Radiozoa
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The Radiolaria, also called Radiozoa, are protozoa of diameter 0.1–0.2 mm that produce intricate
mineral In geology and mineralogy, a mineral or mineral species is, broadly speaking, a solid chemical compound with a fairly well-defined chemical composition and a specific crystal structure that occurs naturally in pure form.John P. Rafferty, ed. (2 ...
skeletons, typically with a central capsule dividing the
cell Cell most often refers to: * Cell (biology), the functional basic unit of life Cell may also refer to: Locations * Monastic cell, a small room, hut, or cave in which a religious recluse lives, alternatively the small precursor of a monastery ...
into the inner and outer portions of
endoplasm Endoplasm generally refers to the inner (often granulated), dense part of a cell's cytoplasm. This is opposed to the ectoplasm which is the outer (non-granulated) layer of the cytoplasm, which is typically watery and immediately adjacent to the ...
and ectoplasm. The elaborate mineral skeleton is usually made of
silica Silicon dioxide, also known as silica, is an oxide of silicon with the chemical formula , most commonly found in nature as quartz and in various living organisms. In many parts of the world, silica is the major constituent of sand. Silica is ...
. They are found as
zooplankton Zooplankton are the animal component of the planktonic community ("zoo" comes from the Greek word for ''animal''). Plankton are aquatic organisms that are unable to swim effectively against currents, and consequently drift or are carried along by ...
throughout the global ocean. As zooplankton, radiolarians are primarily
heterotroph A heterotroph (; ) is an organism that cannot produce its own food, instead taking nutrition from other sources of organic carbon, mainly plant or animal matter. In the food chain, heterotrophs are primary, secondary and tertiary consumers, but ...
ic, but many have photosynthetic
endosymbiont An ''endosymbiont'' or ''endobiont'' is any organism that lives within the body or cells of another organism most often, though not always, in a mutualistic relationship. (The term endosymbiosis is from the Greek: ἔνδον ''endon'' "within ...
s and are, therefore, considered
mixotroph A mixotroph is an organism that can use a mix of different sources of energy and carbon, instead of having a single trophic mode on the continuum from complete autotrophy at one end to heterotrophy at the other. It is estimated that mixotrophs comp ...
s. The skeletal remains of some types of radiolarians make up a large part of the cover of the ocean floor as
siliceous ooze Siliceous ooze is a type of biogenic pelagic sediment located on the deep ocean floor. Siliceous oozes are the least common of the deep sea sediments, and make up approximately 15% of the ocean floor. Oozes are defined as sediments which contain ...
. Due to their rapid change as species and intricate skeletons, radiolarians represent an important diagnostic
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
found from the Cambrian onwards.


Description

Radiolarians have many needle-like
pseudopod A pseudopod or pseudopodium (plural: pseudopods or pseudopodia) is a temporary arm-like projection of a eukaryotic cell membrane that is emerged in the direction of movement. Filled with cytoplasm, pseudopodia primarily consist of actin filament ...
s supported by bundles of microtubules, which aid in the radiolarian's buoyancy. The cell nucleus and most other organelles are in the endoplasm, while the ectoplasm is filled with frothy
vacuole A vacuole () is a membrane-bound organelle which is present in plant and fungal cells and some protist, animal, and bacterial cells. Vacuoles are essentially enclosed compartments which are filled with water containing inorganic and organic m ...
s and
lipid Lipids are a broad group of naturally-occurring molecules which includes fats, waxes, sterols, fat-soluble vitamins (such as vitamins A, D, E and K), monoglycerides, diglycerides, phospholipids, and others. The functions of lipids includ ...
droplets, keeping them buoyant. The radiolarian can often contain symbiotic algae, especially
zooxanthellae Zooxanthellae is a colloquial term for single-celled dinoflagellates that are able to live in symbiosis with diverse marine invertebrates including demosponges, corals, jellyfish, and nudibranchs. Most known zooxanthellae are in the genus ''S ...
, which provide most of the cell's energy. Some of this organization is found among the
heliozoa 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 f ...
, but those lack central capsules and only produce simple scales and spines. Some radiolarians are known for their resemblance to
regular polyhedra A regular polyhedron is a polyhedron whose symmetry group acts transitively on its flags. A regular polyhedron is highly symmetrical, being all of edge-transitive, vertex-transitive and face-transitive. In classical contexts, many different equival ...
, such as the icosahedron-shaped ''Circogonia icosahedra'' pictured below.


