Pedinellales | |
---|---|
Light micrograph of Pteridomonas danica | |
Scientific classification | |
Domain: | Eukaryota |
Clade: | Diaphoretickes |
Clade: | SAR |
Clade: | Stramenopiles |
Phylum: | Gyrista |
Subphylum: | Ochrophytina |
Class: | Dictyochophyceae |
Order: | Pedinellales Zimmermann, Moestrup & Hallfors 1985 |
Family: | Actinomonadaceae Kent 1880 |
Genera | |
Synonyms | |
As class
As subclass
As order
As suborder |
Pedinellales (ICN) or Pedinellida (ICZN) is a group of single-celled algae found in both marine environments and freshwater. [3]
These are found in both freshwater and marine environments, and most genera are sessile, attached by posterior stalks. The flagellum is at the anterior of the cell, and the tentacles surround it, often capturing small prey drawn in by its current. The colored genera are Pedinella , Apedinella , Pseudopedinella , and Mesopedinella . Several more genera have lost their chloroplasts and feed entirely by phagocytosis. These are Parapedinella , Actinomonas , and Pteridomonas .
It also appears that certain heliozoa are actually derived pedinellids. Ciliophrys alternates between a mobile flagellate stage and a heliozoan feeding stage, where the body is contracted with extended axopods all over its surface, and the flagellum is curled up into a tight figure eight. The actinophryids, Actinophrys and Actinosphaerium, exist only in a heliozoan form with no flagellum and with more elaborate bundles of microtubules supporting their axopods. Their inclusion was argued by Mikrjukov and Patterson, who coined the term actinodine to refer specifically to this extended group.
Pedinellids were classified as heliozoans by some authors. The colored pedinellids were originally treated as a family of golden algae in the order Ochromonadales, promoted to an order Pedinellales by Zimmerman in 1984. Their relationship to the silicoflagellates became apparent some time later, and Patterson defined this rankless group for the two in 1994. Moestrup treated it as the class Dictyochophyceae, previously restricted to the silicoflagellates, while Cavalier-Smith defined a new class Actinochrysophyceae for them.
Pedinellids have been known since the 19th century, but were classified as either algae or heliozoan protozoa due to the absence of chloroplasts in some species. [4] Before the taxonomic revisions that took place in the late 20th century, pedinellids were treated as a single family Pedinellaceae, described in 1910 by Pascher. Some authors classified pedinellids within the order Ochromonadales, together with a variety of unrelated heterokonts such as Synuraceae and Bicosoecaceae, as part of the class Chrysophyceae or golden algae. [5] In 1985, phycologists Birthe Zimmermann, Øjvind Moestrup and Guy Hällfors classified them as an independent order, Pedinellales. [6] The same year, John J. Lee and coauthors segregated the phagotrophic (i.e. without chloroplasts) pedinellids as a different order Ciliophryida under the polyphyletic class Heliozoea. Other authors, such as Thomas Cavalier-Smith, went a step further and proposed treating pedinellids as a separate class, under the name of Pedinellea (spelt Pedinellophyceae under botanical nomenclature). [7]
In 1995, Charles J. O'Kelly and Daniel E. Wujek classified the Dictyochophyceae in three orders: Rhizochromulinales (containing the sole genus Rhizochromulina ), Pedinellales (containing all pedinellids) and Dictyochales (silicoflagellates), separating them from golden algae. [8] Simultaneously, Cavalier-Smith proposed a slightly different classification where he created a new class Actinochrysea to embrace these three groups, avoiding the usage of Dictyochophyceae as a name. He considered pedinellids as subclass Pedinellidae, with two orders: Pedinellales (with chloroplasts), containing the families Pedinellaceae and Apedinellaceae; and Ciliophryales (without chloroplasts), containing Actinomonadaceae and Ciliophryaceae. The remaining two groups, Rhizochromulinales and Dictyochales, were transferred to a separate subclass Dictyochidae. [2] The next year, Cavalier-Smith and Ema E. Chao recognized that the chloroplast losses were polyphyletic, and the family Actinomonadaceae was transferred to Pedinellales on the basis of a molecular phylogenetic analysis. [9]
In 2006, Cavalier-Smith once again rearranged the classification. He lowered Actinochrysea to a subclass Actinochrysia within a new class Hypogyristea, which also included pelagophytes as subclass Pelagophycia. He lumped together all pedinellid genera in the family Actinomonadaceae and new suborder Actinomonadineae, with the exception of Ciliophrys, which he included with the non-pedinellid Rhizochromulina in two families of a new suborder Ciliophryineae. [1] He maintained this classification in later years. [10] In 2018, he finally recognized class Dictyochophyceae, and created the subclass Pedinellia or Pedinellophycidae (equivalent to his earlier Actinochrysea/Actinochrysia) to lump together pedinellids and silicoflagellates. Pelagophytes remained under the subclass Pelagophycidae, and two genera of more uncertain placement under a third subclass Sulcophycidae: Olisthodiscus and Sulcochrysis . [11]
