The Glyceriforms: Stabby Worms and Grabby Worms

Historically, the annelid worms have been considered a difficult group to classify. Whereas most of the recognised families have been fairly well established, higher taxa uniting these families have tended to be a bit on the vague side. Nevertheless, there are some supra-familial groups that can be considered well established, one such group being the Glyceriformia.

Specimen of Goniadidae (head to the right), from NOAA Fisheries.

The glyceriforms are two families of marine worms, the Glyceridae and Goniadidae. More than a hundred species are known in this clade (over forty glycerids and over sixty goniadids), found in habitats ranging from the intertidal to the abyssal. They range in size from about a centimetre in length to well over half a metre. The front end of the body tapers to a narrow, elongate conical point in front of the mouth, bearing two terminal pairs of small, slender appendages that may correspond to the antennae and palps of other worms. Eyes may be present or absent. The pharynx forms a remarkably elongate, eversible proboscis. In Glyceridae, the proboscis ends in a ring of four hook-shaped jaws, all similar to each other. In Goniadidae, the arrangement of jaws is more complex with the usual arrangement being small micrognaths on one side of the ring and larger macrognaths on the other. Glycerids usually have a transparent skin and an overall red or white colour reflecting the coloration of the internal fluids (red-coloured individuals are sometimes known as 'bloodworms', as are many other similarly coloured worm-like invertebrates). Goniadids have a more opaque cuticle and often have an iridescent sheen (Rouse & Pleijel 2001).

Glycera dibranchiata with everted proboscis, from the Yale Peabody Museum.

Glyceriforms most commonly live as burrowers in muddy or sandy substrates though some live on the surface of rocks. Most are carnivores of active invertebrates such as crustaceans or other worms; some may be detritivores. They may be vagile or they may construct permanent galleries of burrows with multiple entrance and exit openings in which they wait to lunge at anything foolish enough to pass nearby. In glycerids, the stabby jaws are associated with venom glands leading to ducts opening through pores on the jaw's underside. In some species, this venom is strong enough to cause a painful reaction in humans (though I haven't come across any references to long-term consequences). Goniadids lack venom glands and seem to rely on the physical use of their jaws to capture prey. As with many other marine worms, reproduction happens via pelagic epitokes. As a suitable time approaches (Prentiss, 2020, records goniadid epitokes emerging only during a full moon), the glyceriform worm undergoes a metamorphosis involving the break-down of the digestive system and enlargement of the parapodia. The transformed epitokes swim towards the surface where they release gametes through ruptures of the body wall, ending their life in a suicidal orgasm.

Close-up on proboscis of Glycera alba, copyright Hans Hillewaert.

Because of their hardened jaws, which are mostly constructed of protein but partially mineralised, glyceriforms have quite a good fossil record compared to many other worms (Böggemann 2006). Fossilised glyceriform jaws have been found as far back as the Triassic and are little different from those of modern glyceriforms. Body fossils are, unsurprisingly, much rarer but a worm from the Carboniferous Mazon Creek fauna, Pieckonia helenae, has been identified as a stem-group goniadid. The glyceriform body plan seems to have been a very successful one, remaining essentially unchanged over hundreds of millions of years.

REFERENCES

Böggemann, M. 2006. Worms that might be 300 million years old. Marine Biology Research 2: 130–135.

Prentiss, N. K. 2020. Nocturnally swarming Caribbean polychaetes of St. John, U.S. Virgin Islands, USA. Zoosymposia 19: 91–102.

Rouse, G. W., & F. Pleijel. 2001. Polychaetes. Oxford University Press.