PHYLOGENY OF SPIROGYRA AND SIROGONIUM (ZYGNEMATOPHYCEAE) BASED ON RBCL SEQUENCE DATA1

DNA sequence data were obtained for the gene encoding the large subunit of RUBISCO (rbcL) from 26 strains of Spirogyra and seven of Sirogonium, using as outgroups 10 genera in the Zygnematales and Desmidiales (Closterium, Cosmarium, Cylindrocystis, Gonatozygon, Mesotaenium, Netrium, Penium, Zygnema, Zygnemopsis, Zygogonium). Sequence data were analyzed using maximum parsimony (MP), maximum likelihood (ML), and Bayesian inference (BI), with bootstrap replication (MP, ML) and posterior probabilities (BI) as measures of support. MP, ML, and BI analyses of the rbcL data strongly support a single clade containing Spirogyra and Sirogonium. The Spirogyra taxa are monophyletic, with the exception of Spirogyra maxima (Hassall) Wittrock, which is nested within a clade with Sirogonium and shares with them the characters of loosely spiraled chloroplasts (<1 complete turn per cell) and anisogamy of gametangial cells; S. maxima differs from Sirogonium in displaying well‐defined conjugation tubes rather than a tubeless connection involving bending (genuflection) of filaments. The ML and BI analyses place this Sirogonium/Spirogyra maxima clade sister to the remaining Spirogyra. Morphological differences among strains of Spirogyra grouped together on the basis of rbcL data, including laboratory strains derived from clonal cultures (Spirogyra communis, S. pratensis), indicate that some characters (filament width, chloroplast number) used in the traditional taxonomy of this group are poor measures of species identity. However, some characters such as replicate end walls and loose spiraling of chloroplasts may be synapomorphies for Spirogyra clades.     

The relationship between Spirogyra (Zygnematophyceae, Streptophyta) filament type groups and environmental conditions in Central Europe

Species identification of the common filamentous green alga Spirogyra is mainly based on the conjugation process and zygospores. However, this genus is mostly found in its vegetative stage, which complicates studies on the ecological demands for individual species. We therefore used a different approach by assessing the relationship between vegetative Spirogyra filament type groups (morphotypes) and environmental conditions (mainly ions, nutrients, light supply and water temperature). Sampling was done at 133 sites in Central Europe and in total 333 different filament types were classified. Spirogyra was found at pH values between 6.2 and 9.1, while total alkalinity ranged from 0.6 to 7.9 mequiv l⁻¹. The genus is colonizing habitats with a specific conductivity between 75 and 1500 μS cm⁻¹. Total phosphorus amounts varied between 1 and 2240 μg l⁻¹ with a median value of 34 μg l⁻¹, indicating meso- to eutrophic conditions as optimal growth range. Filament type grouping by means of cluster analysis was based on cell cross walls (plane or replicate), average cell widths and average chloroplast numbers and resulted in 10 groups with plane cross walls and three with replicate cross walls. Canonical correspondence analysis revealed nutrients to be the key factor for morphotype occurrence: filaments with increased cell widths preferred elevated nutrient conditions. Other environmental variables (ions, buffer capacity, light supply and water temperature) had no significant effects on morphotype occurrence.     

