Subject: Geology

Institute: Science
Class: M. Sc. Semester II
Paper: GLM-204 (Micropaleontology and Oceanography)
Topic: Radiolarians (Siliceous Microfossils)

Dr. Dinesh Kumar Naik

Assistant Professor
Department of Geology
Institute of Science
Banaras Hindu University
Varanasi 221 005
Email: dnaik.geo@bhu.ac.in
 Siliceous Microfossils

Radiolaria:
Radiolarians are single-celled, marine zooplankton. They belong to the Phylum Radiozoa.
These are pelagic marine mostly unicellular protozoans having a siliceous skeleton with
no external hard part. They range in age from Pre-Cambrian to Recent.
They can be found floating near the surface or at water depths of hundreds of metres.
The size of radiolarian varies from 30 μm to 2 mm in diameter and some of them form
macroscopic colonies that may reach a size of a centimetre or larger.
Morphology:
The living matter of the radiolarian cell comprises two
layers:
1. a mucoid or chitinous sac in the centre, called the
central capsule, holding the intracapsular cytoplasm and
nucleus, and
2. the outer layer of cytoplasm surrounding the central
capsule, known as the extracapsulum.

Thread-like axopodia radiate from the central capsule and

through the extracapsulum.

The intracapsular cytoplasm consists of the nucleus,

vacuoles of varying sizes and the vital organelles such as
mitochondria and Golgi bodies.

The extracapsulum cytoplasm contains frothy gelatinous

bubbles called calymma and algal symbionts.

Some of the radiolarians emit a bluish flash of light

(bioluminescence) to deter predators.

The life cycle of some shallow water radiolarians is

suggested to be 1–3 months.

Binary fission is observed in some species, but it is not

established if they reproduce sexually.
The soft parts of radiolarians are collectively known
as malacoma.

The skeleton of a radiolarian, known as a

scleracoma, is made of amorphous, opaline silica.

An amazing diversity of skeletal structures is found

in the living and fossil radiolarians.
The skeleton is enclosed within a cytoplasmic
sheath called a cytokalymma and not in direct
contact with the seawater.

The skeleton consists of a porous lattice shell of

variable shapes from spherical to spindle and
conical. There may be concentric or overlapping
lattice shells.

The radial elements are hollow to solid spines

(attached at one end only), bars (attached at both
ends to other elements) and simple spicules.
There are three well-recognized divisions of radiolarian:
1. Spumellaria
2. Nassellaria
3. Phaeodaria

The orders Spumellaria and Nassellaria are grouped under the Class Polycystinea,
and Phaeodaria forms a distinct class.

Anderson (1983) discusses the distinguishing morphological characteristics of these types as follows:
The skeletons of Spumellaria are characterized by radial symmetry and comprise
(1) simple, needle-shaped spicules to complex, symmetrically arranged, triradiate spines distributed in the
extracapsulum or clustered around the central capsule,
(2) spheroidal to spherical shells that are either single or multiple concentric, enclosing the central capsule, and
(3) complex polyhedral skeletons resembling lattices or geodesic structures, reinforced in some groups by radial
beams. In the skeletons with concentric shells, the innermost layer is the medullary shell and the outermost
shells are cortical shells. The connections between shells and structures are through bars and beams.

Spumellarian radiolarians
characterized by spherical to ellipsoidal shape and perforate wall (reproduced after Kling 1998, with permission © Elsevier)
The Nassellaria are characterized by axial symmetry and have three types of skeletons:

1. a tripod located near the base of the central capsule and characterized by three divergent bar-
like elements united at a common central point

2. conical or hat-shaped, complexly perforated shells

3. a sagittal ring that reinforces the latticed shell in the medial, sagittal plane. The skeletons of
Nassellaria are often very complex and multi-chambered, differentiated into cephalis, thorax and
abdomen.

Nassellarian radiolarians
characterized by cap-shaped skeletons, small spherical cephalis and one or more post-cephalic chambers
(reproduced after Kling 1998, with permission © Elsevier)
The phaeodarian structures are delicate and formed of 95 % organic matter and 5 %
silica, and, therefore, have poor fossil records.

A large, oblate spheroid, depressed in the direction of the main axis, distinguishes the
Phaeodaria from the other two types.

The skeletons of some Phaeodaria comprise hollow tubes containing living cytoplasm
and organic materials where tubes are joined to one another.

Such fragile skeletons get disarticulated before being buried under the sediments.

In living forms, the Phaedorians are characterized by distinctive and complex

architectures.
 Environmental Significance

Temperature, salinity and nutrient characteristics of the water masses control the
distribution of radiolarian assemblages. The radiolarians, thus, preserve the signatures
of oceanic and climatic changes of the past.

Unlike calcareous microfossils, the radiolarians are preserved in deep-sea sediments

independent of the CCD and have greater potentiality to be used as tools in
biostratigraphy and paleoceanography, especially where calcareous microfossils are
absent due to their dissolution.

The radiolarians range from the Cambrian period, but the skeleton-less forms may have
evolved in the Precambrian.

