| Centrohelids | |
|---|---|
 | |
| Raphidiophrys contractilis | |
Scientific classification  | |
| Domain: | Eukaryota |
| Clade: | Diaphoretickes |
| Phylum: | Haptista |
| Class: | Centroplasthelida Febvre‐Chevalier & Febvre, 1984 [1] |
| Orders [1] [2] | |
Incertae sedis | |
| Synonyms | |
The centrohelids or centroheliozoa are a large group of heliozoan protists. [4] They include both mobile and sessile forms, found in freshwater and marine environments, especially at some depth.
Individuals are unicellular and spherical, usually around 30–80 μm in diameter, and covered with long radial axopods, narrow cellular projections that capture food and allow mobile forms to move about.
A few genera have no cell covering, but most have a gelatinous coat holding scales and spines, produced in special deposition vesicles. These may be organic or siliceous and come in various shapes and sizes. For instance, in Raphidiophrys the coat extends along the bases of the axopods, covering them with curved spicules that give them a pine-treeish look, and in Raphidiocystis there are both short cup-shaped spicules and long tubular spicules that are only a little shorter than the axopods. Some other common genera include Heterophrys , Actinocystis, and Oxnerella .
The axopods of centrohelids are supported by microtubules in a triangular-hexagonal array, which arise from a tripartite granule called the centroplast at the center of the cell. Axopods with a similar array occur in gymnosphaerids, which have traditionally been considered centrohelids (though sometimes in a separate order from the others). This was questioned when it was found they have mitochondria with tubular cristae, as do other heliozoa, while in centrohelids the cristae are flat. Although this is no longer considered a very reliable character, on balance gymnosphaerids seem to be a separate group.
The evolutionary position of the centrohelids is not clear. Structural comparisons with other groups are difficult, in part because no flagella occur among centrohelids, and genetic studies have been more or less inconclusive. Cavalier-Smith has suggested they may be related to the Rhizaria, [5] but for the most part they are left with uncertain relations to other groups. A 2009 paper suggests that they may be related to the cryptophytes and haptophytes (see Cryptomonads-haptophytes assemblage). [6] They are currently classified as Hacrobia, under the Plants+HC clade, although some research studies have found evidence against the monophyly of this group. [7] Centrohelids were previously divided into two orders with contrasting scale morphology and ultrastructure: Pterocystida and Acanthocystida. [8] Posterior molecular studies of 2018 have rearranged the classification of centrohelids into two taxa: Pterocystida and Panacanthocystida, which includes both Acanthocystida and the genus Yogsothoth . [2] [1]
The modern classification of centrohelids, as of 2019: [2] [1]
The gymnosphaerids are a small group of heliozoan protists found in marine environments. They tend to be roughly spherical with radially directed axopods, supported by microtubules in a triangular-hexagonal array arising from an amorphous central granule.
Nucleariida is a group of amoebae with filose pseudopods, known mostly from soils and freshwater. They are distinguished from the superficially similar vampyrellids mainly by having mitochondria with discoid cristae, in the absence of superficial granules, and in the way they consume food.
Heliozoa, commonly known as sun-animalcules, are microbial eukaryotes (protists) with stiff arms (axopodia) radiating from their spherical bodies, which are responsible for their common name. The axopodia are microtubule-supported projections from the amoeboid cell body, and are variously used for capturing food, sensation, movement, and attachment. They are similar to Radiolaria, but they are distinguished from them by lacking central capsules and other complex skeletal elements, although some produce simple scales and spines. They may be found in both freshwater and marine environments.
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.
Cercozoa is a phylum of diverse single-celled eukaryotes. They lack shared morphological characteristics at the microscopic level, and are instead united by molecular phylogenies of rRNA and actin or polyubiquitin. They were the first major eukaryotic group to be recognized mainly through molecular phylogenies. 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.
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.
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.
Chromalveolata was a eukaryote supergroup present in a major classification of 2005, then regarded as one of the six major groups within the eukaryotes. It was a refinement of the kingdom Chromista, first proposed by Thomas Cavalier-Smith in 1981. Chromalveolata was proposed to represent the organisms descended from a single secondary endosymbiosis involving a red alga and a bikont. The plastids in these organisms are those that contain chlorophyll c.
Telonemia is a phylum of microscopic eukaryotes. They are unicellular free-living flagellates with a unique combination of cell structures, including a highly complex cytoskeleton unseen in other eukaryotes. They present characteristics similar to their sister group, the SAR supergroup, such as cortical alveoli, tripartite mastigonemes and filopodia. Together, the two lineages compose the TSAR clade. They are classified in three genera and seven species, although numerous undescribed lineages are known. They are detected in all marine and freshwater environments, where they prey on bacteria and small phytoplankton through phagotrophy.
Imbricatea is a class of Rhizaria characterised by silica scales. It is sometimes described as "Imbricatea/Silicofilosea", due to the similarity of those two groupings. Imbricatea is divided into the orders Euglyphida and Thaumatomonadida.
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.
Telonema is a genus of single-celled organisms.
Raphidiophrys is a genus of centrohelid with radiating axopodia. R. intermedia is found in the bottom sludge of freshwater bodies in Canada, Chile, Argentina, Australia, New Zealand, Malaysia, Russia, and central Europe. Raphidiophrys have bipartite scales are a defining characteristic among species. Differences in type and size of scales are used to differentiate amongst the members of this genus. The genus Raphidiophrys was discovered in 1867 by W. Archer. Raphidiophrys is one of very few centrohelids in which dimorphism has been shown.
Rabdiophrys is a genus of amoeboid rhizarians. It has 19 species, including the species Rabdiophrys anulifera.
Raphidiophryidae is a family of mostly freshwater centrohelids. It is the sister family of Acanthocystidae, sharing the trait of presenting silica scales and comprising the clade Chalarothoracina. Two genera, Raphidiophrys and Polyplacocystis, have been discovered so far.
Cryptista is a clade of alga-like eukaryotes. It is most likely related to Archaeplastida which includes plants and many algae, within the larger group Diaphoretickes.
Haptista is a proposed group of protists made up of centrohelids and haptophytes. Phylogenomic studies indicate that Haptista, together with Ancoracysta twista, forms a sister clade to the SAR+Telonemia supergroup, but it may also be sister to the Cryptista (+Archaeplastida). It is thus one of the earliest diverging Diaphoretickes.
Endohelea is a proposed clade of eukaryotes that are related to Archaeplastida and the SAR supergroup. They used to be considered heliozoans, but phylogenetically they belong to a group of microorganisms known as Cryptista.
Yogsothoth is a genus of centrohelid protists, distinguished by the shape and arrangement of their external scales as well as their colonial life strategy. It was described in November 2018 by Shɨshkin and Zlatogursky, and is part of a newly described clade of centrohelids, determined as such by analysis of molecular data.