Phenology of Sirodotia suecica (Batrachospermaceae, Rhodophyta) in a high-altitude stream in central Mexico

The morphology and phenology of a monoecious population of Sirodotia suecica Kylin was evaluated seasonally in a central Mexican fifth-order high-altitude stream. Abundance of gametophytes is positively correlated with concentrations of total dissolved solids, specific conductivity and total ionic concentration; being present the whole dry season and disappearing in the rainy season. The gametophytes grew in eutrophic circumstances and particular microhabitat conditions: high current velocity (66–122 cm s⁻¹), low irradiance (75–263 µmol photons m⁻² s⁻¹) and shallow depth (7–26 cm). Percent cover of gametophytes ranged from 5–90% and significant differences in abundance were not observed when alga was present. Some morphological and reproductive characteristics seem to be adaptations to high current velocity: abundant secondary branches, spermatangia and carpogonia. 'Chantransia' stage, microscopic creeping filaments associated with the base of the gametophyte, were not observed in natural conditions. In terms of reproductive success, the population studied can be regarded as highly efficient, considering the high fertilized carpogonia rate, similar to monoecious populations in lotic habitats. However S. suecica was not common in the study region because it was restricted to particular microhabitat conditions.     

Phenology of Paralemanea mexicana (Batrachospermales,Rhodophyta) in a high‐altitude stream in central Mexico

The morphology and phenology of Paralemanea mexicana (Kützing) Vis et Sheath was evaluated seasonally in a fifth order high‐altitude stream in central Mexico. The gametophytes grew during oligotrophic and eutrophic conditions, and during particular microhabitat conditions: high current velocity (40–240 cm s^?1), low to medium irradiance (5–973 μmol photons m^?2 s^?1), and shallow depth (1–30 cm). The abundance of gametophytes was positively correlated with low temperature, high current velocity and concentrations of soluble reactive phosphorus. Interestingly, monoecious gametophytes formed two types of branches, true branches with sympodial pattern originating from meristematic cells in variable number, and abundant false branches produced during the development of the ‘Chantransia’ stage in the surface of the gametophyte or by uniseriate filaments arising within the thallus lumen. These filaments generally produce gametophytes and suggest that they could support the germination of carpospores. The frequently whorled branches are the result of a false branching pattern and are exclusive to P. mexicana. These morphological and reproductive characteristics appear to be biomechanical adaptations to avoid detachment and increase reproductive success. Efficient reproductive strategies of P. mexicana observed in this study can be interpreted as adaptations to successfully colonize streams; however, these features may not have been common in the study region due to restricted microhabitat conditions and geographic isolation.     

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.     

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.     

Systematics and distribution of Thorea (Thoreaceae, Rhodophyta) from central Mexico and south-eastern Brazil

Thirteen populations of Thorea were analyzed from central Mexico and south‐eastern Brazil. All populations were considered as belonging to a single species [Thorea hispida (Thore) Desvaux], with wide variation of morphological features. Secondary branches varying in frequency were observed in several populations with an overlapping in the range of branch density for Thorea violacea Bory and T hispida (0–9 and 11–41 per 30 mm, respectively). As this is the most distinguishing character and on the basis of the overlapping (within a same population or even a single plant), we regarded T. violacea as a synonym of T hispida.‘Chantransia’ stage in culture, as well as gametophyte and carposporophyte were described in detail. We confirmed tine coexistence of asexual monosporangia with sexual reproductive structures (carpogonia and spermatangia) and carposporangia. Size, content, arrangement and chromosome number were the most distinctive characteristics among spermatangia, carposporangia and monosporangia. Monosporangia can be promptly differentiated from spermatangia by their granulated content and larger size but they are similar to carposporangia in shape and size; however, monosporangia are not arranged in fascicles. Structures resembling bisporangia were observed in female plants of some populations. Chromosome numbers were n = 4 for spermatangia and fascicle cells, and 2n ca 8 for gonimoblast filaments, carpospores and the ‘Chantransia’ stage cells. The populations of Thorea from central Mexico and south‐eastern Brazil corroborated the known world distribution for T. hispida, consisting dommantly of tropical to subtropical rainforests, sometimes extending into warm temperate areas. Thorea hispida occurred in warm (temperature 17.6–28.O°C), neutral to alkaline (pH 7.0–8.0), high ion content (specific conductance 59–2140 μS cm^?1), moderate flowing (current velocity 17–43 cm/s) and shallow waters (depth <50cm); these data are essentially similar to previous reports.     