Cavalier-Smith's taxonomic scheme is not accepted by the scientific community. Instead, Moestrup's 1995 classification is more commonly supported, where pedinellids are all contained in the order Pedinellales, rendering Ciliophryales a junior synonym. They are considered part of Dictyochophyceae, along with orders Dictyochales (silicoflagellates), Rhizochromulinales (Rhizochromulina), [12] and a more recent fourth order Florenciellales. [13] In addition, pelagophytes are rejected as members of Dictyochophyceae, and instead form an independent class Pelagophyceae. [14] Within pedinellids, there have been several attempts to establish different families according to morphological data such as the presence or absence of stalks and chloroplasts. However, molecular analyses do not support these internal divisions. [12] Consequently, all pedinellids have been grouped into a single family, Actinomonadaceae, which has taxonomic preference due to being the earliest described, and all other proposed families (Pedinellaceae, Apedinellaceae, etc.) have become junior synonyms. [15]
Cavalier-Smith (1995) [2] | Cavalier-Smith (2006) [1] | Current [14] | Includes | |||||
---|---|---|---|---|---|---|---|---|
Pedinellidae | Pedinellales | Pedinellaceae | Pedinellales | Actinomonadineae | Actinomonadaceae | Pedinellales | Actinomonadaceae | Pedinella |
Pseudopedinella | ||||||||
Apedinellaceae | Apedinella | |||||||
Ciliophryales | Actinomonadaceae | Actinomonas | ||||||
Pteridomonas | ||||||||
Ciliophryaceae | Ciliophryineae | Ciliophryaceae | Ciliophrys | |||||
excluded | Rhizochromulinaceae | excluded | Rhizochromulina |
Pedinellids currently sum the following genera: [16] [10]
The Stramenopiles, also called Heterokonts, are a clade of organisms distinguished by the presence of stiff tripartite external hairs. In most species, the hairs are attached to flagella, in some they are attached to other areas of the cellular surface, and in some they have been secondarily lost. Stramenopiles represent one of the three major clades in the SAR supergroup, along with Alveolata and Rhizaria.
The haptophytes, classified either as the Haptophyta, Haptophytina or Prymnesiophyta, are a clade of algae.
The silicoflagellates are a small group of unicellular photosynthetic protists, or algae, belonging to the supergroup of eukaryotes known as Stramenopiles. They behave as plankton and are present in oceanic waters. They are well-known from harmful algal blooms that cause high mortality of fish. Additionally, they compose a rich fossil record represented by their silica skeletons.
The synurids are a small group of heterokont algae, found mostly in freshwater environments, characterized by cells covered in silica scales.
The axodines are a group of unicellular stramenopiles that includes silicoflagellate and rhizochromulinid algae, actinomonad heterotrophic flagellates and actinophryid heliozoa. Alternative classifications treat the dictyochophytes as heterokont algae, or as Chrysophyceae. Other overlapping taxonomic concepts include the Actinochrysophyceae, Actinochrysea or Dictyochophyceae sensu lato. The grouping was proposed on the basis of ultrastructural similarities, and is consistent with subsequent molecular comparisons.
Chromista is a proposed but polyphyletic biological kingdom, refined from the Chromalveolata, consisting of single-celled and multicellular eukaryotic species that share similar features in their photosynthetic organelles (plastids). It includes all eukaryotes whose plastids contain chlorophyll c and are surrounded by four membranes. If the ancestor already possessed chloroplasts derived by endosymbiosis from red algae, all non-photosynthetic Chromista have secondarily lost the ability to photosynthesise. Its members might have arisen independently as separate evolutionary groups from the last eukaryotic common ancestor.
The Chrysophyceae, usually called chrysophytes, chrysomonads, golden-brown algae or golden algae, are a large group of algae, found mostly in freshwater. Golden algae is also commonly used to refer to a single species, Prymnesium parvum, which causes fish kills.
Bicosoecida (ICZN) or Bicosoecales/Bicoecea (ICBN) is an order of Bikosea, a small group of unicellular flagellates, included among the stramenopiles. Informally known as bicosoecids, they are free-living cells, with no chloroplasts, and in some genera are encased in a lorica.
