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 .     

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.     

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.     

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.     

Phylogeny of extant genera in the family Characeae (Charales,Charophyceae) based on rbcL,sequences and morphology

Extant genera of Characeae have been assigned to two tribes: Chareae (Chara, Lamprothamnium, Nitellopsis, and Lychnothamnus) and Nitelleae (Nitella and Tolypella), based on morphology of the thallus and reproductive structures. Character analysis of fossil and extant oogonia suggest that Tolypella is polyphyletic, the genus comprising two sections, one in each of the two tribes. Eleven morphological characters and sequence data for the Rubisco large subunit (rbcL) were used to reconstruct the phylogeny of genera, including the two sections of Tolypella. Parsimony analysis of the rbcL data, with all positions and changes weighted equally, strongly supports the monophyly of the Characeae. The two Tolypella sections form a robust monophyletic group basal to the family. Transversion weighting yielded the same tree but with a paraphyletic Tolypella. The rbcL data strongly support monophyly of tribe Chareae but tribe Nitelleae is paraphyletic. Parsimony analysis of morphological data produced one unrooted tree consistent with monophyly of the two tribes; on this tree the Tolypella sections were paraphyletic. Combining morphological with rbcL data did not change the results derived from rbcL sequences alone. The rbcL data support the monophyly of the Characeae and Coleochaete, which together form a monophyletic sister group to embryophytes.     

基于16S rRNA基因序列对织纹螺属的分子系统学分析

研究利用线粒体16S rRNA基因序列片段,对织纹螺属6亚属11种动物进行分子系统学分析。结果显示,种内遗传距离(0～0.007)与种间遗传距离(0.019～0.088)无重叠,这表明线粒体16S rRNA基因能较好地反映织纹螺种间的亲缘关系; 而亚属内遗传距离(0.018～0.031)与亚属间遗传距离(0.028～0.083)存在重叠,表明线粒体16S rRNA基因不能对一些贝壳形态相似的亚属进行区分。同时,确定了疑难种灰白织纹螺Nassarius(Zeuxis)canaliculatum的分类地位,并与相似种西格织纹螺N.(Z.)siquijorensis进行了对比; 确认了秀丽织纹螺Nassarius(Hima)festivus应属于Hima亚属;建议保留单型亚属Varicinassa的有效性。     

MONOPHYLY OF THE GENUS CLOSTERIUM AND THE ORDER DESMIDIALES (CHAROPHYCEAE, CHLOROPHYTA) INFERRED FROM NUCLEAR SMALL SUBUNIT rDNA DATA

We newly sequenced the nuclear-encoded small subunit (SSU) rDNA coding region for 21 taxa of the genus Closterium. The new sequences were integrated into an alignment with 13 known sequences of conjugating green algae representing six traditional families (i.e. Zygnemataceae, Mesotaeniaceae, Gonatozygaceae, Peniaceae, Closteriaceae, and Desmidiaceae) and five known charophycean sequences as outgroups. Both maximum likelihood and maximum parsimony analyses supported with high bootstrap values one large clade containing all placoderm desmids (Desmidiales). All the Closterium taxa formed one clade with 100% bootstrap support, indicating their monophyly, but not paraphyly, as suggested earlier. As to the taxa within the genus Closterium , we found two clades of morphologically closely related taxa in both maximum likelihood and maximum parsimony trees. They corresponded to the C. calosporum species complex and the C. moniliferum-ehrenbergii species complex. It is of particular interest that the homothallic entity of C. moniliferum v. moniliferum was distinguished from and ancestral to all other entities of the C. moniliferum-ehrenbergii species complex. Superimposing all 50 charophycean sequences on the higher order SSU rRNA structure model of Closterium , we investigated degrees of nucleotide conservation at a given position in the nucleotide sequence. A characteristic "signature" structure to the genus Closterium was found as an additional helix at the tip of V1 region. In addition, eight base deletions at the tip of helix 10 were found to be characteristic of the C. calosporum species complex, C. gracile , C. incurvum , C. pleurodermatum , and C. pusillum v. maius. These taxa formed one clade with an 82% bootstrap value in maximum parsimony analysis.     

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.     