Organization of plastid genomes in the freshwater red algal order Batrachospermales (Rhodophyta)

Little is known about genome organization in members of the order Batrachospermales, and the infra‐ordinal relationship remains unresolved. Plastid (cp) genomes of seven members of the freshwater red algal order Batrachospermales were sequenced, with the following aims: (i) to describe the characteristics of cp genomes and compare these with other red algal groups; (ii) to infer the phylogenetic relationships among these members to better understand the infra‐ordinal classification. Cp genomes of Batrachospermales are large, with several cases of gene loss, they are gene‐dense (high gene content for the genome size and short intergenic regions) and have highly conserved gene order. Phylogenetic analyses based on concatenated nucleotide genome data roughly supports the current taxonomic system for the order. Comparative analyses confirm data for members of the class Florideophyceae that cp genomes in Batrachospermales is highly conserved, with little variation in gene composition. However, relevant new features were revealed in our study: genome sizes in members of Batrachospermales are close to the lowest values reported for Florideophyceae; differences in cp genome size within the order are large in comparison with other orders (Ceramiales, Gelidiales, Gracilariales, Hildenbrandiales, and Nemaliales); and members of Batrachospermales have the lowest number of protein‐coding genes among the Florideophyceae. In terms of gene loss, apcF, which encodes the allophycocyanin beta subunit, is absent in all sequenced taxa of Batrachospermales. We reinforce that the interordinal relationships between the freshwater orders Batrachospermales and Thoreales within the Nemaliophycidae is not well resolved due to limited taxon sampling.     

A re-examination of the morphology, ultrastructure and classification of genera in the Lemaneaceae (Batrachospermales, Rhodophyta)

Vegetative and reproductive morphology and ultrastructure were examined for the three genera of the freshwater red algal family Lemaneaceae: Lemanea (two species, seven populations), Paralemanea (two species, three populations) and Psilosiphon (one species, one population). Psilosiphon is readily distinguished from the other two genera in having an outer cortex composed of well-defined filaments interconnected with a dense medulla (both cell types being little vacuolated), spores cleaving off obliquely, putative spermatangia scattered on the thallus surface and reproduction by adventitious filaments. Based on the distinctness of this genus (and corroborated by molecular phylogenies in preparation), a new family is described, the Psilosiphonaceae. Lemanea and Paralemanea appear to be closely related, with an outer cortex that has cells of increasing size and vacuolation from the periphery to the interior and not in obvious rows, a central lumen that contains few ray cells but no medullary filaments, sexual reproduction with spermatangia in distinct clusters, and carpospores in chains, some of which can germinate in the thallus lumen. Lemanea and Paralemanea have distinguishing characteristics that the other genus does not: for Lemanea, hair cells, ray cells abutting the outer cortex and spermatangia in patches; for Paralemanea, no hair cells or ray cells appressed to the outer cortex, inner cortical filaments surrounding the central axis and spermatangia in rings.     

中国连珠藻属(串珠藻科)的分类研究

记述了中国产连珠藻属Sirodotia Kylin的6个种,即黄山连珠藻S.huangshanensis Z．X．Shi et S．L.Xie,威拉连珠藻S．huillensis(Welwitsch ex W．et G．S．West)Skuja,中华连珠藻S.sinica Jao,纤细连珠藻S.delicatula Skuja,瑞典连珠藻S.suecica Kylin和细连珠藻S．tenuissima(Collins)Skuja ex Flint。其中,黄山连珠藻S．huangshanensis Z．X．Shi et S．L.Xie为新种,威拉连珠藻S.huillensis(Welwitsch ex W．et G．S．West)Skuja,纤细连珠藻S．delicatula Skuja和瑞典连珠藻S.suecica Kylin为中国新记录。     

Temperature effects on photosynthesis in gametophytic and sporophytic stages of the freshwater red alga Sirodotia delicatula (Rhodophyta,Batrachospermales) under a global warming perspective