They dominate the sediments below the carbonate compensation depth and form
radiolarian oozes.
Such oozes are mostly found in the equatorial Pacific below the zones of high
productivity at depths of 3–4 km, but abundant radiolarians also occur at shallower
depths associated with coccolith or planktic foraminiferal oozes.
The radiolarians are exclusively marine.
The alveolar complex containing CO2-saturated water is suggested to be an adaptation for the planktic mode of
life. The vertical movement of radiolarians in seawater is supposed to be facilitated by adjustment in volume of
the CO2 in alveoli. The skeletal structures, such as perforated walls, radiating spines and axopods, are further
adapted for enhancing buoyancy in the water column. Many radiolarians host dinoflagellates as symbionts and,
therefore, dominate the photic zone (<200 m). Some of the radiolarians form colonies of spherical to cylindrical
shapes, centimetres to metres in dimension.

The radiolarians are abundantly present in equatorial latitudes, but they also occur in subpolar seas.
There are distinct assemblages corresponding to different ocean circulation and water mass characteristics.
Seven biogeographic faunal zones are distinguished in the Pacific (Casey 1971). Although several of the species
may occur in more than one faunal zone, a simplified distribution of the most abundant species in selected zones
is listed below:

Subarctic Transition Fauna: high latitude, north of Arctic or Polar Convergence; Spongotrochus
glacialis, Sethophormis rotula, Pterocanium sp.

Central Shallow Fauna: intermediate latitude; Calocyclas amicae, Euchitonia furcata, Eucyrtidium
hertwigii.

Equatorial Fauna: low latitude, bounded by North and South Equatorial Current Systems;
Acrosphaera murrayana, Acrobotrissa cribrosa, Anthocyrtidium cineraria, Lithomelissa monoceras,
Tristylospyris scaphipes.
The statistical analysis of the present-day distribution of radiolarians in
the southern hemisphere provides the following transfer functions on the
basis of factor loadings for the subtropical (A), Antarctic (B) and sub-
Antarctic (C) assemblages:

Tw = 15.266 A +1.542B+ 9.984C -1.984,

Ts = 13.966A - 2.904B+10.545C + 5.103,

where Tw and Ts refer to temperatures in winter (August) and summer

(February), respectively.
The error in both functions is less than 10 %, which is a fairly good
estimate (refer to Lozano and Hays 1976 and Hays et al 1976 for further
details).
The transfer function can estimate Quaternary seawater temperatures
with a fair degree of accuracy.
The radiolarian distributions are also related to upwelling. Radiolarian-
based indices are proposed for estimating paleo-upwelling.

The Upwelling Radiolarian Index (URI), based on assemblages in the

eastern equatorial Pacific, considered 14 upwelling species, including:
Acrosphaera murrayana,
Eucyrtidium erythromystax,
Lamprocyrtis nigriniae and
Pterocorys minythorax (Haslett 2003).
 Biostratigraphy use:
The fossil record of radiolarians ranges from the Cambrian Era, but it is likely that the
skeleton-less forms would have evolved in the Precambrian.

Spumellaria dominated in its early history in the Paleozoic,

but Nassellaria began to diversify after the Carboniferous.

After the Permian–Triassic boundary extinction, the two groups have expanded since the
Jurassic.
The Mesozoic was the main era of radiolarian radiation. The cause of the radiation is not
certain, but one of the reasons was the breakup of the continents and partitioning of the
world’s oceans that strengthened ocean circulation and the upwelling of nutrient
(D’Wever et al. 2003).

Because of their high morphological diversity, radiolarians are good tracers of

evolutionary history.
An intensive study of evolutionary trends of Cretaceous radiolarians reveals that
Spumellaria and Nassellaria respond differently to ecological stress.
The spumellarians are much more extinction resistant during episodes of critical
environmental deterioration, but nassellarians possess a relatively higher evolutionary
rate than spumellarians during periods of radiation (O’Dogherty and Guex 2002).
The diversity of radiolarians was stable for the greater part of the Cenozoic, but a distinct
change in the robustness of the test occurred after the Eocene.

The Oligocene was marked by the demise of many thickly silicified Paleogene
radiolarians.
The average weight of the radiolarian test has decreased four times since the Eocene
epoch (Racki and Cordey 2000).

Variations in temperature, nutrient and intensity of light in the water column are the
significant factors in the evolution of radiolarians and other plankton.

Competition with the co-evolving group also played a major role.

The diminishing weight of the test is attributed to competitive pressure for dissolved silica
in seawater triggered by the explosive radiation of other siliceous plankton, including
diatoms and silicoflagellates.
The radiolarians adapted by decreasing the wall’s thickness, increasing pore size and
decreasing the width of the bars.
A visible expansion in diversity occurred in the Quaternary and the nassellarians, in
particular, diversified more in this period.
There is a fair similarity between the diversity trends of radiolarians and dinoflagellates,
possibly indicating a trophic relationship or symbiotic interdependence (Anderson 1983).