Cercozoa | |
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Cercomonas | |
Scientific classification | |
Domain: | Eukaryota |
Clade: | Diaphoretickes |
Clade: | SAR |
Clade: | Rhizaria |
Phylum: | Cercozoa Cavalier-Smith, 1998 [1] emend. Adl et al., 2005 emend. Cavalier-Smith, 2018 [2] |
Classes | |
Synonyms | |
Cercozoa (now synonymised with Filosa) [2] is a phylum of diverse single-celled eukaryotes. [4] [5] They lack shared morphological characteristics at the microscopic level, [6] and are instead united by molecular phylogenies of rRNA and actin or polyubiquitin. [7] They were the first major eukaryotic group to be recognized mainly through molecular phylogenies. [8] They are the natural predators of many species of bacteria. They are closely related to the phylum Retaria, comprising amoeboids that usually have complex shells, and together form a supergroup called Rhizaria. [2]
The group includes most amoeboids and flagellates that feed by means of filose pseudopods. These may be restricted to part of the cell surface, but there is never a true cytostome or mouth as found in many other protozoa. They show a variety of forms [9] and have proven difficult to define in terms of structural characteristics, although their unity is strongly supported by phylogenetic studies.
Some cercozoans are grouped by whether they are "filose" or "reticulose" in the behavior of their cytoskeleton when moving: [10]
Other important ecological groups are:
As well as being highly diverse in morphology and physiology, Cercozoa also shows high ecological diversity. [12] The phylum Cercozoa includes many of the most abundant and ecologically significant protozoa in soil, marine and freshwater ecosystems. [8]
Soil-dwelling cercozoans are one of the dominant groups of free-living eukaryotic microorganisms found in temperate soils, accounting for around 30% of identifiable protozoan DNA in arid or semi-arid soils and 15% in more humid soils. In transcriptomic analyses they account for 40-60% of all identifiable protozoan RNA found in forest and grassland soils. They also comprise 9-24% of all operational taxonomic units found in the ocean floor. [12]
Some cercozoa are coprophilic or coprozoic, meaning they use feces as a source of nutrients or as transport through animal hosts. The faecal habitat is an understudied reservoir of microbial eukaryotic diversity, dominated by amoeboflagellates from the phylum Cercozoa. Strongly coprophilic examples of cercozoa are the flagellates Cercomonas , Proleptomonas and Helkesimastix , and the sorocarpic amoeba Guttulinopsis . Many new cercozoan lineages, especially among sarcomonads, have been discovered through phylogenetic sampling of feces because they appear preferentially in this medium. [13]
Cercozoan bacterivores (i.e. predators of bacteria) are highly diverse and important in the plant phyllosphere, the leaf surfaces of plants. Particularly sarcomonads, with their ability to cyst, feed and multiply within hours, are perfectly adapted to the fluctuating environmental factors in the phyllosphere. Their predation causes shifts in the bacterial communities: they reduce populations of alphaproteobacteria and betaproteobacteria, which are less resistant to their grazing, in favour of other bacterial populations such as gammaproteobacteria. [14]
Paraphyletic Cercozoa [2] |
Monophyletic Cercozoa [15] |
Originally, Cercozoa contained both Filosa and Endomyxa, according to phylogenetic analyses using ribosomal RNA and tubulin. These analyses also confirmed Cercozoa as the sister group of Retaria within the supergroup Rhizaria. [10] [16]
However, the monophyly of the group was still uncertain. Posterior multigene phylogenetic analyses consistently found Cercozoa to be paraphyletic, because Endomyxa clustered next to Retaria instead of Filosa. [17] [18] [19] Because of this, Endomyxa was excluded from Cercozoa, which became a synonym of Filosa. [2]
More recent phylogenomic analyses with better sampling recovered a sister relationship between Filosa (=Cercozoa) and Endomyxa once again, [15] although the modern classification of eukaryotes retains Endomyxa, Cercozoa and Retaria as separate phyla within Rhizaria. [20]
The phylum Cercozoa previously contained both Filosa and Endomyxa, but in the latest classifications Endomyxa has been excluded, and Cercozoa is now synonymous with Filosa. It is composed of two subphyla: Monadofilosa and Reticulofilosa. According to multigene phylogenetic analyses, Monadofilosa is a robust clade, in which the deepest branching group is Metromonadea, followed by Helkesea as the second group (together forming the paraphyletic Eoglissa) before the divergence of the clade Ventrifilosa (Imbricatea, Sarcomonadea and Thecofilosea). On the other hand, Reticulofilosa is probably paraphyletic, with Granofilosea diverging earlier than Chlorarachnea, which makes Chlorarachnea the sister group of Monadofilosa. [2]
Cercozoa | |
A more recent phylogenomic analysis recovered both Monadofilosa and Reticulofilosa as monophyletic within the clade Filosa. [15]
In addition to the known Granofilosea, Chlorarachnea and Monadofilosa, a variety of clades inside Cercozoa have been discovered in other analyses and have slowly been described and named, such as Tremulida (previously known as Novel Clade 11) [16] and Aquavolonida (Novel Clade 10), [21] although their specific positions among the two main cercozoan subphyla have yet to be refined. These two orders have been classified as the class Skiomonadea, within Reticulofilosa. [2]
The classification of Cercozoa was revised in 2018: [2]
The Rhizaria are a diverse and species-rich supergroup of mostly unicellular eukaryotes. Except for the Chlorarachniophytes and three species in the genus Paulinella in the phylum Cercozoa, they are all non-photosynthethic, but many foraminifera and radiolaria have a symbiotic relationship with unicellular algae. A multicellular form, Guttulinopsis vulgaris, a cellular slime mold, has been described. This group was used by Cavalier-Smith in 2002, although the term "Rhizaria" had been long used for clades within the currently recognized taxon. Being described mainly from rDNA sequences, they vary considerably in form, having no clear morphological distinctive characters (synapomorphies), but for the most part they are amoeboids with filose, reticulose, or microtubule-supported pseudopods. In the absence of an apomorphy, the group is ill-defined, and its composition has been very fluid. Some Rhizaria possess mineral exoskeletons, which are in different clades within Rhizaria made out of opal, celestite, or calcite. Certain species can attain sizes of more than a centimeter with some species being able to form cylindrical colonies approximately 1 cm in diameter and greater than 1 m in length. They feed by capturing and engulfing prey with the extensions of their pseudopodia; forms that are symbiotic with unicellular algae contribute significantly to the total primary production of the ocean.