The increase in Earth's global mean temperature due to human influence has become a major concern, with ecological consequences already being reported around the world. In tropical streams, benthic red algae belonging to the order Batrachospermales contribute to a large fraction of the community energy input and play an important role in sustaining autotrophic food webs. The Batrachospermales also have a unique life history, with heteromorphic life stages (gametophytes and sporophytes) occurring seasonally and with distinct ecological roles. Through “in vivo” chlorophyll fluorescence and dissolved oxygen evolution techniques, we evaluated the effects of projected temperature increases of two future scenarios (RCP 4.5 and RCP 8.5) on the photosynthetic response of Sirodotia delicatula gametophytes and sporophytes. In general, both chlorophyll fluorescence and dissolved oxygen showed that the RCP 8.5 scenario could severely jeopardize the photosynthetic performance of the sporophytes during the summer while also providing a stressful physiological situation to the gametophytes. Due to the crucial role of sporophytes in the development of this species, results indicate that this predicted scenario could affect this species. Given the position of these organisms at the base of the food web in tropical lotic environments, especially in shaded low‐order streams, such impact could generate unforeseen cascade effects on higher trophic levels.     

Phenology of a dioecious population of Batrachospermum delicatulum (Batrachospermales, Rhodophyta) in a stream from southeastern Brazil

The seasonal dynamics of a dioecious population of Batrachospermum delicatulum (Skuja) Necchi and Entwisle was evaluated biweekly during the growth period of the macroscopic gametophyte, from late autumn (May) to early spring (October) in a third-order stream from the northeast region of Sao Paulo State, southeastern Brazil (20?43'S, 49?13'W). The population fluctuated throughout the study period in terms of percentage cover, frequency and chlorophyll content. Percent cover and frequency showed a clear pattern with the lowest values at the initial and final stages of the growth period and the highest in July-August (winter). Gametophyte growth was associated with high illumination and low temperature, which agrees with most observations on Batrachospermales in stream environments. Relatively wide variations in reproductive characteristics were observed during the growth period, which were not correlated with percentage cover, frequency and chlorophyll content. The phenologic pattern observed in this population was characterized by a synchronic development of female/male plant ratio with the number of spermatangia per plant resulting in: (i) a higher proportion of fertilized (carpo-sporophytic) plants associated with more male plants; (ii) higher fertilization rates during the periods with higher production of spermatangia per plant; and (iii) higher production of spermatangia per plant when the population had more male plants. These characteristics were largely associated with water temperature and ion content. In terms of reproductive success, the population studied can be regarded as highly efficient, considering the relatively low proportion of male to female plants and high fertilization rates. These data are consistent with a hypothesis to explain high fertilization rates in dioecious species in lotic habitats consisting of outcrossing among adjacent plants with intermingled male and female branches within an algal cluster. Efficient reproductive strategies have been reported in dioecious, monoecious and even mixed populations of B. delicatulum and can be interpreted as adaptations to successfully colonize streams with different characteristics.     

Daily dynamics of photosynthesis of the freshwater red alga Sirodotia delicatula (Batrachospermales,Rhodophyta)

The daily course of photosynthetic parameters of a population of the freshwater red alga Sirodotia delicatula from São Paulo State, Brazil (20°43′24″S, 49°18′21″W) was investigated under natural and laboratory conditions using dissolved oxygen and in vivo chlorophyll fluorescence techniques. Field specimens in laboratory conditions showed a defined daily pattern for net photosynthesis (NP) with two peaks observed in marine macroalgae and some freshwater red algae: the first (the highest) during the morning, and the second (the lowest and less evident) during the afternoon. Values of electron transport rate did not show a clear pattern of daily variation. NP results suggest the existence of an endogenous rhythm controlling photosynthesis. The study under natural conditions in two contrasting periods (autumn (June) and spring (October)) showed that the daily course of effective and potential quantum yield values was negatively correlated with irradiance and values were similar in the beginning and end of the day. These data evidenced, respectively, high excitement pressure on photosystem II and good recovery capacity (with lower values in spring) and a lack of irreversible photodamage to photosynthetic apparatus due to the prolonged exposure to high irradiances. Non‐photochemical quenching values were also negatively correlated with the irradiance, suggesting a low dissipation capacity of excess energy absorbed by reaction centers. The results evidenced a typical pattern of daily variation with evident response to irradiance.     

中国淡水红藻一个新记录属——托氏藻属

报道了中国淡水红藻一个新记录属——托氏藻属Tuomeya Harvey。标本采于江苏徐州。该属以植物体软骨质, 多层细胞紧密排列组成坚固的皮层, 果胞枝弯曲等特征, 区别于串珠藻科Batrachospermaceae的其他属。除对此种藻类的形态作详细描述外, 还通过rbcL序列比较分析其与北美地区样本的地理差异。该属先前在加拿大、美国和南非有过报道。     

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...     

