Another Origin of Coloniality in Volvocaleans: The Phylogenetic Position of Pyrobotrys Arnoldi (Spondylomoraceae,Volvocales)

ABSTRACT. Colonial volvocaleans (Chlorophyceae) are used as a standard model of multicellular evolution. However, the phylogenetic position of the colonial volvocalean family Spondylomoraceae has yet to be resolved. To examine this, the molecular phylogenies of Pyrobotrys stellata and Pyrobotrys squarrosa were analyzed using combined 18S rRNA, RUBISCO large subunit, and P700 chl a‐apoprotein A2 gene sequences. In the phylogenetic trees, Pyrobotrys belonged to the clade Caudivolvoxa and was not closely related to other colonial volvocalean flagellates. The results indicate that colony formation of Spondylomoraceae independently evolved from unicellular volvocaleans. The phylogenetic position of problematic “Pascherina tetras” SAG 159‐1 was also analyzed.     

Ultrastructure and 18S rDNA sequence analysis of Wobblia lunata gen. et sp. nov., a new heterotrophic flagellate (Stramenopiles, Incertae sedis)

A new heterotrophic flagellate Wobblia lunata gen. et sp. nov. is described. This organism usually attaches to the substratum showing a wobbling motion, and sometimes glides on the substratum or swims freely in the medium. W. lunata has various features characteristic of the stramenopiles. These include a hairy flagellum with tripartite tubular hairs, a mitochondrion with tubular cristae, arrangement of flagellar apparatus components and a double helix in the flagellar transition zone. W. lunata shares a double helix with heterotrophic stramenopiles, including Developayella elegans, oomycetes, hyphochytrids, opalinids and proteromonads, and could be placed in the phylum Bigyra Cavalier-Smith. However, from 18S rDNA tree analysis, these organisms form two distantly-related clades in the stramenopiles, and Wobblia appears at the base of the stramenopiles. Evaluation of morphological features and comparison of 18S rDNA sequences indicate that W. lunata is a member of the stramenopiles, but it is distinct from any other stramenopiles so far described. Its phylogenetic position within the stramenopiles is uncertain and therefore W. lunata is described as a stramenopile incertae sedis.     

Ultrastructure and taxonomy of a marine photosynthetic stramenopile Phaeomonas parva gen. et sp. nov. (Pinguiophyceae) with emphasis on the flagellar apparatus architecture

Phaeomonas parva gen. et sp. nov., a marine photosynthetic stramenopile from oceanic water near the Caroline Islands, is described. Cells are naked and spherical to ovoid. The alga is motile with two laterally inserted flagella during the light period, whereas during the dark period, it absorbs the flagella and rounds up. The anterior (immature, No. 2) long flagellum possesses tubular tripartite mastigonemes. The posterior (mature, No. 1) short flagellum is smooth and has autofluorescence at the base. The cupshaped, yellowish‐brown chloroplast occupies the posterior half of the cell, and a pyrenoid occurs in the inner cavity of the cup‐shaped chloroplast. The flagellar apparatus has several unusual features. Two basal plates and a two‐gyred proximal helix in the flagellar transitional region may suggest that P. parva is related to the Pelagophyceae, Dictyochophyceae and Sulcochrysis biplastida, a photosynthetic stramenopile of uncertain taxonomic position. The R3 and R4 roots form a loop that resembles phagotrophic chrysophytes. However, this resemblance is superficial because Phaeomonas is not phagotrophic, its R3 root has a different number of microtubules and its R3 root does not split to form a food‐uptake mouth. Phaeomonas has a ‘bypassing root’, which is found only with the Phaeophyceae, Giraudyopsis stellifera (Chrysomerophyceae), and Ankylochrysis lutea (probably a member of the Pelagophyceae). The taxonomic position of P. parva could not be determined solely from ultrastructural features. However, molecular phylogeny and biochemical analyses (published separately) strongly supported a relationship between P. parva and four other monotypic strameno‐piles, Glossomastix, Pinguiochrysis, Pinguiococcus and Polypodochrysis. Although these algae are morphologically distinct, they have unusually high percentages of polyunsaturated fatty acids, especially eicosapentoic acid. This unusual assemblage of stramenopiles is classified in a new class, the Pinguiophyceae (published separately), and P. parva is its only biflagellate member.     

