Epifaunal Worm Tubes on Lower Lias Ammonites – Conclusions and References

Table of Contents

  1. Summary & Introduction
  2. Epifaunal Worm Tubes on Lower Lias Ammonites – Results
  3. Epifaunal Worm Tubes on Lower Lias Ammonites – Detailed Observations and interpretation
  4. Epifaunal Worm Tubes on Lower Lias Ammonites – Discussion
  5. Epifaunal Worm Tubes on Lower Lias Ammonites – Conclusions and References

6. Conclusions

1) Epifaunal serpulids that were themselves overgrown by the ammonites and those that reacted to the growth of the ammonites attached to juvenile, living ammonites. Their growth may be used to interpret the growth of the ammonites. Epifaunal serpulids that are attached to one side of, and grew randomly with respect to the ammonite attached after the death of the ammonite.
2) The commonest pattern of growth in serpulids that undoubtedly attached to growing live ammonites was attachment in the umbilical seam on one side, later growth onto, and then around, the venter. Reconstructing this ‘standard pattern’ shows that serpulid growth maintained the worm tube aperture at about 6 o’clock with respect to the orientation of the living ammonite. The worm tube aperture was approximately 105-115o behind the ammonite aperture throughout life. The standard pattern is interpreted as orientating the worm’s feeding currents parallel to those generated by the swimming ammonite and thus increasing food-gathering capacity. The mid-ventral position was doubly advantageous as the worm could deploy its branchia on both sides of the ammonite shell.
3) Growth on ammonites was beneficial to the worms. However, Promicroceras with epifaunal worm tubes died at smaller mean size than unencumbered examples. A significant inverse correlation exists between estimated extra weight of worm tubes and diameter at death of the ammonites. Epifaunal worm tubes were disadvantageous to the ammonites by increasing both weight and drag, and, therefore, were parasitic.
4) We reject the idea that the worms were stealing the ammonite’s food because the two apertures were too far apart, and because worms filter minute particles unlikely to be generated during ammonite feeding.
5) Comparisons with living serpulids tend to confirm that Promicroceras swam backwards, reached full size in two to three years, but continued to live for some time after reaching maturity.

Further Information

For those interested in finding out more about modern serpulid worms the following website is a good place to start.


Paul Davis, The Natural History Museum, London provided a copy of Buys’ paper and we thank Dr. Willem Renema, NCB – Naturalis, Leiden, The Netherlands, for a translation. We also thank several local collectors who contributed specimens.