Genetic variation in Batrachospermum helminthosum (Batrachospermales, Rhodophyta) among and within stream reaches using intersimple sequence repeat molecular markers

Intersimple sequence repeat molecular markers were utilized to investigate the genetic relationship among individual gametophyte thalli of the freshwater red alga Batrachospermum helminthosum among distant stream reaches and within a stream reach. Fifteen thalli per stream reach were sampled from 11 streams throughout the known distribution in eastern North America from three locations in Ohio, and one location in Indiana, Michigan, North Carolina, Tennessee, Louisiana, Rhode Island, Massachusetts and Connecticut. The pairwise φ_ST analysis showed significant genetic differences (P<0.05) among all streams. The partitioning of genetic variation was almost equal within and among streams (45% and 55%, respectively). Genetic variation among populations did not reflect geographic distance, suggesting that long distance dispersal may be important in the distribution of this alga. Fifty‐eight individuals from Five Mile Creek, Ohio, were investigated from three distinct segments of the stream (upstream, middle and downstream) to examine small‐scale genetic variation within a stream reach. A much higher proportion of genetic variation was observed within a stream segment (79%) than among the three segments (21%). Among these stream segments within Five Mile Creek, the individuals were variously related, with genetic similarity and spatial distance uncorrelated. A similar result was obtained among individuals within other stream reaches, suggesting that genetic structure within a stream reach may be more complex than just reproduction among closely located individuals.     

A re-examination of the morphology and reproduction of Nothocladus lindaueri (Batrachospermales, Rhodophyta)

The vegetative morphology and reproduction of the freshwater rhodophyte Nothocladus lindaueri Skuja [=Batrachospermum lindaueri (Skuja) Necchi et Entwisle] were examined by light and electron microscopy. It was confirmed that this alga has a typical batrachospermalean pit plug with two cap layers, the outer one of which is domed. During elongation of hair cells, the primary wall is broken, forming a basal collar. Hair cells have a single nucleus and abundant Golgi bodies, en-doplasmic reticula (ER) and vesicles. Dividing apical cells of the fascicles have a nucleus with art adjacent zone of exclusion, the latter containing a single polar ring. Branched trichogynes and fertilized carpogonia are shown for the first time in this species. Carpogonial branch and involucral cells contain a prominent axial nucleus, proplastids, ER and vesicles. The pit plugs disintegrate among these cells leaving open pit connections. Carpogonia have plentiful mitochondria and vesicles. The wall at the trichogyne apex is thickened and densely stained. The carposporophyte centre consists of a mass of fusion cells with open pit connections, and indeterminate gonimoblast filaments arise from this mass. The combination of a symmetrical carpogonial base, a carposporophyte centre consisting of a mass of fusion cells, and exclusively indeterminate gonimoblast filaments appears to be unique among the members of the Batrachospermaceae. The specimen of N. lindaueri contains epiphytic filaments of Audouinella meiospora producing both spermatangia and monosporangia. Spermatium formation in N. lindaueri remains unknown.     

Variation in gametophyte dominance in populations of Chondrus verrucosus (Gigartinaceae,Rhodophyta)

We describe the abundance, including spatial and temporal variability, of phases of the isomorphic Chondrus verrucosus Mikami from Japan. Chondrus verrucosus occurred in a dense (～90% cover) and temporally stable bed on a small, isolated rocky outcrop (Oyakoiwa) in Shizuoka Prefecture. Small vegetative fronds were always much more abundant than large vegetative and fertile fronds over the spring to late summer periods in 1999 and 2000. Over the same period, fertile carposporophytic fronds were generally more abundant than fertile tetrasporophytic fronds, and fertile male fronds appeared infrequently at low densities. Using the resorcinol‐acetal test, we determined the proportion of gametophytes and tetrasporophytes in three populations of C. verrucosus: Oyakoiwa and Noroshi (Shizuoka) in the summers of 1999 and 2000 and Kamehana Point (Miyagi) in autumn 2000. All populations had a significantly higher proportion of gametophytes than tetrasporophytes in both years, although gametophytic proportions were lower at Noroshi (～70%) than at Oyakoiwa (～80%) and Kamehana Point (～97%). However, examination of all isolated individuals sampled on Noroshi showed equal proportions of each phase in 1999, but gametophyte dominance (74%) in 2000. Differences in dispersal and spore production between phases are discussed as mechanisms potentially contributing to variation in gametophyte dominance.     