Taxonomy

The radiolarians belong to the supergroup
Rhizaria The Rhizaria are an ill-defined but 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 forami ...
together with (
amoeboid An amoeba (; less commonly spelled ameba or amœba; plural ''am(o)ebas'' or ''am(o)ebae'' ), often called an amoeboid, is a type of cell or unicellular organism with the ability to alter its shape, primarily by extending and retracting pseudopo ...
or flagellate) Cercozoa and (shelled amoeboid)
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 for catching food and other uses; and commonly ...
. Traditionally the radiolarians have been divided into four groups—
Acantharea The Acantharea (Acantharia) are a group of radiolarian protozoa, distinguished mainly by their strontium sulfate skeletons. Acantharians are heterotrophic marine microplankton that range in size from about 200 microns in diameter up to several ...
,
Nassellaria Nassellaria is an order of Rhizaria belonging to the class Radiolaria. The organisms of this order are characterized by a skeleton cross link with a cone or ring. Introduction Nassellaria is an order of Radiolaria under the class Polycystina. ...
,
Spumellaria Spumellaria is an order of radiolarians in the class Polycystinea. They are ameboid protists appearing in abundance in the world's oceans, possessing a radially-symmetrical silica (opal) skeleton that has ensured their preservation in fossil rec ...
and Phaeodarea. Phaeodaria is however now considered to be a Cercozoan. Nassellaria and Spumellaria both produce siliceous skeletons and were therefore grouped together in the group
Polycystina 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 ...
. Despite some initial suggestions to the contrary, this is also supported by molecular phylogenies. The Acantharea produce skeletons of
strontium sulfate Strontium sulfate (SrSO4) is the sulfate salt of strontium. It is a white crystalline powder and occurs in nature as the mineral celestine. It is poorly soluble in water to the extent of 1 part in 8,800. It is more soluble in dilute HCl and nitric ...
and is closely related to a peculiar genus, ''
Sticholonche ''Sticholonche'' is a genus of radiolarians with a single species, ''Sticholonche zanclea'', found in open oceans at depths of 99–510 metres. It is generally considered a heliozoan, placed in its own order, called the Taxopodida. However it ...
'' (
Taxopodida ''Sticholonche'' is a genus of radiolarians with a single species, ''Sticholonche zanclea'', found in open oceans at depths of 99–510 metres. It is generally considered a heliozoan, placed in its own order, called the Taxopodida. However it ...
), which lacks an internal skeleton and was for long time considered a
heliozoa 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 f ...
n. The Radiolaria can therefore be divided into two major lineages: Polycystina (Spumellaria + Nassellaria) and Spasmaria (Acantharia + Taxopodida). There are several higher-order groups that have been detected in molecular analyses of environmental data. Particularly, groups related to Acantharia and Spumellaria. These groups are so far completely unknown in terms of morphology and physiology and the radiolarian diversity is therefore likely to be much higher than what is currently known. The relationship between the Foraminifera and Radiolaria is also debated. Molecular trees supports their close relationship—a grouping termed Retaria. But whether they are sister lineages or if the Foraminifera should be included within the Radiolaria is not known.


Biogeography

In the diagram on the right, a Illustrates generalized radiolarian provinces and their relationship to water mass temperature (warm versus cool color shading) and circulation (gray arrows). Due to high-latitude water mass submergence under warm, stratified waters in lower latitudes, radiolarian species occupy habitats at multiple latitudes, and depths throughout the world oceans. Thus, marine sediments from the tropics reflect a composite of several vertically stacked faunal assemblages, some of which are contiguous with higher latitude surface assemblages. Sediments beneath polar waters include cosmopolitan deep-water radiolarians, as well as high-latitude endemic surface water species. Stars in (a) indicate the latitudes sampled, and the gray bars highlight the radiolarian assemblages included in each sedimentary composite. The horizontal purple bars indicate latitudes known for good radiolarian (silica) preservation, based on surface sediment composition. Material was copied from this source, which is available under
Creative Commons Attribution 4.0 International License
Data show that some species were extirpated from high latitudes but persisted in the tropics during the late Neogene, either by migration or range restriction (b). With predicted global warming, modern
Southern Ocean The Southern Ocean, also known as the Antarctic Ocean, comprises the southernmost waters of the World Ocean, generally taken to be south of 60° S latitude and encircling Antarctica. With a size of , it is regarded as the second-small ...
species will not be able to use migration or range contraction to escape environmental stressors, because their preferred cold-water habitats are disappearing from the globe (c). However, tropical endemic species may expand their ranges toward midlatitudes. The color polygons in all three panels represent generalized radiolarian biogeographic provinces, as well as their relative water mass temperatures (cooler colors indicate cooler temperatures, and vice versa). File:Circogoniaicosahedra ekw.jpg, ''Circogonia icosahedra'', radiolarian species shaped like a
regular icosahedron In geometry, a regular icosahedron ( or ) is a convex polyhedron with 20 faces, 30 edges and 12 vertices. It is one of the five Platonic solids, and the one with the most faces. It has five equilateral triangular faces meeting at each vertex. It ...
File:Anthocyrtium hispidum Haeckel - Radiolarian (34986365113).jpg, ''Anthocyrtium hispidum Haeckel''