MOLECULAR PHYLOGENY AND REVISION OF THE GENUS NETRIUM (ZYGNEMATOPHYCEAE,STREPTOPHYTA): NUCLEOTAENIUM GEN. NOV.1

Nuclear‐encoded SSU rDNA, chloroplast LSU rDNA, and rbcL genes were sequenced from 53 strains of conjugating green algae (Zygnematophyceae, Streptophyta) and used to analyze phylogenetic relationships in the traditional order Zygnematales. Analyses of a concatenated data set (5,220 nt) established 12 well‐supported clades in the order; seven of these constituted a superclade, termed “Zygnemataceae.” Together with genera (Zygnema, Mougeotia) traditionally placed in the family Zygnemataceae, the “Zygnemataceae” also included representatives of the genera Cylindrocystis and Mesotaenium, traditionally placed in the family Mesotaeniaceae. A synapomorphic amino acid replacement (codon 192, cysteine replaced by valine) in the LSU of RUBISCO characterized this superclade. The traditional genera Netrium, Cylindrocystis, and Mesotaenium were shown to be para‐ or polyphyletic, highlighting the inadequacy of phenotypic traits used to define these genera. Species of the traditional genus Netrium were resolved as three well‐supported clades each distinct in the number of chloroplasts per cell, their surface morphology (structure and arrangement of lamellae) and the position of the nucleus or nuclear behavior during cell division. Based on molecular phylogenetic analyses and synapomorphic phenotypic traits, the genus Netrium has been revised, and a new genus, Nucleotaenium gen. nov., was established. The genus Planotaenium, also formerly a part of Netrium, was identified as the sister group of the derived Roya/Desmidiales clade and thus occupies a key position in the evolutionary radiation leading to the most species‐rich group of streptophyte green algae.     

Identity and phylogenetic placement of Spirogyra species (Zygnematophyceae,Charophyta) from California streams and elsewhere1

Diversity of the filamentous green algae in the genus Spirogyra (Zygnematophyceae) was investigated from more than 1,200 stream samples from California. We identified 12 species of Spirogyra not previously known for California (CA), including two species new to science, Spirogyra californica sp. nov. and Spirogyra juliana sp. nov. Environmental preferences of the Californian species are discussed in the light of their restricted distribution to stream habitats with contrasting nutrient levels. We also investigated the systematic relationships of Spirogyra species from several continents using the chloroplast‐encoded genes ribulose‐1,5‐bisphosphate carboxylase/hydrogenase large subunit (rbcL) and the beta subunit of the ATP synthase (atpB). Californian species were positioned in most major clades of Spirogyra. The phylogeny of Spirogyra and its taxonomic implications are discussed, such as the benefits of combining structural and molecular data for more accurate and consistent species identification. Considerable infraspecific genetic variation of globally distributed Spirogyra species was observed across continental scales. This finding suggests that structurally similar species from distant regions may be genetically dissimilar and that Spirogyra may contain a large number of cryptic species. Correlating the morphological and genetic variation within the genus will be a major challenge for future researchers.     

CONJUGATION IN SPIROGYRA1

Conjugating filaments of Spirogyra were examined with both light and electron microscopes. Initially 2 or more filaments of Spirogyra were attached by mucilagenous material. Papillae appeared first in one filament and then in adjacent positions on the other filament. Subsequent growth of papillae separated the conjugating filaments; wall microtubules disappeared in papillae as they elongated. Golgi activity then increased markedly only in the male filament; mucilage production by these Golgi coincided with contraction of the male gamete from its cell wall and may be responsible for its subsequent migration. The end walls separating papillae dissolved to form the conjugation tube, allowing gamete union. The male protoplast then migrated through the tube and further cytoplasmic condensation formed an elliptical-shaped zygote. During the migration phase, zygote wall formation was initiated and numerous active Golgi apparently contributed material to it. Early zygote maturation was characterized by rapid wall formation and an increase in lipid droplets.     

USING rbcL SEQUENCES TO TEST HYPOTHESES OF CHLOROPLAST AND THALLUS EVOLUTION IN CONJUGATING GREEN ALGAE (ZYGNEMATALES, CHAROPHYCEAE)