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.     

New features in the vegetative and reproductive morphology of Cystoclonium purpureum (Cystocloniaceae,Rhodophyta) from the North Atlantic Ocean

The aim of this study was to evaluate whether the success of invasive Caulerpa cylindracea (Caulerpa racemosa var. cylindracea) in Mediterranean habitats may be linked to particular nutritional requirements and/or nutrient accumulation/reallocation dynamics within the algal thallus. To this end, a manipulative field experiment was carried out transplanting C. cylindracea and supplying nutrients to subtidal rocky macroalgal assemblages. Morphological features of C. cylindracea were measured, while percentage cover and tissue nitrogen, carbon and phosphorus were determined for C. cylindracea and two co-occurring species (the introduced Laurencia chondrioides and native Flabellia petiolata) in nutrient-enriched and unenriched conditions. Growth of C. cylindracea was greater in the nutrient-enriched plots while no effect of nutrient enrichment was detected in co-occurring macroalgae. A particularly high requirement for nitrogen has been observed in C. cylindracea, but without severe N-limitation, while a general nutritional state of P-limitation was detected in both invasive and co-occurring macroalgae. The absence of inhibition in hypertrophic conditions and under high levels of ammonia together with a strong uptake and quick reallocation of nutrients within the coenocytic thallus probably represent the main advantages over the other co-occurring species which could explain part of the success of C. cylindracea in the Mediterranean Sea.     

Phylogenetic relationships within the tribe Janieae (Corallinales,Rhodophyta) based on molecular and morphological data: a reappraisal of Jania1

Generic boundaries among the genera Cheilosporum, Haliptilon, and Jania—currently referred to the tribe Janieae (Corallinaceae, Corallinales, Rhodophyta)—were reassessed. Phylogenetic relationships among 42 corallinoidean taxa were determined based on 26 anatomical characters and nuclear SSU rDNA sequence data for 11 species (with two duplicate plants) referred to the tribe Corallineae and 15 species referred to the tribe Janieae (two species of Cheilosporum, seven of Haliptilon, and six of Jania, with five duplicate plants). Results from our approach were consistent with the hypothesis that the tribe Janieae is monophyletic. Our data indicate, however, that Jania and Haliptilon as currently delimited are not monophyletic, and that Cheilosporum should not be recognized as an independent genus within the Janieae. Our data resolved two well‐supported biogeographic clades for the included Janieae, an Indian‐Pacific clade and a temperate North Atlantic clade. Among anatomical characters, reproductive structures reflected the evolution of the Janieae. Based on our results, three genera, Cheilosporum, Haliptilon, and Jania, should be merged into a single genus, with Jania having nomenclatural priority. We therefore propose new combinations where necessary of some species previously included in Cheilosporum and Haliptilon.     

Large subunit rDNA and rbcL gene sequence data place Petrohua bernabei gen. et sp. nov. in the Batrachospermales (Rhodophyta), but do not provide further resolution among taxa in this order

The phylogenetic relationship among 12 previously described batrachospermalean taxa and a novel member of the order were investigated using the LSU and rbcL genes separately and in combination. The primary goal of this research was to establish the phylogenetic placement of a previously undescribed freshwater red alga from Chile. The results showed that the new entity with pseudoparenchymatous tube morphology is a member of the Batrachospermales and Petrohua bernabei gen. et sp. nov. is described herein. This is the first record to our knowledge of a Lemanea‐like alga from Chile. It would appear that this thallus construction has evolved at least three times in the Batrachospermales and that the switch from a Batrachospermum‐like construction to a pseudoparenchymatous construction may be a repeated adaptive response to turbulent waterfall habitats. In addition to providing information about a new freshwater red alga, this study sought to determine whether combining the data from two genes would produce a more robust phylogeny, particularly for intermediate nodes, to resolve familial relationships within the order. As with previous analyses, the Batrachospermales was resolved as a clade and support was high for relationships resolved among relatively recent nodes. Unfortunately, combining the LSU and rbcL data did not have the desired effect of more fully resolving intermediate nodes among the Batrachospermales.