Mollusca

The phylum Mollusca contains some of the most familiar invertebrates, including snails, slugs, clams, mussels, and octopuses. In contrast to these well-known molluscs, however, others are almost never seen, such as the aplacophorans and monoplacophorans, the latter of which which were only known from Paleozoic fossils until the first live specimen was discovered in the deep sea in 1952 (UCMP 2008).

Except for the aplacophorans, most molluscs have a well-developed, muscular foot. This structure is used in a multitude of ways, for example: locomotion, clinging to surfaces, burrowing, anchoring in sediment, swimming, and grasping (modified into prehensile tentacles in octopuses). The vast diversity of foot adaptations exemplifies the huge morphological diversity of the mollusc form.

A layer of epidermal tissue called the mantle surrounds the body of molluscs. Specialized glands in the mantle are responsible for the extracellular excretions that form shell structures. In all molluscan groups the shell is produced in layers of (usually) calcium carbonate, either in calcite or aragonite form.

Molluscs have adapted to terrestrial, marine and freshwater habitats all over the globe, although most molluscs are marine. Nearly 100,000 mollusc species are known (excluding the large number of extinct species known only as fossils) and it is clear that many thousands of species of extant species remain undescribed. Around 80% of known molluscs are gastropods (snails and slugs).

Most molluscs undergo spiral cleavage. Development can be direct (proceed right to settling into a juvenile form) or indirect, going through the swimming trochophore larval stage. The trochophore is very similar to the annelid trochophore. Before settling, many groups then go onto a second larval stage which is unique to molluscs: the feeding (usually) and swimming veliger larvae. Molluscs go through the uniquely molluscan process of torsion, usually during the veliger stage of development. Torsion involves counterclockwise rotation of the visceral mass up to 180 degrees with respect to the head and foot, to profoundly change the relative location of the body regions. Many groups then “detort” to some degree later in development or adulthood. Theories as to the evolutionary significance of torsion abound but this phenomenon is not well understood (Brusca and Brusca 2003). In the long run, torsion has allowed for much morphological diversification over the course of evolution.

Molluscs have adapted to terrestrial, marine and freshwater habitats all over the globe, although most molluscs are marine.

Despite the highly diverse forms of the members of this taxon, molluscs share a recognizable and characteristic generalized general body plan, made up of a head, a foot, and viscera contained in a central body. They are generally considered unsegmented, although primitive forms (aplacophorans and polyplacophorans) with repeated body features show intriguing potential for a possibly segmented mollusc-annelid ancestor (e.g. Jacobs et al 2000).

The mollusc head can house various combinations of sensory structures: tentacles, photoreceptors, statocysts, chemoreceptors. In some molluscs these sensory systems can be very well developed (the complex cephalopod eye is a prime example). Also found on the head is a feature unique to molluscs: the radula. Found in the buccal (mouth) cavity, the radula usually exists as a tongue-like plate covered with “teeth” used by herbivores, carnivores and scavengers to scrape food particles into the mouth. Depending on diet and use, tooth number, shape, arrangement, makeup, and growth have adapted diversely. Especially in the gastropods, number and shape of radular teeth are important taxonomic characters. The radula has also been adapted for diverse feeding methods. Some gastropods and cephalopods have a drill-like radula used to bore holes in the shell of prey, sometimes with the aid of acids secreted from an adjacent boring gland. In cone snails the radula is set on the end of a retractable proboscis and is slung out like a harpoon, to inject toxins into the prey, delivered through piercing, hollow teeth. In some cases these toxins are powerful neurotoxins, deathly to humans. Several lineages of molluscs have evolved suspension feeding, especially in the gastropods and bivalves. The radula in these cases is either highly reduced or lost altogether, and in most cases food particles are caught by ctinidia (gills) and moved to the mouth by cilia.

Except for the aplacophorans, most molluscs have a well-developed, muscular foot. This structure is used in a multitude of ways, for example: locomotion, clinging to surfaces, burrowing, anchoring in sediment, swimming, modified into prehensile tentacles (octopus); the vast diversity of foot adaptations exemplifies the huge morphological diversity of the mollusc form.