Sequences for the Rubisco large subunit (rbcL) gene were used to test hypotheses about the evolution of chloroplast shape and thallus type in genera of two families of conjugating green algae (Zygnematales): the Mesotaeniaceae (saccoderm desmids, mostly unicellular) and the Zygnemataceae (strictly filamentous). Unicellular (u) and filamentous (f) genera exhibit a series of three similar chloroplast shapes: ribbonlike (e.g. Spirotaenia [u], Spirogyra [f], and Sirogonium [f], laminate (e.g. Mesotaenium [u] and Mougeotia [f]), and twin-stellate (e.g. Cylindrocystis [u] and Zygnema [f]. Two conflicting phylogenetic hypotheses have been proposed: 1) families are polyphyletic constructs drawn from three lineages, each with unicellular and filamentous taxa characterized by a specific chloroplast shape; or 2) unicells form one monophyletic lineage (Mesotaeniaceae) and filaments form another (Zygnemataceae), with some chloroplast shapes independently derived. The rbcL data strongly refute hypothesis 2 (monophyly of the two traditional families) and support hypothesis 1 in part. Parsimony, maximum likelihood, and neighbor-joining analyses of the rbcL data strongly support monophyly of a clade containing taxa with ribbonlike chloroplasts and, to a lesser extent, monophyly of a second clade of the four genera with the other two chloroplast shapes. Two saccoderm genera (Roya, curved laminate chloroplasts; Netrium, "cucumber"-shaped chloroplasts) are not members of either of these clades, but they are included in a monophyletic Zygnematales .     

Zygnemataceae in quaternary Columbian sediments

Spores of Zygnemataceae are described from Lower Pleistocene, Upper Pleistocene and Holocene sediments from the Colombian Andes. About ten types could be distinguished, most of them referable to the genera Mougeotia, Spirogyra and Debarya. In addition records of Bulbochaete (Oedogoniaceae) and of Closterium (Desmidiaceae) are reported.Spores of Zygnemataceae appear to be of considerable importance for palaeoecological studies. Debarya aff. D. glyptosperma and several other types were up to now found only in deposits associated with a cold to cool high-mountain climate.     

Light-induced Adhesion of Spirogyra Cells to Glass

Adhesion of Spirogyra (tentatively, Spirogyra fluviatilis) cells to glass is described. The cells of an algal filament can adhere to a substrate only when they are located at the end of the filament. Rapid adhesion is induced by blue-violet light (blue adhesion) as well as by temperature shift (about 6 C → about 22 C) or shaking (dark adhesion). Adherent cells detach in 1 hour in the absence of one of these stimuli. Slow adhesion is induced by red light (red adhesion) 1 hour after irradiation, and may be controlled by phytochrome. A cell once caused to adhere by red light does not release from the glass.     

Two new taxa of Spirogyra

Summary A new species and a new variety ofSpirogyra have been recorded.Spirogyra varshaii spec. nov. has been described and compared to two related species ofSpirogyra. This species has a very distinct subdenticulate to denticulate type of ornamentation of the zygospore wall and differs from the other two related speciesS. chakiense andS. verruculosa in having different dimensions of the zygospores and in the number of chloroplasts. A new variety,Spirogyra chakiense var.lucknowense var. nov. has been described. It differs from the speciesS. chakiense in possessing some differences in the dimensions of the cells and the zygospores from the type species.

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Zusammenfassung Eine neue Spezies und eine neue ArtSpirogyra sind eingetragen worden.Spirogyra varshaii Spec. nov. ist beschrieben worden und mit zwei verwandten Spezies verglichen. Diese Spezies hat einen sehr deutlichen unterzahnigen bis zum zahnigen Typus der gestickten Zygosporeswand, und sie unterscheidet sich von den anderen zwei verwandten SpeziesS. chakiense undS. verruculosa daß sie verschiedene Ausdehnungen der Zygosporen und auch eine verschiedene Zahl der Kromatophoren hat. Eine neue ArtSpirogyra chakiense var.lucknowense var. nov. ist auch beschrieben worden. Sie unterscheidet sich von der SpeziesS. chakiense weil sie einigen Unterschieden in Ausdehnungen der Zellen und auch der Zygosporen hat im Vergleich mit der Typusspezies.