The raphidophytes, formally known as Raphidophycidae or Raphidophyceae, are a small group of eukaryotic algae that includes both marine and freshwater species. All raphidophytes are unicellular, with large cells, but no cell walls. Raphidophytes possess a pair of flagella, organised such that both originate from the same invagination. One flagellum points forwards, and is covered in hair-like mastigonemes, while the other points backwards across the cell surface, lying within a ventral groove. Raphidophytes contain numerous ellipsoid chloroplasts, which contain chlorophylls a, c1 and c2. They also make use of accessory pigments including β-carotene and diadinoxanthin. Unlike other heterokontophytes, raphidophytes do not possess the photoreceptive organelle typical of this group.
Ochrophytes, also known as heterokontophytes or stramenochromes, are a group of algae. They are the photosynthetic stramenopiles, a group of eukaryotes, organisms with a cell nucleus, characterized by the presence of two unequal flagella, one of which has tripartite hairs called mastigonemes. In particular, they are characterized by photosynthetic organelles or plastids enclosed by four membranes, with membrane-bound compartments called thylakoids organized in piles of three, chlorophyll a and c as their photosynthetic pigments, and additional pigments such as β-carotene and xanthophylls. Ochrophytes are one of the most diverse lineages of eukaryotes, containing ecologically important algae such as brown algae and diatoms. They are classified either as phylum Ochrophyta or Heterokontophyta, or as subphylum Ochrophytina within phylum Gyrista. Their plastids are of red algal origin.
Phaeothamniophycidae is a subclass of heterokont algae. It contains two orders, Phaeothamniales and Pleurochloridellales, and consists of species separated from Chrysophyceae.
Bigyra is a phylum of microscopic eukaryotes that are found at the base of the Stramenopiles clade. It includes three well-known heterotrophic groups Bicosoecida, Opalinata and Labyrinthulomycetes, as well as several small clades initially discovered through environmental DNA samples: Nanomonadea, Placididea, Opalomonadea and Eogyrea. The classification of Bigyra has changed several times since its origin, and its monophyly remains unresolved.
Opalozoa is a subphylum of heterotrophic protists of the phylum Bigyra, and is the sister group to Sagenista. Opalozoans are non-photosynthetic heterokonts that are ancestrally phagotrophic but many times have evolved to be osmotrophic saprotrophs in the gut of vertebrate animals.
Dictyochophyceae sensu lato is a photosynthetic lineage of heterokont algae.
Dictyocha is a genus of silicoflagellates, marine photosynthetic unicellular protists that take the form of either flagellates or axopodial amoebae. Described by Ehrenberg in 1837, Dictyocha contains many important species of the marine phytoplankton, some of them responsible for algal blooms that are toxic to fish.
Gyrista is a phylum of heterokont protists containing three diverse groups: the mostly photosynthetic Ochrophyta, the parasitic Pseudofungi, and the recently described group of nanoflagellates known as Bigyromonada. Members of this phylum are characterized by the presence of a helix or a double helix/ring system in the ciliary transition region.
Olisthodiscus is a genus of heterokont algae, present in marine or brackish waters. It is the only genus in the family Olisthodiscaceae, the order Olisthodiscales, and the class Olisthodiscophyceae. After a long history of controversial classifications, in 2021 it was recognized as a phylogenetically distinct lineage from the rest of ochrophyte classes.
Heliorapha is a genus of heliozoan protists, amoeboid eukaryotes with stiff axopodia radiating from their cells. It contains one species, Heliorapha azurina. It is classified within a monotypic family Helioraphidae inside the actinophryids, a group of heliozoa that belong to the Ochrophyta along with other protists such as diatoms and brown algae.
Octactis is a genus of silicoflagellates, marine photosynthetic unicellular protists that take the form of either flagellates or axopodial amoebae. Described by Josef Schiller in 1925, Octactis contains various species of marine phytoplankton, some of them responsible for algal blooms that are toxic to fish.
Ochromonadales is an order of single-celled algae belonging to the class Chrysophyceae, also known as golden algae. Initially it contained numerous groups of flagellates that were not closely related. During the late 20th century, advancements in molecular and ultrastructural studies allowed the transfer of many of these groups out of Ochromonadales, and the order was reduced to a single family Ochromonadaceae. They are aquatic single-celled flagellated algae, with two heterokont flagella each, some of which have secondarily lost their chloroplasts and appear colorless.