Who Knows Which Way the Water Flows?

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Dorsal and lateral views of specimens of Trenella bifrons, from Parkhaev (2001).

There is no question that the molluscs are one of the most significant groups of animals in the marine environment. And thanks to the production by many species of mollusc of a hard shell, they are also one of the best-known groups in the fossil record. A rich and detailed picture of molluscan evolution is available to us as far back as the earliest Cambrian. But, of course, the further back in time we go the more questions we have about what the picture means. And it is in the earliest part of their history that the picture becomes the most opaque.

The Trenellidae are part of that early picture. This family of molluscs is known from the early Cambrian (Parkhaev 2002). They are part of the assemblage of early molluscs referred to as the helcionelloids, whose overall position in the molluscan family tree is very much open to question. Helcionelloids are simple, more or less cap-shaped or cone-shaped shells that are usually also tiny. The type species of the Trenellidae, Trenella bifrons, for instance, is only about 1 to 1.5 millimetres along the longest axis, and only one-half to one millimetre tall (Parkhaev 2001). This all adds up to a general shortage of morphological details that might help us pin down which, if any, modern molluscan groups helcionelloids are connected to. Possession of a undivided dorsal shell has lead many to compare them to gastropods. Others have pointed to the monoplacophorans like the modern Neopilina. In both cases, though, the resemblance is fairly superficial and confirming things one way or another would depend on identifying features of the soft anatomy, such as torsion, that are difficult if not impossible to infer from features of the shell alone.

Within the helcionelloids, trenellids are characterised by having the lower rim of one end of the shell's long axis drawn out into a siphonal groove. It seems likely that this groove was somehow involved in the passage of water around the gill(s), but whether its position indicates the front end or the back end of the shell, and whether it was used to draw water in or expel water out, depends again on what each author expects its original soft anatomy to have been. Unfortunately, evidence for the latter in trenellids is almost completely non-existent; while muscle scars have been identified in some helcionelloids, they remain unknown for this family.

The Trenellidae are closely related to, and probably include the ancestors of, the Yochelcionellidae in which the siphonal groove become raised and closed ventrally, turning it into a snorkel-like structure (one yochelcionellid, Yochelcionella daleki, has been featured on this site before). However, comparing trenellids to yochelcionellids raises something of a question in my mind. In general, mollusc shells grow through secretion from the mantle around the shell's rim only, meaning that once shell growth has passed a certain section the mollusc usually cannot go back and rearrange it. Assuming that helcionelloids grew in the usual molluscan manner, surely yochelcionellids would have gone through a stage in their development before the lower part of the 'snorkel' was closed off where they looked a heck of a lot like a trenellid? Is it even possible to distinguish a mature trenellid from a juvenile yochelcionellid?

REFERENCES

Parkhaev, P. Yu. 2001. Trenella bifrons: a new helcionelloid mollusk from the Lower Cambrian of South Australia. Paleontological Journal 35 (6): 585–588.

Parkhaev, P. Yu. 2002. Phylogenesis and the system of the Cambrian univalved mollusks. Paleontological Journal 36 (1): 25–36.