Andrew, C., Howe, P., Paul, C.R.C. and Donovan, S.K., 2010. Fatally bitten ammonites from the Lower Lias of Lyme Regis, Dorset. Proceedings of the Yorkshire Geological Society 58, 81-94. doi: 10.1144/pygs.58.1.276
Bailey-Brock, J.H., 1976. Habitats of tubicolous polychaetes from the Hawaiian Islands and Johnston Atoll. Pacific Science 30, 69-81.
Buys, J., 1973. Symbiosen van serpula’s met ammonitien uit de Onder Lias van Dorset (Zuid-Engeland). Grundboor en Hamer 1973, 62-67.
Chapman, N.D., Moore, C.G., Harries, D.B. and Lyndon, A.R., 2007. Recruitment patterns of Serpula vermicularis L. (Polychaeta, Serpulidae) in Loch Creran, Scotland. Estuarine, Coastal and Shelf Science 73, 598-606.
Cotter, E., O’Riordan, R.M.and Myers, A.A., 2003. Recruitment patterns of serpulids (Annelida: Polychaeta) in Bantry Bay, Ireland. Journal of the Marine Biological Association of the United Kingdom 83, 41-48.
Cox, B.M., Sumbler, M.G. and Ivimey-Cook, H.C., 1999. A formational framework for the Lower Jurassic of England and Wales (onshore area). British Geological Survey Research Report, RR/99/01, 28 pp.
Dales, R.P., 1957. Some quantitative aspects of feeding in sabellid and serpulid fan worms. Journal of the Marine Biological Association of the United Kingdom 36, 309-316.
Davies, B.R., Stuart, V. and Villiers, M. de, 1989. The filtration activity of a serpulid polychaete population (Ficopomatus enigmaticus (Fauvel)) and its effects on water quality in a coastal marina. Estuarine, Coastal and Shelf Science 29, 613-620.
Dietl, J., Nascimento, C. and Alexander, R., 2000. Influence of ambient flow around the horseshoe crab Limulus polyphemus on the distribution and orientation of selected epizoans. Estuaries 23, 509-520.
Fauchald, K. and Jumars, P.A., 1979. The diet of worms: a study of polychaete feeding guilds. Oceanography and Marine Biology, Annual Reviews 17, 193-284.
Gray, J., 1928. Ciliary Movement. Cambridge University Press, Cambridge, 162 pp.
Hill, M.B., 1967. The life cycles and salinity tolerance of the serpulids Mercierella enigmatica Fauvel and Hydroides uncinata (Philippi) at Lagos, Nigeria. Journal of Animal Ecology 36, 303-321.
Jacobs, D.K. and Chamberlain, J.A. Jr, 1996. Buoyancy and hydrodynamics in ammonoids. 169-224. In Landman, N.H., Tanabe, K. and Davis, R.A., (eds). Ammonoid Paleobiology. Plenum Press, New York, xxiv + 857 pp.
Kupriyanova, E.K., Nishi, E., Ten Hove, H.A. and Rzhavsky, A.V., 2001. Life-history patterns in serpulimorph polychaetes: ecological and evolutionary perspectives. Oceanography and Marine Biology, Annual Review, 39, 1-101.
Lang, W.D. and Spath, L.F., 1926. The black marl of Black Ven and Stonebarrow, in the Lias of the Dorset Coast. Quarterly Journal of the Geological Society, London 82, 144-187.
Lang, W.D., Spath, L.F., Cox, L.R. and Muir-Wood, H.M.,1928. The Belemnite Marls of Charmouth, a series in the Lias of the Dorset coast. Quarterly Journal of the Geological Society, London 84, 179-257.
Lange, W., 1932. Über Symbiosen von Serpula mit Ammoniten im unteren Lias Norddeutschlands. Zeitschrift der Deutschlands Geologischen Gesellschaft 84, 229-234.
Maeda, H. and Seilacher, A. 1996. Ammonoid taphonomy. 543-578. In Landman, N.H., Tanabe, K. and Davis, R.A., (eds). Ammonoid Paleobiology. Plenum Press, New York, xxiv + 857 pp.
Marsden, J.R. and Meeuwig, J. 1990. Preferences of planktotrophic larvae of the tropical serpulid Spirobranchus giganteus (Pallas) for exudates of corals from a Barbados reef. Journal of Experimental Marine Biology and Ecology 137, 95-104.
Medernach, L., Jordana, E., Grémaire, A., Nozias, C., Charles, F. and Amouroux, J.M., 2000. Population dynamics, secondary production and calcification in a Mediterranean population of Ditrupa arietina (Annelida: Polychaeta). Marine Ecology Progress Series 199, 171-184.
Meischner, D., 1968. Perniziöse Epökie von Placunopsis auf Ceratites. Lethaia 1, 156-174.
Merkt, J., 1966. Über Austern und Serpeln als Epöken auf Ammonitengehäusen. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen 125, 467-479.
Page, K.N., 2010. Stratigraphical framework. 33-53. In Lord, A.R., Davis P.G., (eds) Fossils from the Lower Lias of the Dorset Coast. Palaeontological Association, ‘Field Guides to Fossils’, 13, viii + 436 pp.
Paul, C.R.C. and Simms, M.J., 2011. Epizoans on ammonites from the Lower Jurassic of the Severn Basin, southern England, and their palaeoenvironmental and taphonomic significance. Proceedings of the Geologists’ Association, doi:10.1016/j.pgeola.2011.11.008
Saunders, W.B. and Spinosa, C., 1979. Nautilus movement and distribution in Palau, Western Caroline Islands. Science 204, 1199-1201.
Schindewolf, O.H., 1934. Über Epöken auf Cephalopoden-Gehäusen. Paläontologische Zeitschrift 16, 15-31.
Seilacher, A., 1960. Epizoans as a key to ammonoid ecology. Journal of Paleontology 34, 189-193.
Seilacher, A., 1982. Ammonite shells as habitats in the Posidonia Shales of Holzmaden – floats or benthic islands? Neues Jahrbuch für Geologie und Paläontologie, Monatshefte 1982, 98-114.
Simms, M.J., Chidlaw, N., Morton, N. and Page, K.N. (eds), 2004. British Lower Jurassic Stratigraphy. Geological Conservation Review Series, 30. Joint Nature Conservation Committee, Peterborough, xvi + 458 pp.
Trueman,, A.E., 1941. The ammonite body-chamber, with special reference to the buoyancy and mode of life of the living ammonite. Quarterly Journal of the Geological Society, London 96, 339-383.
Wani, R., Kase, T., Shigeta, Y. and Ocampo, R. de, 2005. A new look at ammonoid taphonomy, based on field experiments with modern chambered nautilus. Geology 33, 849-852.