The Rhizaria are an ill-defined but species-rich supergroup of mostly
unicellular
A unicellular organism, also known as a single-celled organism, is an organism that consists of a single cell, unlike a multicellular organism that consists of multiple cells. Organisms fall into two general categories: prokaryotic organisms and ...
eukaryote
Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacte ...
s. Except for the
Chlorarachniophytes
The chlorarachniophytes are a small group of exclusively marine algae widely distributed in tropical and temperate waters. They are typically mixotrophic, ingesting bacteria and smaller protists as well as conducting photosynthesis. Normally the ...
and three species in the genus Paulinella in the phylum
Cercozoa
Cercozoa is a phylum of diverse single-celled eukaryotes. They lack shared morphological characteristics at the microscopic level, and are instead defined by molecular phylogenies of rRNA and actin or polyubiquitin. They were the first major eu ...
, 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 also 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
In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to have ...
), 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 exoskeleton (thecae or loricas), which is in different clades within Rhizaria made out of opal (), celestite (), or calcite (). It 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.
Groups
The three main groups of Rhizaria are:
*
Cercozoa
Cercozoa is a phylum of diverse single-celled eukaryotes. They lack shared morphological characteristics at the microscopic level, and are instead defined by molecular phylogenies of rRNA and actin or polyubiquitin. They were the first major eu ...
– various amoebae and flagellates, usually with filose pseudopods and common in soil
*
Foraminifera
Foraminifera (; Latin for "hole bearers"; informally called "forams") are single-celled organisms, members of a phylum or class of amoeboid protists characterized by streaming granular Ectoplasm (cell biology), ectoplasm for catching food and ot ...
– amoeboids with reticulose pseudopods, common as marine
benthos
Benthos (), also known as benthon, is the community of organisms that live on, in, or near the bottom of a sea, river, lake, or stream, also known as the benthic zone.Radiolaria – amoeboids with axopods, common as marine
plankton
Plankton are the diverse collection of organisms found in Hydrosphere, water (or atmosphere, air) that are unable to propel themselves against a Ocean current, current (or wind). The individual organisms constituting plankton are called plankt ...
A few other groups may be included in the Cercozoa, but some trees appear closer to the Foraminifera. These are the
Phytomyxea
The Phytomyxea are a class of parasites that are cosmopolitan, obligate biotrophic protist parasites of plants, diatoms, oomycetes and brown algae. They are divided into the orders Plasmodiophorida (ICZN, or Plasmodiophoromycota, ICBN) and Pha ...
and
Ascetosporea
The Ascetosporea are a group of eukaryotes that are parasites of animals, especially marine invertebrates. The two groups, the haplosporids and paramyxids, are not particularly similar morphologically, but consistently group together on molecula ...
, parasites of plants and animals, respectively, and the peculiar amoeba '' Gromia''. The different groups of Rhizaria are considered close relatives based mainly on genetic similarities, and have been regarded as an extension of the Cercozoa. The name Rhizaria for the expanded group was introduced by Cavalier-Smith in 2002, who also included the
centrohelid
The centrohelids or centroheliozoa are a large group of heliozoan protists. They include both mobile and sessile forms, found in freshwater and marine environments, especially at some depth.
Characteristics
Individuals are unicellular and spher ...
s and
Apusozoa
The Apusozoa are an Obazoa phylum comprising several genera of flagellate eukaryotes. They are usually around 5–20 μm in size, and occur in soils and aquatic habitats, where they feed on bacteria. They are grouped together based on the prese ...
.
A noteworthy order that belongs to
Ascetosporea
The Ascetosporea are a group of eukaryotes that are parasites of animals, especially marine invertebrates. The two groups, the haplosporids and paramyxids, are not particularly similar morphologically, but consistently group together on molecula ...
is the
Mikrocytida
The Ascetosporea are a group of eukaryotes that are parasites of animals, especially marine invertebrates. The two groups, the haplosporids and paramyxids, are not particularly similar morphologically, but consistently group together on molecula ...
. These are parasites of
oyster
Oyster is the common name for a number of different families of salt-water bivalve molluscs that live in marine or brackish habitats. In some species, the valves are highly calcified, and many are somewhat irregular in shape. Many, but not al ...
s. This includes the causative agent of Denman Island Disease, ''Mikrocytos mackini'' a small (2−3 μm diameter) amitochondriate protistan.
Evolutionary relationships
Rhizaria are part of the SAR supergroup (Stramenopiles, Alveolates, Rhizaria), a grouping that had been presaged in 1993 through a study of mitochondrial morphologies. SAR is currently placed in the
Diaphoretickes
Diaphoretickes () is a major group of eukaryotic organisms, with over 400,000 species. The majority of the earth's biomass that carries out photosynthesis belongs to Diaphoretickes.
Diaphoretickes includes:
* Archaeplastida (comprising red alg ...
Cryptista
Cryptista is a clade of algae-like eukaryotes. It is most likely related to Archaeplastida which includes plants and many algae, within the larger group Diaphoretickes.
Although it has sometimes placed along with Haptista in the group Hacrobia, ...
, Haptista, and several minor clades.
Historically, many rhizarians were considered
animal
Animals are multicellular, eukaryotic organisms in the Kingdom (biology), biological kingdom Animalia. With few exceptions, animals Heterotroph, consume organic material, Cellular respiration#Aerobic respiration, breathe oxygen, are Motilit ...
s because of their motility and heterotrophy. However, when a simple animal-plant dichotomy was superseded by a recognition of additional kingdoms, taxonomists generally placed amoebae in the kingdom
Protista
A protist () is any eukaryotic organism (that is, an organism whose cells contain a cell nucleus) that is not an animal, plant, or fungus. While it is likely that protists share a common ancestor (the last eukaryotic common ancestor), the excl ...
