PHYLOGENY OF THE CONJUGATING GREEN ALGAE BASED ON CHLOROPLAST AND MITOCHONDRIAL NUCLEOTIDE SEQUENCE DATA1

The conjugating green algae represent a lineage of charophyte green algae known for their structural diversity and unusual mode of sexual reproduction, conjugation. These algae are ubiquitous in freshwater environments, where they are often important primary producers, but few studies have investigated evolutionary relationships in a molecular systematic context. A 109‐taxon data set consisting of three gene fragments (two from the chloroplast and one from the mitochondrial genome) was used to estimate the phylogeny of the genera of the conjugating green algae. Maximum likelihood (ML), maximum parsimony (MP), and Bayesian inference (BI) were used to estimate relationships from the 4,047 alignable nucleotides. This study confirmed the polyphyly of the Zygnemataceae and Mesotaeniaceae with respect to one another. The Peniaceae were determined to be paraphyletic, and two genera traditionally classified among the Zygnematales appear to belong to the lineage that gave rise to the Desmidiales. Six genera, Euastrum, Cosmarium, Cylindrocystis, Mesotaenium, Spondylosium, and Staurodesmus, were polyphyletic in this analysis. These findings have important implications for the evolution of structural characteristics in the group and will require some taxonomic changes. More work will be required to delineate lineages of Zygnematales in particular and to identify structural synapomorphies for some of the newly identified clades.     

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.     

PHYLOGENY OF THE CONJUGATING GREEN ALGAE (ZYGNEMOPHYCEAE) BASED ON rbc L SEQUENCES

Sequences of the gene encoding the large subunit of RUBISCO (rbcL) for 30 genera in the six currently recognized families of conjugating green algae (Desmidiaceae, Gonatozygaceae, Mesotaeniaceae, Peniaceae, and Zygnemataceae) were analyzed using maximum parsimony and maximum likelihood; bootstrap replications were performed as a measure of support for clades. Other Charophyceae sensu Mattox and Stewart and representative land plants were used as outgroups. All analyses supported the monophyly of the conjugating green algae. The Desmidiales, or placoderm desmids, constitute a monophyletic group, with moderate to strong support for the four component families of this assemblage (Closteriaceae, Desmidiaceae, Gonatozygaceae, and Peniaceae). The analyses showed that the two families of Zygnematales (Mesotaeniaceae, Zygnemataceae), which have plesiomorphic, unornamented and unsegmented cell walls, are not monophyletic. However, combined taxa of these two traditional families may constitute a monophyletic group. Partitioning the data by codon position revealed no significant differences across all positions or between partitions of positions one and two versus position three. The trees resulting from parsimony analyses using first plus second positions versus third position differed only in topology of branches with poor bootstrap support. The tree derived from third positions only was more resolved than the tree derived from first and second positions. The rbcL‐based phylogeny is largely congruent with published analyses of small subunit rDNA sequences for the Zygnematales. The molecular data do not support hypotheses of monophyly for groups of extant unicellular and filamentous or colonial desmid genera exhibiting a common cell shape. A trend is evident from simple omniradiate cell shapes to taxa with lobed cell and plastid shapes, which supports the hypothesis that chloroplast shape evolved generally from simple to complex. The data imply that multicellular placoderm desmids are monophyletic. Several anomalous placements of genera were found, including the saccoderm desmid Roya in the Gonatozygaceae and the zygnematacean Entransia in the Coleochaetales. The former is strongly supported, although the latter is not, and Entransia's phylogenetic position warrants further study.     

PHYLOGENY OF THE DASYCLADALES (CHLOROPHYTA,ULVOPHYCEAE) BASED ON ANALYSES OF RUBISCO LARGE SUBUNIT (rbcL) GENE SEQUENCES1