     

Molecular Phylogeny of Conjugating Green Algae (Zygnemophyceae, Streptophyta) Inferred from SSU rDNA Sequence Comparisons

Abstract Nuclear-encoded SSU rDNA sequences have been obtained from 64 strains of conjugating green algae (Zygnemophyceae, Streptophyta, Viridiplantae). Molecular phylogenetic analyses of 90 SSU rDNA sequences of Viridiplantae (inciuding 78 from the Zygnemophyceae) were performed using complex evolutionary models and maximum likelihood, distance, and maximum parsimony methods. The significance of the results was tested by bootstrap analyses, deletion of long-branch taxa, relative rate tests, and Kishino–Hasegawa tests with user-defined trees. All results support the monophyly of the class Zygnemophyceae and of the order Desmidiales. The second order, Zygnematales, forms a series of early-branching clades in paraphyletic succession, with the two traditional families Mesotaeniaceae and Zygnemataceae not recovered as lineages. Instead, a long-branch Spirogyra/Sirogonium clade and the later-diverging Netrium and Roya clades represent independent clades. Within the order Desmidiales, the families Gonatozygaceae and Closteriaceae are monophyletic, whereas the Peniaceae (represented only by Penium margaritaceum) and the Desmidiaceae represent a single weakly supported lineage. Within the Desmidiaceae short internal branches and varying rates of sequence evolution among taxa reduce the phylogenetic resolution significantly. The SSU rDNA-based phylogeny is largely congruent with a published analysis of the rbcL phylogeny of the Zygnemophyceae (McCourt et al. 2000) and is also in general agreement with classification schemes based on cell wall ultrastructure. The extended taxon sampling at the subgenus level provides solid evidence that many genera in the Zygnemophyceae are not monophyletic and that the genus concept in the group needs to be revised.     

SYSTEMATICS OF THE ZYGNEMATACEAE (CHLOROPHYCEAE). III. TRANSEAU COLLECTION OF THE ZYGNEMATACEAE: AN OLD COLLECTION WITH MODERN USES

Edgar Nelson Transeau, botanist-ecologist-phycologist, accumulated a worldwide collection of algal specimens for the compilation of a monograph on the algal family Zygnemataceae, published in 1951. Following his death on January 26, 1960 this Collection remained at The Ohio State University, Columbus until 1978 when it was transferred to the University of Arizona, Tucson. A set of glycerine-preserved specimens on microscope slides is the major item of the Collection with the number of species represented as follows: Spirogyra (138), Mougeotia (52), Zygnema (45), Zygnemopsis (16), Sirogonium (9), Debarya (9), Zygogonium (4), Mougeotiopsis (1) and Entransia (1). All specimens of Pleurodiscus, Sirocladium, Temnogametum and Temnogyra are unidentified. With the Transeau Collection it was demonstrated that an old collection need not be an historical relic, but does have continued usefulness. Potential future use is a powerful justification for maintaining seemingly useless collections whose accumulation often represents the expenditure of untold time, effort and money on the part of many persons and institutions worldwide. Further, as man continues to alter his environment and species are destroyed, collections with long histories become increasingly more valuable. Specimens of the Transeau Collection were subjected to modern techniques, not available earlier, resulting in improved characterization of zygospore-wall structure by scanning electron microscopy and the determination of nuclear DNA by cytofluorometry.     

Spiral cell wall growth in zygnemataceae

Using two species ofSpirogyra and one species ofZygnema, it was demonstrated on a quantitative basis that these algal filaments grow while twisting around their own axis. The sense of spiral growth of the cell wall inSpirogyra-1 was always left-handed being coincident with the sense of chloroplast helix. InSpirogyra-2, the growth vector of the cell wall was likewise left-handed in most cases, but there occurred right-handed growth also. InZygnema both left-handed and right-handed senses of spiral growth were found in nearly equal frequencies. Besides the natural cell wall growth, the effects of longitudinal tension and turgor pressure on elongation and twisting of the filaments were briefly studied. It was shown that the cell wall of Zygnemataceae exhibited mechanical anisotropy in helical direction.     

