Radiolaria Polycystina
Introduction

Polycystine radiolarians are exclusively marine, pelagic, solitary or colonial protists provided with actinopods. Polycystines comprise the Order Collodaria, a small group lacking a skeleton (Actissa princeps ov; Collozoum inerme; Collozoum inerme 2; Collozoum serprentinum; Collozoum longiforme; Collozoum sp.; Rhaphidozoum sp.), or provided only with scattered mono- or polyaxonic spicules (Thalassoxanthium medusinum ov; Thalassoxanthium cervicorne ov; Lampoxanthium pandora ov; Rhaphidozoum neapolitanum; Sphaerozoum geminatum); and the Order Spumellaria and Order Nassellaria, most of which have a well developed siliceous latticed or spongy skeleton (see Superorder Polycystina forthe outline classification).

Solitary [l][m]Glossary[/m][r]species[/r]species (the greatest majority) range between 20-30 µm to about 300 µm, but colonies (some Collodaria and the spumellarian family Collosphaeridae) may in exceptional cases be as long as 3 m (Swanberg, 1979). The siliceous skeletons of the polycystines are a major contributor to the sedimentary flux, their earliest records dating back to the Cambrian. Paleozoic, Mesozoic and Cenozoic sequences furnish detailed records for evolutionary, stratigraphic and paleoecologic analyses.

A distinguishing feature of all radiolarians (polycystines and phaeodarians) is the central capsule membrane, a proteinaceous perforated membrane that divides the cytoplasm into two areas: the endoplasm or intracapsular cytoplasm, and the calymma or extracapsular cytoplasm (Spumellaria). This central capsule is either spherical (in many of Order Spumellaria), or elongated and pyriform (in most of Order Nassellaria: Nassellaria). The intracapsular cytoplasm contains reserve substances and major cytoplasmic organelles (nucleus or nuclei, mitochondria, and other organelles, except for the digestive vacuoles), and is generally believed to be responsible for the functions of reproduction, biochemical synthesis and energy production. The calymma is the frothy or web-like extracapsular cytoplasm where the digestive vacuoles are located. Algal symbionts, when present, are enclosed within vacuoles usually located in the calymma. Colonial forms have a gelatinous sheath containing numerous central capsules interconnected by a rhizopodial network (Collozoum inerme; Collozoum inerme 2; Collozoum serprentinum; Collozoum longiforme; Collozoum sp.; Rhaphidozoum neapolitanum; Collozoum sp.; Rhaphidozoum sp.).

Polycystine skeletons are typically constructed of a network of structures which can be either connected at both ends with other elements - the bars (nodes/bars intro), or formations attached to the rest of the shell by one end only - the spines (Stylatractus spp. group; Cromyechinus icosacanthus intro; types of spines). All skeletal elements are composed of amorphous silica (SiO2 nH2O). There is a perplexing variety of shapes in which these bars and spines can be arranged in order to form the skeleton, from simple latticed spheres (Collosphaera huxleyi) or a few anastomosed spines (Tetraplecta pinigera), to elaborate constructions with several concentric spheres (Cromyechinus icosacanthus intro) or multilocular conical structures with protruding latticed or solid appendages known as wings, feet, teeth, etc.

Illustrations of characters used for the identification of nassellarian radiolarians:

main spines/skeleton
main spines/skeleton 2
main spines/skeleton 3
types of cephalis
type of cephalis 2
type of cephalis 3
undivided cephalis
undivided cephalis 2
trilocular cephalis
multilocular cephalis
shell structure
shell structure 2
shell structure 3
shell structure 4
pore arrangement
pore arrangement 2
A. denticulata group? intro

Very little is known about the reproduction of the Radiolaria. In addition to vegetative reproduction (Hollande and Enjumet, 1953), including binary fission at various ontogeneting stages (Anderson and Swanberg, 1981; Anderson and Gupta, 1998), the production of biflagellated swarmers was observed, but it is not known if the swarmers are asexual dissemules or motile gametes (Anderson, 1983a). Although no direct estimates have been made so far, it is generally assumed that individual radiolarian life spans are around 2-4 weeks (Anderson 1983a; Caron and Swanberg, 1990).

Polycystines consume a wide variety of prey including bacteria, algae, protists, copepods, appendicularians, and other small zooplankton (Anderson 1983a, 1993; Caron and Swanberg, 1990). Algal symbionts, when present, secrete photosynthetic products that are assimilated by the host as a nutritional source (Anderson 1983b).

The first published descriptions of Radiolaria date back to the early XIX century. Between approximately 1850 and 1900, C.G. Ehrenberg, J. Müller, R. Hertwig, A. Popofsky, and especially E. Haeckel described thousands of new species and provided the first comprehensive classification systems (Riedel, 1967a). After a period of little activity, interest in the Radiolaria was renewed around 1950, and somewhat later further fostered by the rich sedimentary materials recovered by the Deep Sea Drilling Project.

Because of their application to stratigraphy, polycystine studies have traditionally been within the realm of geologists/paleontologists, with biologically-oriented publications representing less than 10% of the overall total produced to date (A. Sanfilippo, pers. comm., 1997). The directory included in the 1994 issue of Radiolaria (Newsletter for the International Association of Radiolarian Paleontologists) lists 400-plus names; however, only 100-150 of these are primarily concerned with radiolarian studies. Almost all these workers are geologists focusing their interest on stratigraphic and paleoceanographic problems, especially dealing with Paleozoic and Mesozoic deposits; interest in Cenozoic faunas has been dwindling over the last few years. Biologically-oriented research based on samples from the water-column has even fewer specialists, and at present they are probably less than 10-20 world-wide. Since 1834 approximately 3500 works on polycystine radiolarians have been published (over half of these on Cenozoic faunas, about 35% on Mesozoic, and 15% on Paleozoic; A. Sanfilippo, pers. comm., 1997).