The phylogeny of the green algal Order Dasycladales was inferred by maximum parsimony and Bayesian analyses of chloroplast‐encoded rbcL sequence data. Bayesian analysis suggested that the tribe Acetabularieae is monophyletic but that some genera within the tribe, such as Acetabularia Lamouroux and Polyphysa Lamouroux, are not. Bayesian analysis placed Halicoryne Harvey as the sister group of the Acetabularieae, a result consistent with limited fossil evidence and monophyly of the family Acetabulariaceae but was not supported by significant posterior probability. Bayesian analysis further suggested that the family Dasycladaceae is a paraphyletic assemblage at the base of the Dasycladales radiation, casting doubt on the current family‐level classification. The genus Cymopolia Lamouroux was inferred to be the basal‐most dasycladalean genus, which is also consistent with limited fossil evidence. Unweighted parsimony analyses provided similar results but primarily differed by the sister relationship between Halicoryne Lamouroux and Bornetella Munier‐Chalmas, thus supporting the monophyly of neither the families Acetabulariaceae nor Dasycladaceae. This result, however, was supported by low bootstrap values. Low transition‐to‐transversion ratios, potential loss of phylogenetic signal in third codon positions, and the 550 million year old Dasycladalean lineage suggest that dasyclad rbcL sequences may be saturated due to deep time divergences. Such factors may have contributed to inaccurate reconstruction of phylogeny, particularly with respect to potential inconsistency of parsimony analyses. Regardless, strongly negative g₁ values were obtained in analyses including all codon positions, indicating the presence of considerable phylogenetic signal in dasyclad rbcL sequence data. Morphological features relevant to the separation of taxa within the Dasycladales and the possible effects of extinction on phylogeny reconstruction are discussed relative to the inferred phylogenies.     

PHYSIOLOGICAL RESPONSES TO TEMPERATURE AND IRRADIANCE IN SPIROGYRA (ZYGNEMATALES, CHAROPHYCEAE)1

Spirogyra Link (1820) is an anabranched filamentous green alga that forms free-floating mats in shallow waters. It occurs widely in static waters such as ponds and ditches, sheltered littoral areas of lakes, and stow-flowing streams. Field observations of its seasonal distribution suggest that the 70-μm-wide filament form of Spirogyra should have a cool temperature and high irradiance optimum for net photosynthesis. Measurements of net photosynthesis and respiration were marie at 58 combinations of tight and temperature in a controlled environment facility. Optimum conditions were 25°C and 1500 μmol photons m⁻² s⁻¹, at which net photosynthesis averaged 75.7 mg O₂ gdm⁻¹ h⁻¹. Net photosynthesis was positive at temperatures from 5° to 35°C at most irradiances except at combinations of extremely low irradiances and high temperatures (7 and 23 μmol photons m⁻² s⁻¹ at 30°C and 7, 23, and 35 μmol photons m⁻² s⁻¹ at 35°C). Respiration rates increased with both temperature and prior irradiance. Light-enhanced respiration rates were significantly greater than dark respiration rates following irradiances of 750 μmol photons m⁻² s⁻¹ or greater. Polynomials were fitted to the data to generate response surfaces; such response surfaces can be used to represent net photosynthesis and respiration in ecological models. The data indicate that the alga can tolerate the cool water and high irradiances characteristic of early spring but cannot maintain positive net photosynthesis under conditions of high temperature and low light (e.g. when exposed to self-shading ).     

EFFECTS OF TEMPERATURE AND IRRADIANCE ON THE SEASONAL VARIATION OF A SPIROGYRA (CHLOROPHYTA) POPULATION IN A MIDWESTERN LAKE (U.S.A.)