Radiolarian shells

Radiolarians are unicellular predatory protists encased in elaborate globular shells usually made of silica and pierced with holes. Their name comes from the Latin for "radius". They catch prey by extending parts of their body through the holes. As with the silica frustules of diatoms, radiolarian shells can sink to the ocean floor when radiolarians die and become preserved as part of the ocean sediment. These remains, as
microfossils A microfossil is a fossil that is generally between 0.001 mm and 1 mm in size, the visual study of which requires the use of light or electron microscopy. A fossil which can be studied with the naked eye or low-powered magnification, ...
, provide valuable information about past oceanic conditions.Wassilieff, Maggy (2006
"Plankton - Animal plankton"
''Te Ara - the Encyclopedia of New Zealand''. Accessed: 2 November 2019.
File:Mikrofoto.de-Radiolarien 6.jpg, Like diatoms, radiolarians come in many shapes File:Theocotylissa ficus Ehrenberg - Radiolarian (34638920262).jpg, Also like diatoms, radiolarian shells are usually made of silicate File:Acantharian radiolarian Xiphacantha (Haeckel).jpg, However
acantharian The Acantharea (Acantharia) are a group of radiolarian protozoa, distinguished mainly by their strontium sulfate skeletons. Acantharians are heterotrophic marine microplankton that range in size from about 200 microns in diameter up to several ...
radiolarians have shells made from
strontium sulfate Strontium sulfate (SrSO4) is the sulfate salt of strontium. It is a white crystalline powder and occurs in nature as the mineral celestine. It is poorly soluble in water to the extent of 1 part in 8,800. It is more soluble in dilute HCl and nitric ...
crystals
File:Spherical radiolarian 2.jpg, Cutaway schematic diagram of a spherical radiolarian shell File:Cladococcus abietinus.jpg, ''Cladococcus abietinus''


Fossil record

The earliest known radiolaria date to the very start of the Cambrian period, appearing in the same beds as the first
small shelly fauna The small shelly fauna, small shelly fossils (SSF), or early skeletal fossils (ESF) are mineralized fossils, many only a few millimetres long, with a nearly continuous record from the latest stages of the Ediacaran to the end of the Early Cambri ...
—they may even be terminal Precambrian in age. They have significant differences from later radiolaria, with a different silica lattice structure and few, if any, spikes on the
test Test(s), testing, or TEST may refer to: * Test (assessment), an educational assessment intended to measure the respondents' knowledge or other abilities Arts and entertainment * ''Test'' (2013 film), an American film * ''Test'' (2014 film), ...
. About ninety percent of known radiolarian species are extinct. The skeletons, or tests, of ancient radiolarians are used in geological dating, including for
oil exploration Hydrocarbon exploration (or oil and gas exploration) is the search by petroleum geologists and geophysicists for deposits of hydrocarbons, particularly petroleum and natural gas, in the Earth using petroleum geology. Exploration methods Vis ...
and determination of ancient climates. Some common radiolarian fossils include '' Actinomma'', '' Heliosphaera'' and '' Hexadoridium''.


See also

*
Radiolarite Radiolarite is a siliceous, comparatively hard, fine-grained, chert-like, and homogeneous sedimentary rock that is composed predominantly of the microscopic remains of radiolarians. This term is also used for indurated radiolarian oozes and ...


References

* * * *


External links

*Radiolarians
*
Radiolaria.org
*
Tree Of Life—Radiolaria
{{Authority control * Amoeboids Extant Cambrian first appearances Bikont subphyla Taxa named by Thomas Cavalier-Smith