A PHYLOGENETIC STUDY OF THE CORALLINALES (RHODOPHYTA) BASED ON NUCLEAR SMALL-SUBUNIT rRNA GENE SEQUENCES

Conflicting classifications for the Corallinales were tested by analyzing partial sequences for the nuclear small-subunit ribosomal RNA (SSU) gene of 35 species of coralline algae. Parsimony and likelihood analyses of these data yielded congruent hypotheses that are inconsistent with classifications for the group that include as many as eight subfamilies. Four major clades are resolved within the order, including the early-diverging Sporolithaceae as well as the Melobesioideae and Corallinoideae. The fourth clade, which is supported robustly, includes both nongeniculate and geniculate species classified in the subfamilies Mastophoroideae, Metagoniolithoideae, Lithophylloideae, and Amphiroideae. Molecular and morphological data support the proposal that the latter two subfamilies are sister taxa. Although relationships among some genera are not resolved clearly, the order of branching of taxa among and within the four principal lineages is concordant with paleontological evidence for the group. Relationships inferred among genera within each of the clades is discussed. Seven morphological characters delimiting higher taxonomic groups within the order were combined with the sequence data, analyzed, and optimized onto the resulting tree(s). Except for the presence or absence of genicula, all other characters were found to be phylogenetically informative. Genicula are nonhomologous structures that evolved independently in the Amphiroideae, Corallinoideae, and Metagoniolithoideae. The phenetic practice of separating coralline algae into two categories solely on the basis of the presence or absence of genicula does not accurately reflect the evolutionary history of the group.     

PHYLOGENY OF EUGLENOPHYCEAE BASED ON SMALL SUBUNIT rDNA SEQUENCES: TAXONOMIC IMPLICATIONS1

Small subunit rDNA sequences of 42 taxa belonging to 10 genera were used to infer phylogenetic relationships among euglenoids. Members of the phototrophic genera Euglena, Phacus, Lepocinclis, Colacium, Trachelomonas, and Strombomonas plus the osmotrophs Astasia longa, Khawkinea quartana, and Hyalophacus ocellatus were included. Six major clades were found in most trees using multiple methods. The utility of Bayesian analyses in resolving these clades is demonstrated. The genus Phacus was polyphyletic with taxa sorting into two main clades. The two clades correlated with overall morphology and corresponded in large part to the previously defined sections, Pleur‐ aspis Pochmann and Proterophacus Pochmann. Euglena was also polyphyletic and split into two clades. In Bayesian analyses species with less plastic pellicles and small disk‐like chloroplasts diverged at the base of the tree. They grouped into a single clade which included the two Lepocinclis spp., which also are rigid and bear similar chloroplasts. The metabolic Euglena species with larger plastids bearing pyrenoids and paramylon caps arose near the top of the tree. The loricates Strombomonas and Trachelomonas formed two well‐ supported, but paraphyletic, clades. The strong support for the individual clades confirmed the value of using lorica features as taxonomic criteria. The separation of the osmotrophic species A. longa, K. quartana, and H. ocellatus into different clades suggested that the loss of the photosynthetic ability has occurred multiple times.     

DISTRIBUTION,MORPHOLOGY, AND PHYLOGENY OF KLEBSORMIDIUM (KLEBSORMIDIALES,CHAROPHYCEAE) IN URBAN ENVIRONMENTS IN EUROPE1

Klebsormidium is a cosmopolitan genus of green algae, widespread in terrestrial and freshwater habitats. The classification of Klebsormidium is entirely based on morphological characters, and very little is understood about its phylogeny at the species level. We investigated the diversity and phylogenetic relationships of Klebsormidium in urban habitats in Europe by a combination of approaches including examination of field‐collected material, culture experiments conducted in many different combinations of factors, and phylogenetic analyses of the rbcL gene. Klebsormidium in European cities mainly occurs at the base of old walls, where it may produce green belts up to several meters in extent. Specimens from different cities showed a great morphological uniformity, consisting of long filaments 6–9 μm in width, with thin‐walled cylindrical cells and smooth wall, devoid of false branches, H‐shaped pieces, and biseriate parts. Conversely, the rbcL phylogeny showed a higher genetic diversity than expected from morphology. The strains were separated in four different clades supported by high bootstrap values and posterior probabilities. In culture, these clades differed in several characters, such as production of a superficial hydro‐repellent layer, tendency to break into short fragments, and inducibility of zoosporulation. On the basis of the taxonomic information available in the literature, most strains could not be identified unambiguously at the species level. The rbcL phylogeny showed no correspondence with classification based on morphology and suggested that the identity of many species, in particular the type species K. flaccidum (kütz.) P.C. Silva, Mattox et W. H. Blackw., needs critical reassessment.     