Although Spirogyra Link (1820) is a common mat‐forming filamentous alga in fresh waters, little is known of its ecology. A 2‐year field study in Surrey Lake, Indiana, showed that it grew primarily in the spring of each year. The population consisted of four morphologically distinct filamentous forms, each exhibiting its own seasonal distribution. A 45‐μm‐wide filament was present from February to late April or early May, a 70‐μm‐wide form was present from late April to mid‐June, a 100‐μm‐wide form was present from February to mid‐June, and a 130‐μm‐wide form appeared only in February of 1 of 2 study years. The 70‐ and 100‐μm‐wide forms contributed to the peak amount of biomass observed in late May and early June. Multiple regression analysis indicated that the presence of the 45‐, 70‐, and 100‐μm‐wide forms was negatively correlated with temperature. Presence of the 130‐μm‐wide form was negatively correlated with irradiance. Isolates of these filament forms were exposed to temperature (15, 25, and 35° C)/irradiance (0, 60, 200, 400, 900, and 1500 μmol·m^?2·s^?1) combinations in the laboratory. Growth rates of the 45‐μm‐wide form were negative at all irradiances at 35° C, suggesting that this form is susceptible to high water temperatures. However, growth rates of the other forms did not vary at the different temperatures or at irradiances of 60 μmol·m^?2·s^?1 or above. Net photosynthesis was negative at 35° C and 1500 μmol·m^?2·s^?1 for the 100‐ and 130‐μm‐wide forms but positive for the 70‐μm‐wide form. All forms lost mat cohesiveness in the dark, and the 100‐ and 130‐μm‐wide forms lost mat cohesiveness under high irradiances and temperature. Thus, the morphological forms differed in their responses to irradiance and temperature. We hypothesize that the rapid disappearance of Spirogyra populations in the field is due to loss of mat cohesiveness under conditions of reduced net photosynthesis, for example, at no to low light for all forms or at high light and high temperatures for the 100‐ and 130‐μm‐wide forms. Low light conditions can occur in the interior of mats as they grow and thicken or under shade produced by other algae.     

A MOLECULAR PHYLOGENY OF THE HETEROKONT ALGAE BASED ON ANALYSES OF CHLOROPLAST-ENCODED rbcL SEQUENCE DATA1

Nearly complete ribulose-1,5-bisphosphate carboxylase/ oxygenase (rbcL)sequences from 27 taxa of heterokont algae were determined and combined with rbcL sequences obtained from GenBank for four other heterokont algae and three red algae. The phylogeny of the morphologically diverse haterokont algae was inferred from an unambiguously aligned data matrix using the red algae as the root, Significantly higher levels of mutational saturation in third codon positions were found when plotting the pair-wise substitutions with and without corrections for multiple substitutions at the same site for first and second codon positions only and for third positions only. In light of this observation, third codon positions were excluded from phylogenetic analyses. Both weighted-parsimony and maximum-likelihood analyses supported with high bootstrap values the monophyly of the nine currently recognized classes of heterokont algae. The Eustigmatophyceae were the most basal group, and the Dictyochophyceae branched off as the second most basal group. The branching pattern for the other classes was well supported in terms of bootstrap values in the weightedparsimony analysis but was weakly supported in the maximum-likelihood analysis (<50%). In the parsimony analysis, the diatoms formed a sister group to the branch containing the Chrysophyceae and Synurophyceae. This clade, charactetized by siliceous structures (frustules, cysts, scales), was the sister group to the Pelagophyceae/Sarcinochrysidales and Phaeo-/Xantho-/ Raphidophyceae clades. In the latter clade, the raphido-phytes were sister to the Phaeophyceae and Xanthophyceae. A relative rate test revealed that the rbcL gene in the Chrysophyceae and Synurophyceae has experienced a significantly different rate of substitutions compared to other classes of heterokont algae. The branch lengths in the maximum-likelihood reconstruction suggest that these two classes have evolved at an accelerated rate. Six major carotenoids were analyzed cladistically to study the usefulness of carotenoid pigmentation as a class-level character in the heterokont algae. In addition, each carotenoid was mapped onto both the rbcL tree and a consensus tree derived from nuclear-encoded small-subunit ribosomal DNA (SSU rDNA) sequences. Carotenoid pigmentation does not provide unambiguous phylogenetic information, whether analyzed cladistically by itself or when mapped onto phylogenetic trees based upon molecular sequence data.     

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 .     

MOLECULAR PHYLOGENY OF STAURASTRUM MEYEN EX RALFS AND RELATED GENERA (ZYGNEMATOPHYCEAE,STREPTOPHYTA) BASED ON CODING AND NONCODING RDNA SEQUENCE COMPARISONS1