Use of algal monocultures by larvae of Hydroptila waubesiana and Oxyethira pallida (Trichoptera: Hydroptilidae)

Larvae of the microcaddisflies Hydroptila waubesiana Betten and Oxyethira pallida (Banks) were given monocultures of eight genera of algae to test their suitability as food sources, and to investigate the degree of trophic specialization exhibited by these larvae. Only those larvae given green algae (Chlorophyta) survived and emerged as adults. Oxyethira pallida consumed Zygnema and Spirogyra, and exhibited the shortest larval-pupal period on Spirogyra (P < 0.05). Similarly, H. waubesiana consumed Zygnema and Cladophora, but exhibited the shortest larval-pupal period on Cladophora (P < 0.001). Diatoms and Hydrodictyon allowed H. waubesiana to attain partial growth only, while O. pallida did not consume either alga. Oscillatoria and Anabaena were not consumed by either species. The larvae of the two species tested appear to obtain adequate nourishment from only a narrow spectrum of algae, and one alga allowed significantly faster growth compared to the others tested in each case. This revised version was published online in July 2006 with corrections to the Cover Date.     

Wachstum und Stärkebildung einiger Konjugaten auf Kosten organisch gebundenen Kohlenstoffes

Zusammenfassung Von zehn untersuchten Stoffen (Glyzerin, Mannit, Dulzit, Glukose, Fruktose, Mannose, Galaktose, Saccharose, Maltose und Laktose) erwiesen sich nur Glukose, Maltose und Saccharose und nur beiMesotaenium caldariorum als geeignete Kohlenstoffquellen zur Unterhaltung von Wachstum und zur Bildung von Stärke. Sowohl Wachstum als auch Stärkebildung gehen in diesem Falle im Licht und im Dunkeln vor sich, wenn sich auch im einzelnen ein deutlicher Einfluß des Lichtes dabei beobachten läßt. Wachstum und Stärkebildung können nur in hypotonischen Lösungen stattfinden. Das Wachstumsoptimum liegt wesentlich niedriger als die plasmolytische Grenzkonzentration, das Optimum der Stärkebildung scheint mit der plasmolytischen Grenzlösung zusammen zu fallen.Cosmarium Botrytis, Zygnema sp., Zygnema peliosporum undSpirogyra varians können keinen der genannten Stoffe weder im Licht noch im Dunkeln, weder in hypo-noch hypertonischen Lösungen, weder zum Wachstum noch zur Stärkebildung ausnützen. Außer diesen in bakterienfreier Kultur befindlichen Algen wurde Freilandmaterial vonSpirogyra majuscula, Spirogyra Weberi undSpirogyra fallax (?) für einige Stärkebildungsversuche herangezogen. Auch sie verliefen negativ. Einige meiner Versuchsergebnisse legen die Vermutung nahe, daß die früheren Angaben durch eine unzutreffende Deutung von Versuchsergebnissen nach mangelhafter Entstärkung zustande gekommen sind.     

Species associations and metal contents of algae from rivers polluted by heavy metals