Phaeodarea or Phaeodaria is a group of amoeboid cercozoan organisms. They are traditionally considered radiolarians, but in molecular trees do not appear to be close relatives of the other groups, and are instead placed among the Cercozoa. They are distinguished by the structure of their central capsule and by the presence of a phaeodium, an aggregate of waste particles within the cell.
Cercomonads are small amoeboflagellates, widespread in aqueous habitats and common in soils.
The tectofilosids are a group of filose amoebae with shells. These are composed of organic materials and sometimes collected debris, in contrast to the euglyphids, which produce shells from siliceous scales. The shell usually has a single opening, but in Amphitrema and a few other genera it has two on opposite ends. The cell itself occupies most of the shell. They are most often found on marsh plants such as Sphagnum.
Monadofilosa is a grouping of Cercozoa. These organisms are single-celled amoeboid protists.
Telonemia is a phylum of microscopic eukaryotes commonly known as telonemids. They are unicellular free-living flagellates with a unique combination of cell structures, including a highly complex cytoskeleton unseen in other eukaryotes.
Retaria is a clade within the supergroup Rhizaria containing the Foraminifera and the Radiolaria. In 2019, the Retaria were recognized as a basal Rhizaria group, as sister of the Cercozoa.
Endomyxa is a group of eukaryotic organisms in the supergroup Rhizaria. They were initially a subphylum of Cercozoa and later a subphylum of Retaria, but several analyses have proven they are a phylogenetically separate lineage, and Endomyxa is currently regarded as its own phylum.
Thecofilosea is a class of unicellular testate amoebae belonging to the phylum Cercozoa. They are amoeboflagellates, organisms with flagella and pseudopodia, distinguished from other cercozoa by their scale-lacking test composed of organic material. They are closely related to the Imbricatea, a group of testate amoebae with tests composed of inorganic silica scales.
The spongomonads are a group of flagellated protists in the phylum Cercozoa. Taxonomically, they compose the family Sarcomonadidae and order Sarcomonadida. They were originally placed among the Reticulofilosa, but were later transferred to Monadofilosa. It includes only two genera:
The sarcomonads or class Sarcomonadea are a group of amoeboid biciliate protists in the phylum Cercozoa. They are characterized by a propensity to move through gliding on their posterior cilium or through filopodia, a lack of scales or external theca, a soft cell surface without obvious cortical filamentous or membranous skeleton, two cilia without scales or hairs, tubular mitochondrial cristae, near-spherical extrusomes, and a microbody attached to the nucleus.
Halvaria is a taxonomic grouping of protists that includes Alveolata and Stramenopiles (Heterokonta).
The vampyrellids, colloquially known as vampire amoebae, are a group of free-living predatory amoebae classified as part of the lineage Endomyxa. They are distinguished from other groups of amoebae by their irregular cell shape with propensity to fuse and split like plasmodial organisms, and their life cycle with a digestive cyst stage that digests the gathered food. They appear worldwide in marine, brackish, freshwater and soil habitats. They are important predators of an enormous variety of microscopic organisms, from algae to fungi and animals. They are also known as aconchulinid amoebae.
The plasmodiophores are a group of obligate endoparasitic protists belonging to the subphylum Endomyxa in Cercozoa. Taxonomically, they are united under a single family Plasmodiophoridae, order Plasmodiophorida, sister to the phagomyxids.
Cryptofilida is an order of small heterotrophic protists in the phylum Cercozoa. They are filose amoebae that lack cilia and gliding, and are instead characterized by movement through branching or unbranched granular filopodia that are appressed to the substrate during their feeding.
The glissomonads are a group of bacterivorous gliding flagellated protists that compose the order Glissomonadida, in the amoeboflagellate phylum Cercozoa. They comprise a vast, largely undescribed diversity of soil and freshwater organisms. They are the sister group to cercomonads; the two orders form a solid clade of gliding soil-dwelling flagellates called Pediglissa.
Euglyphia is a group of imbricate protists in the phylum Cercozoa. They are unicellular eukaryotes characterized by a cell body covered in large imbricate scales, and an apical aperture through which they extend either filose pseudopodia or two cilia of different sizes that are not used for gliding.
Pediglissa is a subclass of phagotrophic protists that inhabit soil or freshwater habitats. They were defined in 2018 according to phylogenetic analyses that showed a clade containing the orders Cercomonadida and Glissomonadida. They're the sister group of Paracercomonadida.
Helkesida is a group of microscopic protists belonging to the supergroup Rhizaria, both discovered through molecular phylogenetic analyses. It contains amoeboid flagellates with two flagella. They are either free-living, mostly on fecal matter, or live inside the gut of animals. Among these amoebae, one lineage has independently evolved aggregative multicellularity similarly to slime moulds.
Limnofila is a genus of heterotrophic protists that live in freshwater habitats and feed on bacteria. They are also present in the soil ecosystem, where they play an important role as predators of bacteria. They are classified as a single family Limnofilidae and order Limnofilida.