Nuclear‐encoded small subunit rDNA, 1506 group I intron, and internal transcribed spacer sequences were obtained from 39 strains representing five core desmid genera, Staurastrum, Staurodesmus Teil., Cosmarium Corda ex Ralfs, Xanthidium Ehr. ex Ralfs, and Euastrum Ehr. ex Ralfs (Desmidiaceae, Zygnematophyceae), and used individually and concatenated to assess phylogenetic relationships between putatively allied members of the family. To identify positional homology between divergent noncoding sequences, secondary structure models were generated and their reliability assessed by screening the alignment for compensating base changes. The phylogeny based on coding and noncoding sequence comparisons confidently resolved a monophyletic core of the genus Staurastrum but also revealed the artificial nature of the traditional genus. Twenty distinct species representing a wide range of morphotypes of Staurastrum formed a strongly supported generic clade that was further split into three well‐resolved lineages. The phylogenetic relationships revealed within Staurastrum were in conflict with all previous formal or informal classifications of the genus. The genera Staurodesmus and Cosmarium were shown to be highly polyphyletic, and some morphologically similar taxa displayed high sequence divergence that exceeded generic boundaries. Apparently, the taxonomic significance of some morphological characters in Staurastrum and other desmid genera has been greatly overestimated.     

黑龙江省双星藻科新种

三江水绵图1 Spirogyra sanjingensis sp. nov. Celluis vegetativis 104-133μlatis, 142—295μlongis; dissepimentis planis; chromatophoris 9-10(11), anfractibus 1-1.5; conjugations scalari, tubo ex utroque cellula mascula et feminca emisso; cellulis fructiferis cylindricis, levi-     

PHYLOGENY OF THE BASAL LINEAGES OF STREPTOPHYTA BASED ON RBCL AND ATPB GENE SEQUENCE DATA

The streptophytes comprise the Charophyceae sensu Mattox and Stewart (a morphologically diverse group of fresh‐water green algae) and the embryophytes (land plants). Several charophycean groups are currently recognized. These include the Charales, Coleochaetales, Chlorokybales, Klebsormidiales and Zygnemophyceae (Desmidiales and Zygnematales). Recently, SSU rRNA gene sequence data allied Mesostigma viride (Prasinophyceae) with the Streptophyta. Complete chloroplast sequence data, however, placed Mesostigma sister to all green algae, not with the streptophytes. Several morphological, ultrastructural and biochemical features unite these lineages into a monophyletic group including embryophytes, but evolutionary relationships among the basal streptophytes remain ambiguous. To date, numerous studies using SSU rRNA gene sequences have yielded differing phylogenies with varying degrees of support dependent upon taxon sampling and choice of phylogenetic method. Like SSU data, chloroplast DNA sequence data have been used to examine relationships within the Charales, Coleochaetales, Zygnemophyceae and embryophytes. Representatives of all basal streptophyte lineages have not been examined using chloroplast data in a single analysis. Phylogenetic analyses were performed using DNA sequences of rbcL (the genes encoding the large subunit of rubisco) and atpB (the beta‐subunit of ATPase) to examine relationships of basal streptophyte lineages. Preliminary analyses placed the branch leading to Mesostigma as the basal lineage in the Streptophyta with Chlorokybus, the sole representative of the Chlorokybales, branching next. Klebsormidiales and the enigmatic genus Entransia were sister taxa. Sister to these, the Charales, Coleochaetales, embryophytes and Zygnemophyceae formed a monophyletic group with Charales and Coleochaetales sister to each other and this clade sister to the embryophytes.     

PHYLOGENY OF CHLAMYDOMONAS (CHLOROPHYTA) BASED ON CLADISTIC ANALYSIS OF NUCLEAR 18S rRNA SEQUENCE DATA1

The genus Chlamydomonas Ehrenberg may contain as many as 450 described species. Morphological, physiological and molecular data show that variation among some Chlamydomonas species can he great, leading to speculation that multiple, generic-level lineages exist within this genus. The most recent systematic studies of Chlamydomonas have led to proposals of nine distinct morphological and 15 distinct sporangial autolysin groups. Partial sequences from the nuclear small subunit rRNAs from 14 Chlamydomonas species representing 12 autolysin and four morphological groups, and from three flagellates thought to he related to Chlamydomonas were determined in a phylogenetic study of relationships among these algae. Sequence comparisons among some Chlamydomonas species revealed differences comparable to the sequence divergence between soybeans and cycads. Cladistic analysis of the sequence data suggests that multiple lineages exist among species of Chlamydomonas. Some of these lineages represent alliances of both Chlamydomonas and non-Chlamydomonas taxa; thus, the current taxonomy does not reflect natural, or monophyletic, groups. Collectively, these lineages may represent distinct families or even orders.     