MOLECULAR PHYLOGENY OF THE GREEN ALGAL ORDER PRASIOLALES (TREBOUXIOPHYCEAE,CHLOROPHYTA)1

The systematics of the Prasiolales was investigated by phylogenetic inference based on analyses of the rbcL and 18S rRNA genes for representatives of all four genera currently attributed to this order (Prasiococcus, Prasiola, Prasiolopsis, Rosenvingiella), including all type species. The rbcL gene had higher sequence divergence than the 18S rRNA gene and was more useful for phylogenetic inference at the ranks of genus and species. In the rbcL gene phylogeny, three main clades were observed, corresponding to Prasiola, Prasiolopsis, and Rosenvingiella. Prasiococcus was nested among species of Prasiola occurring in subaerial and supralittoral habitats. Trichophilus welckeri Weber Bosse, a subaerial alga occurring in the fur of sloths in Amazonia, was closely related to Prasiolopsis ramosa Vischer. The species of Prasiola were grouped into three well‐supported clades comprising (i) marine species, (ii) freshwater and terrestrial species with linear blades, and (iii) terrestrial species with rounded or fan‐shaped blades. Sequence divergence was unexpectedly low in the marine group, which included species with different morphologies. For the 18S rRNA gene, the phylogenetic analyses produced several clades observed for the rbcL gene sequence analysis, but, due to very little sequence variation, it showed considerably lower resolution for inference at the species and genus levels. Due to the low support of some internal branches, the results of the analyses did not allow an unambiguous clarification of the origin and the early evolution of the Prasiolales.     

DISTRIBUTION OF THE XANTHOPHYLL LOROXANTHIN IN DESMIDS (CHAROPHYCEAE,CHLOROPHYTA)1

Forty-seven species of desmids, representing all four families, were examined for the presence of the xanthophyll loroxanthin by reverse-phase high-performance liquid chromatography. In the Desmidiaceae 28 of the 35 species examined possessed loroxanthin, and in the Mesotaeniaceae two of the six examined had loroxanthin present. All six species of the families Peniaceae and Closteriaceae examined possessed loroxanthin. Although the distribution of loroxanthin appears to be disjunct in the desmids and does not have strict taxonomic significance, it does follow a coherent pattern consistent with current ideas on desmid phylogeny. This pattern suggests that loroxanthin synthesis probably evolved once in the desmid lineage, with one or more subsequent reversals.     

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.     

PHYLOGENY OF THE BATRACHOSPERMALES (RHODOPHYTA) INFERRED FROM rbcL AND 18S RIBOSOMAL DNA GENE SEQUENCES

The sequence data from the large subunit of ribulose-1,5-bisphosphate carboxylase/oxygenase ( rbc L) gene and 18S ribosomal DNA (small subunit) of taxa in the freshwater rhodophyte order Batrachospermales were used to construct phylogenetic hypotheses. Taxa examined in this study represent four families, eight genera, and six sections of the genus Batrachospermum . In addition, Rhododraparnaldia oregonica Sheath, Whittick et Cole, was included in the analysis because it shares particular ultrastructural, reproductive, and morphological characteristics with members of the Batrachospermales and Acrochaetiales. The trees generated from each gene, as well as a combined data set, were largely congruent. Rhododraparnaldia consistently occurs on an early branch within the Acrochaetiales – Palmariales clade and does not appear to be a member of the Batrachospermales. In addition, Thorea violacea Bory de St. Vincent was not closely related to the other taxa of the Batrachospermales in all trees and hence the family Thoreaceae does not appear to be a natural grouping within this order. All other taxa analyzed, which are presently classified within this order, formed a monophyletic clade in most analyses. Psilosiphon scoparium Entwisle was not closely allied with the taxa of the Lemaneaceae, lending support to the newly proposed family Psilosiphonaceae. Sequence data from the remaining taxa of the Lemaneaceae support the concept of a derived monophyletic clade. The genus Batrachospermum appears to comprise many morphologically similar but distantly related taxa, which will need further investigation to resolve their taxonomic status. Tuomeya, Sirodotia and Nothocladus are retained at the generic level until further data are obtained.     

