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One of the defining characteristics of the Phylum Mollusca is the possession of a shell, which serves both as a protective covering and an exoskeleton. We've all seen snails, and some people may have noticed that snails often withdraw entirely into their shells and even have a little door that they can use to seal up the opening of the shell. That little door is called the operculum. Opercula occur in non-molluscan animals, too, such as some of the tube-dwelling polychaete worms and some of the thecate hydroids. Snail opercula come in lots of different shapes, depending on the aperture of their owner's shell.

Calliostoma ligatum at Mitchell's Cove
2 April 2017
© Allison J. Gong

Given the enormous morphological diversity within the Mollusca it shouldn't be surprising that their shells vary immensely in prominence and shape. In fact, molluscan shells demonstrate quite beautifully the relationship between form and function. The benthic and most familiar molluscs, the gastropod snails, generally have coiled shells. Notable exceptions to this generality are the marine opisthobranchs (nudibranchs and sea hares) and the terrestrial slugs. And for the most part snail shells look recognizably like snail shells, even though some are plain coils, others may be flattened (e.g., abalones), and still others may be crazily ornamented. Aquatic animals crawl around in water, which helps to support the weight of heavily calcified shells. Terrestrial snails, on the other hand, live in a much less dense medium (air) and have lighter, less calcified shells. The trade-off for a more easily transportable shell is that air is also very drying, and a thinner shell provides less protection from desiccation.

I should state for the record right now that I'm not talking about the many molluscs that don't have shells at all, or that have much reduced shells.

View into the exhalant opening of a mussel (Mytilus sp.) at the Santa Cruz Yacht Harbor
29 August 2015
© Allison J. Gong

The bivalve molluscs (mussels, clams, oysters, etc.) live inside a pair of shells. They are sedentary animals, living either attached to a hard surface or buried in sand or mud. Not being able to run from predators (although some scallops can swim!), their only defense is the toughness of their shells and the strength of the adductor muscles that hold the shells closed. Most bivalves feed by sucking water into the shells through an incurrent siphon, using their gills to filter food particles from the water, and expelling the water through an excurrent siphon. To do so they must open their shells enough to extend their siphons, or at least expose inhalant and exhalant openings, to the water current surrounding them.

So, snails have one shell and bivalves have two. Some of the most interesting molluscs, in terms of shell morphology, are the chitons. The Polyplacophora (Gk: 'many plate bearer') have a shell that is divided into eight dorsal plates. This makes them immediately distinguishable from just about any other animal.

Chiton (Lepidozona mertensii) at Point Piños
6 February 2016
© Allison J. Gong

Chitons live from shallow water to the deep sea, but the majority of species live in the intertidal. This is a high-energy habit characterized by the bashing of waves as the tide rises and falls twice daily. Any organism living here must be able to hang on for dear life or risk being swept away to certain death. Chitons are certainly well equipped to survive in this habitat. They have a low profile, offering minimal resistance to the waves. Rather than stand tall and face the brunt of the wave energy, chitons cling tightly to the rocks and let the waves wash over them.

Tonicella lokii at Monastery Beach
27 October 2015
© Allison J. Gong

The chiton's shell, divided into eight articulating plates, gives the animal a much more flexible shell than is found in any other mollusc. This allows them to conform to the topography of the rocks, giving them an even lower profile than, say, a limpet of the same overall shape and size.

While most chitons are pretty sedentary, at least during the low tides when we can see them, some of them can move pretty quickly when they want. So what, exactly, motivates a chiton to run? One species, Stenoplax heathiana, lives on the underside of rocks in the intertidal; it comes out at night to forage on algal films and retreats back under its rock with the dawn. I've seen them at Pistachio Beach, where I turned over rocks and watched them run away from the light. This video is shot in real-time; the chitons are really running fast!

When the eight shell plates are visible it's easy to identify a chiton as a chiton. But not all chitons are quite so obliging with their most chiton-ish characteristic, and one is downright misleading.

Below is Katharina tunicata, one of the largest chitons on our coast. Its shell plates are barely visible, as they are almost entirely covered by the animal's mantle, the layer of tissue that covers the visceral mass and encloses an open space called the mantle cavity in which the gills are located. In chitons, the mantle extends onto the dorsal side of the animal and is called the girdle. Katharina's girdle is smooth and feels like wet leather.

The chiton Katharina tunicata at the Great Tidepool in Pacific Grove
26 October 2015
© Allison J. Gong

The largest chiton in the world is the gumboot chiton, Cryptochiton stelleri, and it lives on our coast. This beast is about the size of a football, reaching a length of 30 cm or so. It lives mostly in subtidal kelp forests, but can be found in the very low intertidal, which is where I usually see it. At first encounter it's hard to figure out what this animal is. It certainly doesn't look like a chiton.

A large Cryptochiton stelleri at Mitchell's Cove
6 June 2016
© Allison J. Gong

If anything, it looks like a mostly deflated football, doesn't it? Turning it over to look at the underside doesn't help much, either, although this photo does give an idea of how big the animal can get:

Ventral view of Cryptochiton stelleri at Pigeon Point
24 April 2016
© Allison J. Gong

Cryptochiton goes one beyond Katharina and covers its plates entirely. Just looking at the animal you'd have no idea that there are eight plates underneath the tough reddish-brown mantle, but you can feel them if you run your finger along the midline of the dorsum. Living subtidally as it does, Cryptochiton doesn't have the ability to cling tightly to rocks that its intertidal relatives do, and it tends to get washed off its substrate and cast onto the beach during storms. I've never seen one on the beach that wasn't very dead. Once a friend and I were trudging back up the beach after working a low tide, and encountered a dead softball-sized Cryptochiton. I mentioned that it would be nice to have a complete set of shell plates from one of these animals. My friend always carries a knife in her pocket, so we started an impromptu dissection right there on the beach. It didn't take long to learn that the mantle of a gumboot chiton is really tough and difficult to cut through with a pocket knife. And even once we got through the mantle, dissecting the plates from the underlying tissue wasn't going to happen with the tools we had with us. Besides, the stench was godawful even with our unusual tolerance for the smell of dead sea things. We abandoned that corpse.

Single plate of the gumboot chiton, Cryptochiton stelleri
16 May 2017
© Allison J. Gong

Many beachcombers have found white butterfly-shaped objects in the sand, but not known what they are. They are definitely calcareous and feel like bone, but what kind of animal makes a bone shaped like this? Turns out this object is one of the shell plates from C. stelleri. They wash up frequently, never attached to their neighbors so they provide no clue as to what organism they came from.

In order to obtain a complete set of Cryptochiton plates, I'd have to start with an intact chiton corpse. I did happen upon another dead Cryptochiton on a beach somewhere I was allowed to collect organisms, and I brought it back to the marine lab. I remember spending a smelly afternoon cutting the plates out of the corpse and removing as much of the tissue as I could, then feeding the plates to various hungry anemones to take care of the rest. Some of the plates got a little broken during the extraction process, but I do have my very own full set!

Shell plates of Cryptochiton stelleri
16 May 2017
© Allison J. Gong

Some day I will figure out a way to mount those plates permanently.

One final question to ponder. Does a chiton have one shell, or eight shells?

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