SUMMARY. The algal flora of the Rivers Hayle and Gannel. whieh drain copper and lead mining regions of Cornwall, are described and compared. Although fluctuations in the rarer members of the algal communities were observed, the dominant filamentous algae at most sites did not change over the year of study. Both the total algal abundance and the number of species were depressed at high metal sites. Associations of species that were evident in field samples were confirmed and correlated with water metal levels by principal component analysis. The close similarity between the flora of similar sites on the copper-polluted River Hayle and the lead-polluted River Gannei implies that the degree of metal pollution, rather than the polluting metal per se, determines the species present. All mine sites were characterized by a Microspora -community whereas a Zygnemales community of Spirogyra and Mougeotia species was typical of low metal pollution. Moderately polluted sites downsteam of the mines had an intermediate flora of Zygnemaies, Microsporales. Ulotrichales and gelatinous Volvocales and Tetrasporales species. No species could be said to invariably indicate metal pollution; the most abundant species at highly contaminated sites were also those with the widest distributions. Field samples of filamentous algae (mainly Spirogyra, Zygogonium, Mougeotia and Microspora species) contained metal concentrations several orders of magnitude greater than ambient levels. For copper and iron, both algal metal contents and concentration ratios (μg g-1 algae/μg ml-1 water) were positive functions of water metal levels, although algal iron appeared to plateau at water concentrations of about 1 mg ml-1. Algal lead concentrations, but not the concentration ratios, also were positively correlated with water lead levels. Thus, for these three metals, the algal metal contents were indicative of ambient conditions. In contrast, algal zinc concentrations were nearly constant and consequently the concentration ratios for this metal were inversely related to water levels. This result suggests that unlike the uptake of other metals, uptake of zinc by these aigae may be strictly regulated.     

Seasonal algal succession and cultural eutrophication in a north temperate lake

A year-round study of the algal composition of a previously uninvestigated north temperate, dimictic lake revealed the abundance of many pollution-tolerant forms, including such toxin-producing blue-greens as Anabaena and Aphanizomenon. A significant hypolimnetic oxygen deficit (0.4 ppm, 3.5% saturation) in late summer and the rate of oxygen depletion from 1 August to 19 September (0.13 mg/ml/day) further indicate eutrophic conditions. Of the three groups considered in this study (Chlorophyta, Bacillariophyta and Cyanophyta), the Cyanophyta were less diverse in terms of relative numbers of genera but produced the largest blooms. Chlorophyta had the greatest number of genera. Diatoms dominated in winter and spring, Chlorophyta in summer and fall and blue-greens in late summer, fall and winter. Spirogyra and Oscillatoria were the most ubiquitous members of the algal flora. Important perennials included Oscillatoria, Spirogyra, Closterium, Fragilaria, Meridion, Tabellaria and Cymbella. No unialgal blooms ever occurred. The accelerated rate of eutrophication in recent years is due primarily to excess nutrient loading resulting from input of raw domestic sewage. The completion of a sanitary sewer system is expected to alleviate the excessive nutrient loading and thereby slow the eutrophication process.     

PLOIDAL CHANGES IN CLONAL CULTURES OF SPIROGYRA COMMUNIS AND IMPLICATIONS FOR SPECIES DEFINITION

A clonal culture of Spirogyra filaments of initially uniform width produced filaments of three additional significantly different widths. Group I filaments of the original clone were 30.9 ± 0.7 μm wide (mean ± SD, N = 50). Group I filaments produced Group II filaments (22.0 ± 1.1 μm) through vegetative growth and sexual reproduction. Zygospores from homothallic Group I filaments produced germlings representative of Groups I and II; zygospores from homothallic Group II filaments produced germlings representative of Group II only. Germlings of Groups III (27.7 ± 1.0 μm) and IV (44.9 ± 0.8 μm) were produced in the cross of I × II. Viable zygospores from homothallic Group III filaments were obtained. Cells of Group IV filaments were initially binucleate and did not conjugate. Of the six intergroup crosses possible, four resulted in conjugation-tube formation only; two crosses yielded zygospores (I × II and III × IV). Germlings from the successful cross of Groups III and IV produced filaments of all four groups. Chromosome counts were: Group I (24), Group II (12), Group III (18), and Group IV (24, one nucleus). Relative nuclear fluorescence values of mithramycin-stained DNA were (mean ± SD, N ≥ 30): Group I (11.1 ± 1.4), Group II (5.7 ± 0.7), Group III (8.8 ± 1.3), and Group IV (10.0 ± 0.9, one nucleus). Cytologically, Group II appears to be a diploid (2x), Group I a tetraploid (4x), and Group III a triploid (3x). Systematically, Groups I, II, and III key out to Spirogyra singularis, S. communis, and S. fragilis, respectively, using Transeau's mongraph of the family Zygnemataceae. These species are interpreted to represent a species complex of S. communis (whose name has priority) with the ancestral haploid (x = 6) missing.     