Morphological evolution of silica scales in the freshwater genus Synura (Stramenopiles)

A high degree of morphological variability is expressed between the ornately sculptured siliceous scales formed by species in the chrysophycean genus, Synura. In this study, we aimed to uncover the general principles and trends underlying the evolution of scale morphology in this genus. We assessed the relationships among thirty extant Synura species using a robust molecular analysis that included six genes, coupled with morphological characterization of the species‐specific scales. The analysis was further enriched with addition of morphological information from fossil specimens and by including the unique modern species, Synura punctulosa. We inferred the phylogenetic position of the morphologically unique S. punctulosa, to be an ancient Synura lineage related to S. splendida in the section Curtispinae. Some morphological traits, including development of a keel or a labyrinth ribbing pattern on the scale, appeared once in evolution, whereas other structures, such as a hexagonal meshwork pattern, originated independently several times over geologic time. We further uncovered numerous construction principles governing scale morphology and evolution, as follows: (i) scale roundness and pore diameter decreased during evolution; (ii) elongated scales became strengthened by a higher number of struts or ribs; (iii) as a consequence of scale biogenesis, scales with spines possessed smaller basal holes than scales with a keel and; and (iv) the keel area was proportional to scale area, indicating its potential value in strengthening the scale against breakage.     

EFFECT OF TAXON SAMPLING,CHARACTER WEIGHTING,AND COMBINED DATA ON THE INTERPRETATION OF RELATIONSHIPS AMONG THE HETEROKONT ALGAE1

Nuclear ribosomal small subunit and chloroplast rbcL sequence data for heterokont algae and potential outgroup taxa were analyzed separately and together using maximum parsimony. A series of taxon sampling and character weighting experiments was performed. Traditional classes (e.g. diatoms, Phaeophyceae, etc.) were monophyletic in most analyses of either data set and in analyses of combined data. Relationships among classes and of heterokont algae to outgroup taxa were sensitive to taxon sampling. Bootstrap (BS) values were not always predictive of stability of nodes in taxon sampling experiments or between analyses of different data sets. Reweighting sites by the rescaled consistency index artificially inflates BS values in the analysis of rbcL data. Inclusion of the third codon position from rbcL enhanced signal despite the superficial appearance of mutational saturation. Incongruence between data sets was largely due to placement of a few problematic taxa, and so data were combined. BS values for the combined analysis were much higher than for analyses of each data set alone, although combining data did not improve support for heterokont monophyly.     

Ultrastructure of Prorodon (Ciliophora, Prostomatida) Ii. Oral Cortex and Phylogenetic Conclusions

ABSTRACT. The ultrastructure of the oral region and the ultrastructural architecture of the basket of Prorodon aklitolophon and Prorodon teres are described. the oral region of Prorodon consists of: 1) A circumoral kinety at the anterior pole of the cell surrounding the typically slit-shaped cytostomial funnel. This kinety is composed of inversely oriented dikinetids in which both kinetosomes are ciliated and are associated with a very short postciliary microtubular ribbon and a few transverse microtubules; 2) Three brush rows aligned in parallel and extended meridionally in the anterior part of the cell. the individual brush rows consist of dikinetids, but in contrast to the dikinetids around the cytopharynx they are not inverted and only the anterior kinetosomes bear specialized short brush cilia and are associated with a divergent-tangential transverse microtubular ribbon. the posterior kinetosome is non-ciliated and bears a prominent convergent postciliary microtubular ribbon. Schematized dikinetid patterns of both oral regions of Prorodon are provided. In addition, a three-dimensional reconstruction of the basket of the genus Prorodon based on serial thin sections is presented. A phylogenetic tree, mainly based on stomatogenic data, is given to show the phylogenetic relationships of some prostomatid genera as well as the hypothesized sistergroup relationship of colpodid and prostomatid ciliates.