the great variation existing in the organization of the procarp in members of this order.     

Phylogeny and morphology of the freshwater red alga Nemalionopsis shawii (Rhodophyta,Thoreales) from Nepal

Phylogenetic relationships and morphological characters are presented for a population of Nemalionopsis shawii Skuja collected from Nepal. Molecular data (sequences of rbcL and cox1) were generated and morphological characters were described in detail. The rbcL sequence analyses showed that specimens from Nepal are most similar to N. shawii from Indonesia and Japan and that these entities form a clade with high support (>95% bootstrap and 0.95 posterior probability). The cox1 barcode sequence, however, only had 90.9–91.9% identity with specimens of N. shawii from Hawaii. The rbcL sequence of the specimen from Nepal was positioned in a clade having sequence identity of 99.3–99.7% with three samples: N. shawii from Indonesia and two from Japan identified as N. tortuosa. The comparison of morphological characters of Nemalionopsis from Nepal allowed unequivocal identification with N. shawii. Identifications from previous studies using molecular data were mistaken since most reports of N. shawii are actually of N. tortuosa or vice‐versa. This confusion of names presumably occurred because most specimens previously sequenced were from culture collections or from ‘Chantransia’ stages. Small tufts of ‘Chantransia’ stage were observed growing epiphytically on gametophytes and on the basal system. Carpogonia and spermatangia were fully described in specimens from Nepal. Monosporangia were not observed, whereas carposporangia were unequivocally described for the first time in the genus. An unusual flat strap‐like basal system was observed, interpreted as an additional mode of maintenance in nature under unfavorable environmental conditions.     

Morphology of the crustose coralline alga Pseudolithophyllum muricatum (Corallinales,Rhodophyta) responds to 30 years of ocean acidification in the Northeast Pacific

As the process of ocean acidification alters seawater carbon chemistry, physiological processes such as skeletal accretion are expected to become more difficult for calcifying organisms. The crustose coralline red algae (Corallinales, Rhodophyta) form an important guild of calcifying primary producers in the temperate Northeast Pacific. The morphology of important ecological traits, namely, skeletal density and thallus thickness near the growing edge, was evaluated in Pseudolithophyllum muricatum (Foslie) Steneck & R.T. Paine, the competitively dominant alga within this guild. P. muricatum shows a morphological response to increased ocean acidification in the temperate Northeast Pacific. Comparing historical (1981–1997) and modern (2012) samples from the field, crust thickness near the growing edge was approximately half as thick in modern samples compared with historical samples, while crust calcite density showed no significant change between the two sample groups. Morphological changes at the growing edge have important consequences for mediating competitive interactions within this guild of algae, and may affect the role of crustose coralline algal beds as hosts to infaunal communities and facilitators of recruitment in many invertebrate and macroalgal species.