Silicon Rockets

In a previous post, I spoke of the radiolarians, marine protists renowned for their intricate skeletons, and the major radiolarian group known as the Spumellaria. Standing in contrast to the spumellarians is another major group, the Nassellaria. Like spumellarians, nassellarians have a skeleton of silica but whereas the basic shape of spumellarian skeleton is a sphere, that of nassellarians is a cone, bell or some similar shape, arranged along a longitudinal axis. The origination point of the skeleton is at or near the top of the cone and is known as the cephalis (from the Greek for 'head'). There may be an apical spine rising above the cephalis. Below it, the skeleton is commonly divided into recognisable sections referred to as the thorax, abdomen and post-abdominal segments (if present). The nucleus of the cell is more or less associated with the cephalis, contained within it at least during the juvenile stage of development though it may shift below the cephalis as the cell matures (Suzuki et al. 2009).

Skeleton of a Eucyrtidium sp., copyright Picturepest.

As is commonly the case with unicellular organisms, radiolarian taxonomy has been influenced by disagreements about which features should be regarded as more significant. Some would arrange taxa based on the overal formation of the skeleton. Others would focus on the development of the initial embryonic spicule around which the cephalis develops. A recent phylogenetic analysis of living nassellarians by Sandin et al. (2019), based on both morphological and molecular data, found that overall skeleton morphology was a much better indication of relationships than the internal structure. One well supported subgroup of the Nassellaria is the superfamily Eucyrtidioidea.

Eucyrtidioids have a fossil record going back to the Triassic (Afanasieva et al. 2005). The cephalis is spherical and clearly distinguished from the following segments by a constricted basal aperture. The test is usually multi-segmented; members of the subfamily Theocotylinae may have just two segments but other members of Eucyrtidiidae have up to ten segments. Fossil families assigned to Eucyrtidioidea by Afanasieva et al. (2005) may have up to twenty (but as Afanasieva et al.'s concept of Eucyrtidioidea was not found to be monophyletic by Sandin et al., the affinities of these fossil families perhaps warrant re-investigation). Segments are commonly divided by distinct inner rings. The skeleton lacks feet, the term used for protruding spines around the basal aperture of the skeleton found in many other nassellarians.

The phylogeny of nassellarians indicated by Sandin et al. (2019) places the Eucyrtidiidae as the sister taxon to other living nassellarians. Other living families included in the Eucyrtidioidea by Afanasieva et al. (2005) were placed in more nested positions. The implication is that the multi-segmented condition may be ancestral for crown Nassellaria. Segments are added progressively during the life of the radiolarian, leading the organism to look quite different at different ages. Indeed, this metamorphosis is pronounced enough that one of the earliest influential researchers on radiolarians, Ernst Haeckel (he of Kunstformen der Natur fame), made the mistake of classifying different ages as different species, genera and even families. Our understanding may be better than in Haeckel's time but there may still be a lot to learn about these intricate organisms.

REFERENCES

Afanasieva, M. S., E. O. Amon, Y. V. Agarkov & D. S. Boltovskoy. 2005. Radiolarians in the geological record. Paleontological Journal 39 (Suppl. 3): S135–S392.

Sandin, M. M., L. Pillet, T. Biard, C. Poirier, E. Bigeard, S. Romac, N. Suzuki & F. Not. 2019. Time calibrated morpho-molecular classification of Nassellaria (Radiolaria). Protist 170: 187–208.

Suzuki, N., K. Ogane, Y. Aita, M. Kato, S. Sakai, T. Kurihara, A. Matsuoka, S. Ohtsuka, A. Go, K. Nakaguchi, S. Yamaguchi, T. Takahashi & A. Tuji. 2009. Distribution patterns of the radiolarian nuclei and symbionts using DAPI-fluorescence. Bulletin of the National Museum of Nature and Science, Series B 35 (4): 169–182.