Almost since the creation of the genus Euglena (Ehrenberg), the taxa assigned to it have been separated, split apart, and reorganized into new genera based on morphological relationships, resulting in the creation of the genera Phacus (Dujardin), Lepocinclis (Perty), Astasia (Pringsheim), and Khawkinea ( Jahn and McKibben) based on intuitive methods. In an effort to assess the validity of these genera, we have used small subunit (SSU) rDNA data to generate a phylogenetic framework for these genera, with particular attention to the genus Euglena. Using the conserved sequence areas, we performed a phylogenetic analysis using parsimony, maximum likelihood, and distance methods. These different criteria have resulted in trees of the same topology. The euglenoid clade was composed of phagotrophic euglenoids at the base, which gave rise to phototrophs that in turn gave rise to osmotrophs. Among the photosynthetic taxa, the biflagellate form diverged prior to the uniflagellate form. Additionally, the need for a revision in the taxonomy of some of these genera was demonstrated. Currently, taxa from the photosynthetic genera Euglena, Phacus, and Lepocinclis do not form monophyletic clades, but are intermixed with each other as well as with the osmotrophic taxa, Astasia and Khawkinea.
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.
Previous studies using the nuclear SSU rDNA and partial LSU rDNA have demonstrated that the euglenoid loricate taxa form a monophyletic clade within the photosynthetic euglenoid lineage. It was unclear, however, whether the loricate genera Trachelomonas and Strombomonas were monophyletic. In order to determine the relationships among the loricate taxa, SSU and LSU nuclear rDNA sequences were obtained for eight Strombomonas and 25 Trachelomonas strains and combined in a multigene phylogenetic analysis. Conserved regions of the aligned data set were used to generate maximum-likelihood (ML) and Bayesian phylogenies. Both methods recovered a strongly supported monophyletic loricate clade with Strombomonas and Trachelomonas species separated into two sister clades. Taxa in the genus Strombomonas sorted into three subclades. Within the genus Trachelomonas, five strongly supported subclades were recovered in all analyses. Key morphological features could be attributed to each of the subclades, with the major separation being that all of the spine-bearing taxa were located in two sister subclades, while the more rounded, spineless taxa formed the remaining three subclades. The separation of genera and subclades was supported by 42 distinct molecular signatures (33 in Trachelomonas and nine in Strombomonas). The morphological and molecular data supported the retention of Trachelomonas and Strombomonas as separate loricate genera.
The euglenoid genus Monomorphina was defined by Mereschowsky in 1877 to include rigid euglenoids that were pyriform in lateral view, had a hyaline spine at the posterior end, and one to few parietal chloroplasts typically without pyrenoids. The genus included taxa previously assigned to Phacus Dujardin or Euglena Ehrenberg. The general structure of Monomorphina aenigmatica comb. nov. is described on the basis of light microscopy and scanning and transmission electron microscopy. Cells were pear-shaped in lateral view, rounded at the anterior end and narrowed posteriorly, tapering into a long twisted tail. The pellicle had helically arranged strips spiralled in a counter-clockwise fashion. A distinctive feature of M. aenigmatica was the presence of a single chloroplast bearing a pyrenoid, capped with a paramylon plate. The large parietal chloroplast extended along most of the cell with three prominent cup-shaped paramylon caps on the external face. In transverse section, the chloroplast appeared C-shaped. Because of the ambiguity surrounding the original descriptions used to diagnose this taxon, we designated an epitype for Monomorphina aenigmatica. Morphological features of this species were compared to other members of the genus.
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