Saponin-induced release of single cells from filaments and rhizoid differentiation in<Emphasis Type="Italic"> Spirogyra</Emphasis>

Some species of Spirogyra living in streams can anchor to the substratum by differentiating a rhizoid from a terminal cell of a filament. Rhizoid differentiation occurs in the light but not in the dark. When a filament of Spirogyra sp. competent for rhizoid differentiation was incubated in a medium containing 0.1% saponin, terminal cells were released one by one, forming single cells. Single cells effectively differentiated to be rhizoids when saponin in the incubation medium was removed. The single-cell system developed in the present study seems suitable for analysis of gene expression during rhizoid differentiation of Spirogyra.     

SYSTEMATICS OF THE GENUS ZYGNEMA (ZYGNEMATOPHYCEAE,CHAROPHYTA) FROM CALIFORNIAN WATERSHEDS1

Natural populations of Zygnema were collected from 80 stream sites across California, and eight species were identified and characterized morphologically. Generic and infrageneric concepts of Zygnema and Zygogonium were tested with cox3 and rbcL gene sequence analysis. Strains of Zygnema were positioned in a single monophyletic clade sister to Zygogonium tunetanum Gauth.‐Lièvre. In both the rbcL and cox3 phylogenies, strains of Zygnema formed two major clades. The first clade contained species that have zygospores with a blue‐colored mesospore or akinetes with a colorless mesospore. The second clade contained species that have a yellow or brown mesospore. The existing taxonomic concepts for Zygnema classification are not consistent with our molecular phylogeny and do not correspond to natural groups. We propose that mesospore color may be useful in the infrageneric classification of Zygnema. Newly described Zygnema aplanosporum sp. nov. and Zygnema californicum sp. nov. have zygospores with a blue mesospore formed in the conjugation tube and separated by a cellulosic sporangial wall. Z. aplanosporum also possessed a combination of vegetative and reproductive features characteristic of Zygogonium, such as presence of short branches, rhizoidal outgrowths, thickened vegetative cell walls, purple‐colored cell content, small compressed‐globular chloroplasts as well as predominant asexual reproduction. Z. aplanosporum and Z. californicum were deeply embedded in a larger clade of Zygnema both in rbcL and cox3 analyses. Based on our observations, there are no features or combination of features that separate Zygnema and Zygogonium. Therefore, we conclude that Zygogonium is probably a synonym of Zygnema.     

Hidden genetic diversity in the green alga Spirogyra (Zygnematophyceae, Streptophyta)

ABSTRACT: BACKGROUND: The unbranched filamentous green alga Spirogyra (Streptophyta, Zygnemataceae) is easily recognizable based on its vegetative morphology, which shows one to several spiral chloroplasts. This simple structure falsely points to a low genetic diversity: Spirogyra is commonly excluded from phylogenetic analyses because the genus is known as a long-branch taxon caused by a high evolutionary rate. RESULTS: We focused on this genetic diversity and sequenced 130 Spirogyra small subunit nuclear ribosomal DNA (SSU rDNA) strands of different origin. The resulting SSU rDNA sequences were used for phylogenetic analyses using complex evolutionary models (posterior probability, maximum likelihood, neighbor joining, and maximum parsimony methods). The sequences were between 1672 and 1779 nucleotides long. Sequence comparisons revealed 53 individual clones, but our results still support monophyly of the genus. Our data set did not contain a single slow-evolving taxon that would have been placed on a shorter branch compared to the remaining sequences. Out of 130 accessions analyzed, 72 showed a.