Wooster’s Fossils of the Week: The mysterious Paleozoic encrusters Ascodictyon and Allonema

The above pair of fossils are small sclerobionts commonly found on hard substrates in shallow marine sediments through much of the Paleozoic, especially the Silurian and Devonian. Paul Taylor and I have been studying them for a few years now and our first paper on them was published this summer (Wilson and Taylor, 2014). Ascodictyon (Silurian-Carboniferous) is on the left and Allonema (Silurian-Permian) is on the right. Both are calcitic encrusters and look, at least in this view, very different from each other. We present evidence in our paper, though, that strongly suggests Ascodictyon and Allonema are actually manifestations of the same organism. What that organism is, exactly, still eludes us. We are persuaded at the very least that they are not bryozoans as originally described by Nicholson, Ulrich and Bassler. Since they are so common their identity is important for studies of fossil diversity and paleoecology.
The above view through a light microscope of Ascodictyon and Allonema shows the perspective paleontologists have had of these encrusters until recently. The clear calcite skeletons sitting on a calcitic brachiopod shell (this is from the Devonian of Michigan) makes for little contrast and poor resolution, and the microscope-camera combination has a very limited depth of field. The rest of the images in this post were made with a Scanning Electron Microscope (SEM) expertly operated by Paul. The difference in morphological detail is not just astonishing, it is a revolution in the study of tiny fossils like this.
This is a typical view of Ascodictyon. It consists of stellate clusters of inflated vesicles (like little calcite balloons) connected by thin calcitic tubes called stolons. (Ascodictyon siluriense from the Silurian of the England.)

This is a typical Allonema. The primary form is a series of porous vesicles attached in chains like sausages. (Allonema waldronense from the Silurian of Gotland, Sweden.)

Here is where these obscure little encrusters get interesting. This is a specimen from the Silica Shale (Middle Devonian) exposed in Michigan. It was collected in a beautiful suite of fossils by that intrepid citizen scientist, Brian Bade. It consists of Allonema sausages connected to Ascodictyon stolons which are themselves connected to Ascodictyon stellate vesicle clusters. Clear evidence that Allonema and Ascodictyon are end members of a morphological continuum produced by the same organism.

A critical feature we see in this Ascodictyon/Allonema complex is the occurrence of “sockets” at the bases of vesicles like the above from the Silica Shale. These are almost certainly places where some erect portion of the organism extended above the substrate. Maybe these were feeding devices? Reproductive parts? We’ve found no trace of them.

Our hypothesis is that Allonema (left) and Ascodictyon (right, both from the Silurian of Gotland, Sweden) are the basal parts of some as yet unknown erect organism. They may have stored nutrients for the creature. We are convinced they were not bryozoans, foraminiferans, corals or sponges. Unfortunately we can only classify them as incertae sedis or Microproblematica. At some point we’ll have to figure out how to name this complex with two genera and over a dozen species.

It was fun work, and the project continues. For more detail, see Wilson and Taylor (2014).

References:

Nicholson H.A. and Etheridge R. 1877. On Ascodictyon, a new provisional and anomalous genus of Palæozoic fossils. J. Nat. Hist., Series 4, 19: 463-468.

Ulrich E.O. and Bassler R.S. 1904. A revision of the Paleozoic Bryozoa. Smith. Misc. Coll. (Quart.) 45: 256-294.

Wilson M.A. and Taylor P.D. 2001. “Pseudobryozoans” and the problem of encruster diversity in the Paleozoic. PaleoBios 21 (Supplement to No. 2): 134-135.

Wilson, M.A. and Taylor, P.D. 2014. The morphology and affinities of Allonema and Ascodictyon, two abundant Palaeozoic encrusters commonly misattributed to the ctenostome bryozoans. In: Rosso, A., Wyse Jackson, P.N. and Porter, J. (eds.), Bryozoan Studies 2013. Studi trentini di scienze naturali 94: 259-266.