Field of Science

Fusulinoids: Complex Forams of the Late Palaeozoic

Among the most characteristic fossils of the latter part of the Palaeozoic are the group of Foraminifera known as the fusulinoids. These forams, known from around the middle of the Carboniferous to the end of the Permian, can be extremely abundant. Indeed, I get the impression that some fossil deposits are pretty much made of fusulinoids. Fusulinoids did not merely thrive in their environment; they were the environment.

Limestone block dominated by fusulinids, copyright James St John. Field of view is about 3.9 cm across.

Fusulinoids are distinguished from other forams by their test composition, built from minute granules of calcite, and complex internal structure. Externally, fusulinoids (defined here to exclude their forerunners, the endothyroids) were fairly conservative, with a planispiral, usually involute test (that is, each successive whorl covers the last). The last whorl ended on a transverse wall without a defined aperture; instead, the only connection between the interior and exterior of the test was by a series of pores in said wall. Early forms were disc-shaped; later species could be more globular or fusiform. Some of the later fusulinoids also reached gigantic sizes by single-celled organism standards: whereas the earliest fusulinoids were only a fraction of a millimetre across, the late Permian Polydiexodina could be up to six centimetres along their longest axis (Loeblich & Tappan 1964). Internally, fusulinoids had an incredibly complicated and varied structure which I'm not going to go into too much detail about here, primarily because I barely understand a word of it myself. Any description of fusulinoid morphology quickly devolves into madly throwing about terms like chomata, parachomata, spirotheca, tectorium, and the like, and your humble narrator feeling the need to go look at something else.

Cutaway diagram of a fusulinid, showing an example of internal structure, from here.

I have to go into some detail, though, because some features of the fusulinoid wall structure may explain their success. The ancestral state for the fusulinoid test wall involved a thin layer of solid calcite, the tectum. In most species, the inside of the tectum was coated with a thicker, less dense layer. As the test wall becomes more derived, this inner layer becomes more or less translucent, or pierced by tubular alveoli to produce a honeycomb-like appearance. It has been suggested that these modifications may have been adaptations to accomodating symbiotic microalgae, striking a balance between maintaining the protective test and allowing optimal transmission of light. Microalgal associations with fusulinoids may be corroborated by the discovery of minute fossils of probable planktonic relationships such as Ovummuridae preserved within fusulinoid tests (Vachard et al. 2004).

Ecologically, fusulinoids were restricted to off-shore marine habitats, being mostly found preserved in limestones and calcareous shales. They are absent from deposits that would have been formed in brackish water, and while they may be found in sandstones it is debatable whether such occurrences represent life associations or post-mortem transport (Loeblich & Tappan 1964). Fusulinoids would therefore have been ecologically similar to the inhabitants of modern-day photic zone coral reefs, another reflection of their probable co-dependence with photosynthetic microalgae. However, as successful as the advanced fusulinoids were in their time, they did not make it past the massive extinction event at the end of the Permian. This was not the end of giant and complex forams entirely—indeed, some later forms such as the alveolinids would evolve morphologies very similar to those of fusulinoids—but it was the end of these particular giant forams.


Loeblich, A. R., Jr & H. Tappan. 1964. Treatise on Invertebrate Paleontology pt C. Protista 2. Sarcodina, chiefly "thecamoebians" and Foraminiferida vol. 1. The Geological Society of America and The University of Kansas Press.

Vachard, D., A. Munnecke & T. Servais. 2004. New SEM observations of keriothecal walls: implications for the evolution of Fusulinida. Journal of Foraminiferal Research 34 (3): 232–242.


  1. Did you pick a picture with Cyrillic text specifically to underscore the incomprehensibility of their anatomy?

    As large forams, they presumably had many nuclei, and thus arguably "cheated" on the single-cell front. Can a hard line be drawn between multinucleate unicells and syncytia?

  2. The Cyrillic-text diagram just happened to be the best I could find, but I did make a conscious decision not to attempt to translate the labels.

    As for the number of nuclei, I should note that it turns out I've been wrong in the past in assuming that large forams must have been multinucleate; some surprisingly large ones have just a single nucleus. In the case of fusulinoids, the centre of the test is a more or less globular chamber called the proloculus. This was presumably the embryonic chamber and has generally been presumed to hold the nucleus, though just how much of the proloculus the nucleus would have taken up is of course an unknown. I don't think there's a dividing line between a multinucleate unicell and a syncytium; I think they're just alternate terms for the same thing.

  3. Well, presumably nobody would call a syncytium that's part of a larger multicellular organism (e.g. a human muscle fibre) a "multinucleate unicell", so the terms aren't quite synonymous.

    But Polydiexodina is a candidate 6cm uninucleate? Are there any modern parallels?

  4. Bathysiphon filiformis reaches up to 50 mm in length with a single nucleus (which is large enough to be visible with the naked eye, a bit over half a millimetre across). Mind you, Bathysiphon has a much simpler structure than fusulinoids, being a largely undifferentiated tube. I'm also told by my colleague who shares an office with me and works on foram biostratigraphy that modern nummulitids (which are somewhat comparable to fusulinoids) have only a single nucleus each.


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