Radiolaria

Radiolaria Temporal range: Cambrian – Recent PreꞒ Ꞓ O S D C P T J K Pg N
Radiolaria illustration from the Challenger expedition 1873–76
Scientific classification
Domain:	Eukaryota
Clade:	Diaphoretickes
Clade:	SAR
Phylum:	Retaria
Subphylum:	Radiolaria Cavalier-Smith, 1987
Classes
Polycystinea Nassellaria Spumellaria Collodaria Acantharea Taxopodida (Sticholonche) Excluded: Phaeodaria (now in Cercozoa)

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The Radiolaria, also called Radiozoa, are unicellular eukaryotes of diameter 0.1–0.2 mm that produce intricate mineral skeletons, typically with a central capsule dividing the cell into the inner and outer portions of endoplasm and ectoplasm. The elaborate mineral skeleton is usually made of silica.^[1] They are found as zooplankton throughout the global ocean. As zooplankton, radiolarians are primarily heterotrophic, but many have photosynthetic endosymbionts and are, therefore, considered mixotrophs. The skeletal remains of some types of radiolarians make up a large part of the cover of the ocean floor as siliceous ooze. Due to their rapid change as species and intricate skeletons, radiolarians represent an important diagnostic fossil found from the Cambrian onwards.

Radiolarians have many needle-like pseudopods 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 vacuoles and lipid droplets, keeping them buoyant. The radiolarian can often contain symbiotic algae, especially zooxanthellae, which provide most of the cell's energy. Some of this organization is found among the heliozoa, but those lack central capsules and only produce simple scales and spines.

Some radiolarians are known for their resemblance to regular polyhedra, such as the icosahedron-shaped Circogonia icosahedra pictured below.

The radiolarians belong to the supergroup Rhizaria together with (amoeboid or flagellate) Cercozoa and (shelled amoeboid) Foraminifera.^[2] Traditionally the radiolarians have been divided into four groups—Acantharea, Nassellaria, Spumellaria and Phaeodarea. Phaeodaria is however now considered to be a Cercozoan.^[3][4] Nassellaria and Spumellaria both produce siliceous skeletons and were therefore grouped together in the group Polycystina. Despite some initial suggestions to the contrary, this is also supported by molecular phylogenies. The Acantharea produce skeletons of strontium sulfate and is closely related to a peculiar genus, Sticholonche (Taxopodida), which lacks an internal skeleton and was for long time considered a heliozoan. The Radiolaria can therefore be divided into two major lineages: Polycystina (Spumellaria + Nassellaria) and Spasmaria (Acantharia + Taxopodida).^[5][6]

There are several higher-order groups that have been detected in molecular analyses of environmental data. Particularly, groups related to Acantharia^[7] and Spumellaria.^[8] 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 support their close relationship—a grouping termed Retaria.^[9] But whether they are sister lineages or whether the Foraminifera should be included within the Radiolaria is not known.

Class	Order	Image	Families	Genera	Species	Description
Polycystinea	Nassellaria					...
	Spumellaria					...
	Collodaria					...
Acantharea						...
Sticholonchea	Taxopodida		1	1	1	...

In the diagram on the right, a Illustrates generalized radiolarian provinces ^[10][11] 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.^[12][13]

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 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).^[13]

• 
  Circogonia icosahedra, radiolarian species shaped like a regular icosahedron
• 
  Anthocyrtium hispidum Haeckel

Radiolarian shapes

          Drawings by Haeckel 1904 (click for details)

Radiolarians are unicellular predatory protists encased in elaborate globular shells (or "capsules"), 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, provide valuable information about past oceanic conditions.^[14]

• 
  Like diatoms, radiolarians come in many shapes
• 
  Also like diatoms, radiolarian shells are usually made of silicate
• 
  However acantharian radiolarians have shells made from strontium sulfate crystals

• 
  Cutaway schematic diagram of a spherical radiolarian shell
• 
  Cladococcus abietinus

Bernard Richards, worked under the supervision of Alan Turing (1912–1954) at Manchester as one of Turing's last students, helping to validate Turing’s theory of morphogenesis.^{[18][19][20][21]}

"Turing was keen to take forward the work that D’Arcy Thompson had published in On Growth and Form in 1917".^[20]

Turing and radiolarian morphology

Shell of a spherical radiolarian

Shell micrographs

Computer simulations of Turing patterns on a sphere
closely replicate some radiolarian shell patterns^[22]

External videos
Radiolarian geometry
Ernst Haeckel's radiolarian engravings

• Spine variations in radiolarians as discovered by HMS Challenger in the 19th century and drawn by Ernst Haeckel
• 
  Cromyatractus tetracelyphus with 2 spines
• 
  Circopus sexfurcus with 6 spines
• 
  Circopurus octahedrus with 6 spines and 8 faces
• 
  Circogonia icosahedra with 12 spines and 20 faces
• 
  Circorrhegma dodecahedra with 20 (incompletely drawn) spines and 12 faces
• 
  Cannocapsa stethoscopium with 20 spines

The gallery shows images of the radiolarians as extracted from drawings made by the German zoologist and polymath Ernst Haeckel in 1887.

• Turing, Alan (1952). "The Chemical Basis of Morphogenesis" (PDF). Philosophical Transactions of the Royal Society of London B. 237 (641): 37–72. Bibcode:1952RSPTB.237...37T. doi:10.1098/rstb.1952.0012. JSTOR 92463. S2CID 120437796.
• Richards, Bernard (2005-2006) "Turing, Richards and Morphogenesis", The Rutherford Journal, Volume 1.

Fossil radiolarian

X-ray microtomography of Triplococcus acanthicus

This is a microfossil from the Middle Ordovician with four nested spheres. The innermost sphere is highlighted red. Each segment is shown at the same scale.^[23]

The earliest known radiolaria date to the very start of the Cambrian period, appearing in the same beds as the first small shelly fauna—they may even be terminal Precambrian in age.^{[24][25][26][27]} They have significant differences from later radiolaria, with a different silica lattice structure and few, if any, spikes on the test.^[26] 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 and determination of ancient climates.^[28]

Some common radiolarian fossils include Actinomma, Heliosphaera and Hexadoridium.

• Radiolarite

Wikimedia Commons has media related to Radiolaria.

Wikispecies has information related to Radiolaria.

• ^[1]Radiolarians
• Brodie, C. (February 2005). "Geometry and Pattern in Nature 3: The holes in radiolarian and diatom tests". Micscape (112). ISSN 1365-070X.
• Radiolaria.org
• Haeckel, Ernst (1862). Die Radiolarien (Rhizopoda radiaria). Berlin. Archived from the original on 2009-06-19. Retrieved 2007-09-07.{{cite book}}: CS1 maint: location missing publisher (link)
• Radiolaria—Droplet
• Tree Of Life—Radiolaria