SYSTEMATICS OF THE GRACILARIACEAE (GRACILARIALES,RHODOPHYTA): A CRITICAL ASSESSMENT BASED ON RBCL SEQUENCE ANALYSES1

Generic concepts in the economically important agarophyte red algal family Gracilariaceae were evaluated based on maximum parsimony, Bayesian likelihood, and minimum evolution analyses of the chloroplast‐encoded rbc L gene from 67 specimens worldwide. The results confirm the monophyly of the family and identify three large clades, one of which corresponds to the ancestral antiboreal genera Curdiea and Melanthalia, one to Gracilariopsis, and one to Gracilaria sensu lato, which contains nine distinct independent evolutionary lineages, including Hydropuntia. The species currently attributed to Hydropuntia comprise a single well‐supported clade composed of two distinct lineages. The two most basal clades within Gracilaria sensu lato deserve generic rank: a new genus centered around G. chilensis Bird, McLachlan et Oliveira and G. aff. tenuistipitata Chang et Xia and a resurrected Hydropuntia encompassing primarily Indo‐Pacific (G. urvillei [Montagne] Abbott, G. edulis [S. Gmelin] P. Silva, G. eucheumatoides Harvey, G. preissiana [Sonder] Womersley, and G. rangiferina [Kützing] Piccone) and western Atlantic species (G. cornea J. Agardh, G. crassissima P. et H. Crouan in Mazé et Schramm, G. usneoides [C. Agardh] J. Agardh, G. caudata J. Agardh, and G. secunda P. et H. Crouan in Mazé et Schramm). Cystocarpic features within the Gracilaria sensu lato clades appear to be more phylogenetically informative than male characters. The textorii‐type spermatangial configuration is represented in two distinct clusters of Gracilaria. The rbc L genetic divergence among the Gracilariaceae genera ranged between 8.46% and 16.41%, providing at least 2.5 times more genetic variation than does the 18S nuclear rDNA. rbc L also resolves intrageneric relationships, especially within Gracilaria sensu lato. The current number of gracilariacean species is underestimated in the western Atlantic because of convergence in habit and apparent homoplasy in vegetative and reproductive anatomy.     

PHYLOGENY AND SYSTEMATICS OF THE GENUS MONOMORPHINA (EUGLENACEAE) BASED ON MORPHOLOGICAL AND MOLECULAR DATA1

Morphological studies of 16 strains belonging to the genus Monomorphina revealed a single, parietal, orbicular chloroplast in their cells. The chloroplast has a tendency to be perforated and disintegrates in aging populations and thus may appear to be many chloroplasts under the light microscope. A single chloroplast in the cells of Cryptoglena skujae is also parietally located and highly perforated. It never forms a globular and closed structure, but is open from the side of the furrow, resembling the letter C. We have verified the Monomorphina pyrum group (M. pyrum–like) on the basis of phylogenetic analysis of SSU rDNA and morphological data. The strain CCAC 0093 (misidentified as M. reeuwykiana) diverges first on the SSU rDNA phylogenetic tree. The rest of the M. pyrum–like strains form a tight cluster, subdivided into several smaller ones. Because morphological differences between the M. pyrum–like strains (including the strain CCAC 0093) do not conform to the tree topology, we suggest that they all (except the strain CCAC 0093) belong to M. pyrum. We designate a new species, M. pseudopyrum, for the strain CCAC 0093, solely on the basis of molecular characters. We also suggest that M. reeuwykiana and similar species should stay in Phacus and Lepocinclis unless detailed molecular and morphological studies show otherwise. Emended diagnoses of the genera Monomorphina and Cryptoglena and the species M. aenigmatica are also proposed, as well as the delimitation of an epitype for M. pyrum, the type species for the genus Monomorphina.     