SPECIES PHYLOGENY OF COSMARIUM AND STAURASTRUM (DESMIDIACEAE) BASED ON RBC L SEQUENCES

Cosmarium and Staurastrum are the two most diverse genera of placoderm desmids (Family Desmidiaceae), with approximately 1100 and 800 species, respectively. Phylogenetic analysis of relationships of species has been extremely difficult. In a monograph of North American placoderm desmids, Prescott et al. described early phylogenetic work that concluded Staurastrum to be polyphyletic and certainly polymorphic. Likewise, Cosmarium has also been viewed as polyphyletic, and a number of workers have proposed splitting these genera. The classical view of West and West grouped species within each genus into two divisions and 6–8 sections based on wall features and semicell shape. We sequenced rbc L from 18 species of Cosmarium (2 divisions, 7 sections) and 12 species of Staurastrum (2 divisions and 7 sections) and performed a phylogenetic analysis (parsimony, maximum likelihood, bootstrap) using other placoderm desmids and Zygnematales as outgroups. The results exhibit little support for the monophyly of sections or divisions of the two genera. Furthermore, although there is support for the monophyly of clades within each genus, there is also support for a separate clade containing species from both genera.     

STUDIES ON ULTRASTRUCTURE AND THREE‐GENE PHYLOGENY OF THE GENUS MALLOMONAS (SYNUROPHYCEAE)1

The genus Mallomonas, a common and often abundant member of the planktic community in many freshwater habitats worldwide, consists of 180 species divided into 19 sections and 23 series. Classification of species is based largely on ultrastructural characteristics of the siliceous scales and bristles that collectively form a highly organized covering over the cell. However, the relative importance of the different siliceous features of the scales, such as the dome, V rib, and secondary structures, as well as the different types of scales, in understanding the evolution and phylogeny of the genus is little known. In this study, we investigated the scale and bristle ultrastructure, along with sequences of three genes, for 19 isolates (18 species) of Mallomonas (18 isolates were from Korean habitats). The isolates represented nine of the 19 sections. Sequences for both the nuclear SSU and LSU rDNA and plastid LSU of RUBISCO (rbcL) genes for each of the 19 Mallomonas isolates and four outgroups were determined. Bayesian and maximum‐likelihood (ML) analyses of the data revealed that Mallomonas consists of two strongly supported clades. Mallomonas bangladeshica (E. Takah. et T. Hayak.) Siver et A. P. Wolfe was at the base of the first clade that included taxa from the sections Planae and Heterospinae, both of which lack a V rib on the shield of the scales. Our results indicated that the sections Planae and Heterospinae should be combined. The second clade, with Mallomonas insignis Penard and Mallomonas punctifera Korshikov at the base, contained taxa from the sections Mallomonas, Striatae, Akrokomae, Annulatae, Torquatae, Punctiferae, and Insignes, all of which have V ribs or well‐developed marginal ribs on the scales. Sister relationships between Mallomonas and Striatae were strongly supported, but interrelations among the remaining sections were not resolved, probably due to inclusion of too few species. Our results suggest that the current classification of the genus Mallomonas at the section level will require some revision. Additional species will need to be added in future analyses.     

画眉科鸟类系统发育及分类地位商榷

原记录在中国有分布的鹛类计142种,占世界鹛类纪录种总数284种的50%.由于该类群的种类繁多,形态和生态习性多样,其分类问题历来受分类学者的关注,且争议较大.本文尝试以分子生物学手段来解决传统分类中遗留的问题.在系统发育分析中共有20属59种鹛类作为内群、14个相关物种作为外群,分子标记分别采用线粒体细胞色素b(Cty 6,846 bp)和核内重组引导基因(RAG-1,870 bp),以最大似然法和贝叶斯法重建系统发育树.结果表明,画眉科鸟类明显被分为2大支系,一支由噪鹛Garrulax、希鹛Minla、薮鹛Liocichla、相思鸟Leiothrix、奇鹛Heterophasia、穗鹛Stachyris、钩嘴鹛Pomatorhinus、凤鹛Yuhina、及部分雀鹛Alcippe等属的种类构成,另一支由鸦雀属Paradoxornis和山鹛Rhopophilus pekinensis、及部分雀鹛属的种类构成,2支都有相当高的支持率;各个属之间的相互关系大致得到解决,传统分类体系中噪鹛属、雀鹛属、鸦雀属、凤鹛属、穗鹛属的单系性不能成立;证实绣眼鸟属Zosteropidae、山鹛和林莺属Sylvia鸟类的亲缘关系较近,而白腹凤鹛Yuhina.zantholeuca、鹏鹛属Pteruthius和文须雀Panurus biarmicus与鹛类的亲缘关系较远;并提出小鳞胸鹪鹛Pnoepyga pusilla有可能不是画眉科种类,其分类地位需要重新认识.     

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.