A layer of epidermal tissue called the mantle surrounds the body of molluscs. Specialized glands in the mantle are responsible for the extracellular excretions that form shell structures. The ancestral mollusc is thought to have one shell capped over the body like a limpet, and from that a diverse number of shell arrangements have evolved. Molluscs may have have one, two, or eight (in chitons) shells. Aplacophorans have no shell, but have instead minute aragonite spicules imbedded within the mantle. Secondary loss or much reduced shell vestiges have also occurred independently in multiple mollusc lineages (for example nudibranchs, slugs, cephalopods). Shells usually provide external protection, but there have been several independent internalizations within cephalopods and opisthobranchia. In all molluscan groups the shell is produced in layers of (usually) calcium carbonate, either in calcite or aragonite form. The wide range of pigmentation, shape, size, sculpturing, and twisting of sea shells is, of course, well known. There is much recent developmental work describing gene expression in shell formation, and the roles of highly conserved regulatory genes such as engrailed and Hox genes have been examined (e.g. Jacobs et al 2000, Samadi and Steiner 2009).

Between the mantle and the body proper is the mantle cavity, which may be organized as one or two separate spaces or grooves. Many important functions occur in the mantle cavity: the ctenidia (gills) are positioned here and the body systems, namely the nephridia (kidney like organs), the gut and the reproductive organs open up into this space. In aquatic molluscs cilia on the surface of the mantle and organs maintain water flow through the mantle cavity to take away wastes and bring in oxygenated water (and food particles for those suspension feeding molluscs). Molluscs have an open circulatory system with a full heart (with the exception of the cephalopods, which have a closed circulatory system). Their nervous system is well developed, usually consisting of a dorsal ganglion, a ring of nerves around the esophagus, and two pairs of lateral nerve cords running the length of the body, which are connected transversely in a ladder-like arrangement. There is an enormous range of nervous system development in the molluscs, from the poorly developed ganglia of the aplacophorans to the extreme cephalization of the cephalopods. Important work in the fields of neurobiology has been carried out on the squid Doryteuthis pealeii (formerly Loligo pealeii) and on Aplysia sea slugs.

Like other systems, reproduction is highly variable among molluscs. Molluscs can be dioecious or simultaneously or sequentially hermaphroditic. Gametes are freely spawned in some groups, others have internal fertilization and complex mating behaviors, many produce egg capsules, egg cases, or brood chambers.

Molluscs range in size from almost microscopic to animals 20 meters long (giant squid) or weighing 450 pounds (giant clams).

The molluscs demonstrate remarkable morphological diversity, a characteristic that has confused molluscan taxonomy from the group’s inception. The Latin root molluscus means soft, and many soft-bodied invertebrates have been added and removed this group until Cuvier’s modern approximation in 1795 (Brusca and Brusca 2003). Mollusca is the second largest invertebrate phylum after the arthropods. Some 93,000 extant species have been described, but the thinking is this number represents only about half of the living species. 70,000 fossil species are also known. Most classifications recognize ten molluscan classes (two extinct). One class, the gastropods (snails and slugs), contains about 80% of mollusc species.

A very rich molluscan fossil record dates back 500 million year to the Precambrian. The evolutionary origins of molluscs are still disputed, but recent well-respected molecular phylogenetic analyses place the molluscs in the Lophotrochozoa, along with annelids, brachiopods, bryozoans and several other phyla (Halanych et al. 1995). Relationships within the Mollusca are also unclear and disputed; some recent analyses challenge whether this enormous phylum is a natural, monophyletic group (Sigwart and Sutton 2007 and references therein).

Non-marine molluscs appear to have a very high extinction rate. Lydeard et al (2004) list terrestrial and fresh water mollusc extinctions as about 40% of total recorded animal extinctions, far greater than marine molluscs.

Many different molluscs have been integrated into human culture since prehistoric times in a plethora of forms: shell money, jewelry and food, crop pests, and disease carriers (Schistosomiasis is a watersnail-born parasite that effects hundreds of millions of people in the world).