PHYLOGENY OF THE DUMONTIACEAE (GIGARTINALES, RHODOPHYTA) AND ASSOCIATED FAMILIES BASED ON SSU rDNA AND INTERNAL TRANSCRIBED SPACER SEQUENCE DATA

Small subunit (SSU) rDNA was sequenced for 25 species in 19 genera of the Gigartinales (Rhodophyta). As well, the internal transcribed spacer (ITS) region was sequenced, and a data matrix of 36 morphological characters was constructed for 16 species of Dumontiaceae. Phylogenetic trees were calculated from a multiple alignment of the SSU sequence data to infer relationships between species of Dumontiaceae and other gigartinalean taxa. The SSU analysis produced a polyphyletic Dumontiaceae. Notably, Acrosymphyton failed to associate with the included Gigartinales, let alone the Dumontiaceae, supporting an earlier proposal to remove it to a new family. The analyses were equivocal about the phylogenetic affinities of Dudresnaya , which clustered with the Kallymeniaceae, and the affinities of the Indo-West Pacific Gibsmithia , Kraftia , and Dasyphloea , the last-mentioned clustering with the Antarctic Gainiaceae, and these four taxa with Portieria (Rhizophyllidaceae). Further investigations are necessary to resolve relationships among these taxa. Rhodopeltis , a genus recently moved to the Dumontiaceae from the Polyideaceae, showed a weak association with the remaining northern Dumontiaceae. The final group consisted of cold-temperate Northern Hemisphere species. Phylogenetic analyses using a combination of SSU, ITS, and morphological data within this clade produced two strongly supported clades, a Dilsea / Neodilsea clade and a Cryptosiphonia / Dumontia clade. Dilsea is derived from a paraphyletic Neodilsea and may itself be polyphyletic. Atlantic and Pacific isolates of Dumontia contorta clearly showed sufficient divergence to warrant recognition as distinct species, and Dumontia alaskana , sp. nov. is proposed for the Pacific species.     

MOLECULAR PHYLOGENY OF THE HAPTOPHYTA BASED ON THE rbcL GENE AND SEQUENCE VARIATION IN THE SPACER REGION OF THE RUBISCO OPERON

A phylogeny of 21 haptophyte algae was inferred by maximum parsimony, neighbor-joining, and maximum likelihood analyses of sequences of the plastid-encoded gene, rbcL. Sequence variation in the spacer region of the RUBISCO operon was also investigated. In all the rbcL trees constructed, the haptophytes form two distinct clades: one includes the Pavlovales and the other includes the Prymnesiales, Coccosphaerales, and Isochrysidales (all sensu Parke and Green 1976 . This relationship coincides with the recent taxonomic treatment splitting the division into two subclasses, the Prymnesidae and Pavlovidae ( Cavalier-Smith 1989 ) or the Prymnesiophycidae and the Pavlovophycidae using botanical suffixes ( Jordan and Green 1994 ), or into two classes, the Patelliferea and the Pavlovea ( Cavalier-Smith 1993 ). In the Prymnesiophycidae, all the coccolithophorids examined are placed in a single clade, which suggests a single origin of the coccolithophorids and the ability of coccolith formation in the haptophytes. The genus Chrysochromulina is polyphyletic. Species of Chrysochromulina with a very long haptonema and a compressed cell body (typical of species including the type C. parva Lackey) form a clade, including Imantonia, that is often classified in the Isochrysidales in the neighbor-joining tree, whereas some species possessing a nontypical cell body and cell covering form a clade with Prymnesium and Platychrysis in all trees. It is suggested that loss of the haptonema in Imantonia and the reduction in Prymnesium and Platychrysis occurred secondarily and independently in two different lineages. Within the coccolithophorids, four clades are recognized: Pleurochrysis, Calyptrosphaera-Cruciplacolithus-Calcidiscus-Umbilicosphaera, Helicosphaera, and Emiliania-Gephyrocapsa. A non-coccolith-bearing haptophyte, Isochrysis, is an ingroup of the Emiliania-Gephyrocapsa clade, suggesting its secondary loss of the ability to form a coccolith. Sequence comparison of the spacer region of RUBISCO operon supports most results obtained in the analysis of rbcL sequences. Monophyly of the Prymnesiales sensu Parke and Green is still unclear because of low (<50%) bootstrap support for this group.     