. When scientists began examining the evolutionary relationships among eukaryotes in the 1970's, it became clear that the kingdom
Protista
A protist () is any eukaryotic organism (that is, an organism whose cells contain a cell nucleus) that is not an animal, plant, or fungus. While it is likely that protists share a common ancestor (the last eukaryotic common ancestor), the excl ...
was
paraphyletic
In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...
. Rhizaria appear to share a common ancestor with Stramenopiles and Alveolates forming part of the SAR (Stramenopiles+Alveolates+Rhizaria) super assemblage. Rhizaria has been supported by molecular phylogenetic studies as a monophyletic group. Biosynthesis of 24-isopropyl cholestane precursors in various rhizaria suggests a relevant ecological role already during the
Ediacaran
The Ediacaran Period ( ) is a geological period that spans 96 million years from the end of the Cryogenian Period 635 million years ago (Mya), to the beginning of the Cambrian Period 538.8 Mya. It marks the end of the Proterozoic Eon, and th ...
Cercozoa
Cercozoa is a phylum of diverse single-celled eukaryotes. They lack shared morphological characteristics at the microscopic level, and are instead defined by molecular phylogenies of rRNA and actin or polyubiquitin. They were the first major eu ...
and Retaria. Subsequently, Cercozoa and Retaria are also
monophyletic
In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
. The following cladogram depicts the evolutionary relationships between all rhizarian
class
Class or The Class may refer to:
Common uses not otherwise categorized
* Class (biology), a taxonomic rank
* Class (knowledge representation), a collection of individuals or objects
* Class (philosophy), an analytical concept used differentl ...
es, and is made after the works of Cavalier-Smith ''et al.'' (2018), Irwin ''et al.'' (2019) and Sierra ''et al.'' (2022):
Gallery
File:Cercomonas sp.jpg, ''Cercomonas'' sp. (
Cercozoa
Cercozoa is a phylum of diverse single-celled eukaryotes. They lack shared morphological characteristics at the microscopic level, and are instead defined by molecular phylogenies of rRNA and actin or polyubiquitin. They were the first major eu ...
:
Cercomonadida
Cercomonads are small flagellates, widespread in aqueous habitats and common in soils.
Characteristics
The cells are generally around 10 μm in length, without any shell or covering. They produce filose pseudopods to capture bacteria, but do no ...
Ebridea
The Ebridea is a group of phagotrophic flagellate eukaryotes present in marine coastal plankton communities worldwide. ''Ebria tripartita'' is one of two (possibly four) described extant species in the Ebridea.
Members of this group are named for ...
Spongomonadea
The spongomonads are a group of flagellated protist
A protist () is any eukaryotic organism (that is, an organism whose cells contain a cell nucleus) that is not an animal, plant, or fungus. While it is likely that protists share a common ...
Phaeodarea
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 ...
)
File:Clathrulina elegans - - Print - Iconographia Zoologica - Special Collections University of Amsterdam - UBAINV0274 113 04 0030.tif, ''
Clathrulina elegans
''Clathrulina elegans'' is a species of heliozoan eukaryote
Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryo ...
Chlorarachnion
The chlorarachniophytes are a small group of exclusively marine algae widely distributed in tropical and temperate waters. They are typically mixotrophic, ingesting bacteria and smaller protists as well as conducting photosynthesis. Normally the ...
'' sp. (Cercozoa: (
Chlorarachniophyta
The chlorarachniophytes are a small group of exclusively marine algae widely distributed in tropical and temperate waters. They are typically mixotrophic, ingesting bacteria and smaller protists as well as conducting photosynthesis. Normally the ...
Vampyrellidae
The family Vampyrellidae is a subgroup of the order Aconchulinida (formerly Vampyrellida) within the phylum Cercozoa. Based on molecular sequence data, the family currently comprises the genus ''Vampyrella'', and maybe several other vampyrellid ...
)
File:Die Gartenlaube (1890) b 083.jpg, '' Gromia'' (Cercozoa:
Gromiidea
Gromiida is an order of cercozoans. It is the only order in the class Gromiidea.
Taxonomy
Class Gromiidea Cavalier-Smith 2003 sensu Bass et al. 2009
* Order Gromiida Claparède & Lachmann 1856 s.s.
** Family Gromiidae Ruess 1862 romiina Delag ...
)
File:Powdery scab (Spongospora subterranea) of potatoes (1914) (14577838889).jpg,
Powdery scab
Powdery scab is a disease of potato tubers. It is caused by the cercozoan ''Spongospora subterranea'' f. sp. ''subterranea'' and is widespread in potato growing countries. Symptoms of powdery scab include small lesions in the early stages of the ...
(Cercozoa:
Plasmodiophorida
The plasmodiophores (also known as plasmophorids or plasmodiophorids) are a group of obligate endoparasitic protists belonging to the subphylum Endomyxa in Cercozoa. Taxonomically, they are united under a single family Plasmodiophoridae, order ...
)
File:Foraminifères de Ngapali.jpg, Foraminiferans ( Retaria:
Foraminifera
Foraminifera (; Latin for "hole bearers"; informally called "forams") are single-celled organisms, members of a phylum or class of amoeboid protists characterized by streaming granular Ectoplasm (cell biology), ectoplasm for catching food and ot ...
)
File:Mikrofoto.de-Radiolarien-3.jpg,
Polycystine
The polycystines are a group of radiolarians. They include the vast majority of the fossil radiolaria, as their skeletons are abundant in marine sediments, making them one of the most common groups of microfossils. These skeletons are composed ...