A REVISED CLASSIFICATION OF THE DICTYOTEAE (DICTYOTALES,PHAEOPHYCEAE) BASED ON rbcL AND 26S RIBOSOMAL DNA SEQUENCE ANALYSES1

Dictyota is a genus of tropical to warm temperate brown algae characterized by parenchymatous, flattened thalli that grow from a single, transversely oriented apical cell. Dictyota is currently distinguished from allied genera of the tribe Dictyoteae (Dilophus, Glossophora, Glossophorella, and Pachydictyon) by the structure of the cortical and medullary layers, as well as the relative abundance of surface proliferations. Even though the traditional classification of the Dictyoteae has repeatedly been criticized in the past, the absence of sound molecular data has so far discouraged any new taxonomic proposals apart from a merger of Dilophus with Dictyota, which has been accepted by only part of the phycological community. Phylogenetic analysis of rbcL gene, partial 26S rDNA sequence, and combined data sets, including four of five generitypes, demonstrates that the traditional classification does not accurately reflect the evolutionary history of the group. None of the genera are resolved as a monophyletic clade. Hence, a merger of Glossophora, Glossophorella, and Pachydictyon in Dictyota is proposed. Two new genera, Canistrocarpus (incorporating D. cervicornis, D. crispata, and D. magneana) and Rugulopteryx (accommodating D. radicans, Dil. suhrii, and Dil. marginata), are proposed. Both genera are supported by molecular indications and a combination of reproductive and vegetative characters. The position of Dil. fastigiatus as a clade sister to Dictyota s.l. and the absence of Dil. gunnianus, the generitype of Dilophus, from the analyses, prevented us from making a more definite statement on the status of the latter genus.     

A MOVING MAT: PHOTOTAXIS IN THE FILAMENTOUS GREEN ALGAE SPIROGYRA (CHLOROPHYTA,ZYGNEMATACEAE)1

A blue light– (peak at 470 nm) induced photomovement was observed in the filamentous eukaryotic algae, Spirogyra spp. When Spirogyra filaments were scattered in a water chamber under a unilateral light source, they rapidly aligned toward the light source in 1 h and bound with neighboring filaments to form thicker parallel bundles of filaments. The filaments in the anterior of the bundles curved toward the light first and then those in the posterior began to roll up toward the light, forming an open‐hoop shape. The bundle of filaments then moved toward the light source by repeated rolling and stretching of filaments. When the moving bundle met other filaments, they joined and formed a bigger mat. The coordination of filaments was essential for the photomovement. The average speed of movement ranged between 7.8 and 13.2 μm·s^?1. The movement was induced in irradiance level from 1 to 50 μmol photons·m^?2·s^?1. The filaments of Spirogyra showed random bending and stretching movement under red or far‐red light, but the bundles did not move toward the light source. There was no distinct diurnal rhythm in the photomovement of Spirogyra spp.     

串珠藻目植物的系统发育-基于rbcL序列的证据

世界范围内报道的全部串珠藻目种类均生活于淡水中,而在淡水红藻中,70%约有130种属于串珠藻目。研究以目前获取的来自世界各大洲串珠藻目植物43种的rbcL基因序列,结合其形态和生物地理特征,构建了该目的系统发育关系,以期探讨整个串珠藻目植物的系统发育关系及发生途径,进而为研究该目以至淡水红藻的起源提供基本资料。运用PAUP*4.0b10和MrBayes 3.0b4等软件对43种串珠藻目植物的叶绿体DNA rbcL基因序列进行系统发育分析,探讨了其主要分类群的系统演化关系。用最大简约法、邻接法和贝叶斯分析方法构建的系统树基本一致,结果显示:(1)基于分子数据分析结果显示,红索藻目植物均独立于串珠藻目植物,构成一个单独的分支,支持红索藻目的建立。(2)鱼子菜科属于串珠藻目植物中较为进化的类群。(3)串珠藻属扭曲组与杂生组的差异度较小,结合其形态特点,倾向于将杂生组并入扭曲组。(4)串珠藻科属于串珠藻目中最大的科,包括较多的种类,其系统关系也较为复杂。因此,串珠藻科系统发育关系的明确有待于进一步结合更多的分子数据和形态学特征加以分析研究。