Spiral feather-duster worm Spirorbis, photographed by Piotr Rotkiewicz.

Belongs within: Sabellida.
Contains: RotulariaSpirorbis, Hydroides, Serpulinae.

The Serpulidae are a mostly marine (occasionally freshwater) group of sessile worms that produce calcareous tubes. As such, they are one of the few worm groups to have an extensive fossil record. Prominent genera include Spirorbis, in which the small tube is coiled in a flat spiral and attached to the substrate (Howell 1962).

Serpularia: a rightly forgotten problematicum
Published 15 August 2015

I think it may be time to rock out something that hasn’t been seen on this site for a while. Horns at the ready…

(Credit, again, to Neil from Microecos). And I’m afraid that may just be the most excitement that we get in this post. While some fossils are problematic because they’re so strange that they can’t be easily compared to living animals, others are problematic simply because they’re rubbish.

In 1840, the palaeontologist Georg Graf zu Münster (‘Graf’ being a German title that generally gets translated as ‘Count’) published his Beiträge zur Petrefakten-Kunde, in which he described a number of fossils held in his collection. This book included a section on fossils from the Ordovician Orthoceratite Limestone of the Fichtel Mountains in Bavaria. Which, close to the end, included this little tidbit:

Unter mehreren Bruchstücken einiger mir noch unbekannten Versteinerungen kommen auch einige röhrenformige Korper vor, welche ich anfänglich für den von Murchison aus der 27sten Tafel abgebildeten Myrianites hielt, allein genaue Untersuchung zeigte, dass diese Korper formliche Schalen hatten und daher vielleicht zu den Serpuliten gehört hatten, daher ich sie vorläufig Serpularia genannt habe. Aus der Taf. IX. Fig. 14 und 15 sind zwei Arten von dergleichen Bruchstücken abgebildet; Fig. II. Serpularia crenata; glatt gebogene Röhre, aus dem Rücken crenulirt. Fig. 15. Serpularia bicrenata; glatte etwas zusammengedrückte ganz grade Röhrchen, die an beiden Seiten crenulirt sind.

Translated with the help of Google Translate, I think this means: “Among several fragments of fossils unknown to me occured a tube-like body, which I initially took for Myrianites as figured by Murchison in the 27th plate, until close examination showed that this body had distinct signs of segmentation and was therefore perhaps one of the Serpulidae. Therefore, I have provisionally called it Serpularia. On Plate IX Figs 14 and 15 are shown two types of the like fragments; Fig. 14, Serpularia crenata: smooth curved tube crenulated from the back. Fig. 15, Serpularia bicrenata: smooth, slightly compressed, quite straight tubes that are crenulated on both sides“.

Münster’s (1840) original figures of the two Serpularia.

As perfunctory as it was, that seems to be all there was to say on the matter. The good Graf’s Serpularia has pretty much never been mentioned again*, beyond being cited to cause a name change in a later homonymous gastropod genus, and a brief listing in Howell’s (1962) coverage of worm fossils for the Treatise on Invertebrate Paleontology that adds nothing to the original description.

*Though if it were to be mentioned again, it would probably have to be under a different name. The name ‘Serpularia‘ had earlier been used by Fries in 1829 for a genus of slime moulds. At the time, slime moulds were treated as fungi, and hence fell under the purview of botanical rather than zoological names, but with the recognition that they are amoebozoans an increasing number of authors would move them into the field of the Zoological Code.

Münster believed that his fossils belonged to the Serpulidae, a family of annelid worms. Annelids, being mostly soft and squishy things that do not stand up well to decay, have a pretty deplorable fossil record, but serpulids are a bit of an exception. These are sessile worms that secrete a calcareous tube in which they live their lives. Unfortunately, while these tubes are eminently fossilisable, they are also a bit nondescript, and have little to mark them as uniquely serpulid.

Because of the dominance of annelids among modern worms, there has been a definite tendency in the past to assume that any given worm-like fossil represents an annelid. Howell’s (1962) aforementioned list of annelids includes the Ediacaran Spriggina (identity still under debate, but probably not an annelid) and the Cambrian Pikaia (now generally regarded as an early chordate). Similarly, any worm-like tube has been assumed a serpulid. But even among annelids, serpulids are not the only tube-bearing worms. At least two other families, the Sabellidae and the Cirratulidae, include species producing calcareous tubes. There are also other groups of non-annelid worms that, though relatively uncommon or unprepossessing today, may have been more prominent in the past. After all, we are talking here about a period of hundreds of millions of years. We know that vertebrates have gone through a great deal of evolutionary change over that period; why should we assume that worms have not?

So while fossils have been assigned to the serpulids going back as far as the Cambrian (if not beyond), there is little reason to take those assignations at face value. When so-called Palaeozoic serpulids have been examined critically in recent years, they have so far proven to lack features that would definitely confirm their identification (Vinn & Mutvei 2009). Weedon (1994) found that Palaeozoic fossils that had been assigned not only to the Serpulidae, but to the modern genus Spirorbis, had a shell microstructure that suggested a relationship to bryozoans or brachiozoans rather than to annelids. Without a similar close analysis, we could not assume a priori that Münster’s Serpularia were not serpulids, but odds would currently be against it.

Systematics of Serpulidae

Characters (from Howell 1962): Body cylindrical; thorax with three to seven segments bearing bundles of hairlike bristles on dorsal side and hooks on ventral side; abdomen with many segments bearing hooks on dorsal side and bundles of hairlike bristles on ventral surface; numerous tentacles around mouth and calcareous or horny operculum; builds calcareous tube that is circular, polygonal, or triangular in cross-section and may be ornamented on outside with concentric raised rings or longitudinal ridges or keels; usually attached for part or all of its length to substratum, sometimes free throughout entire length.

    |--+--Chitinopoma serrula (Stimpson 1854)RP07
    |  `--Protula Risso 1826RP07, H62
    |       |--*P. rudolphi Risso 1826H62
    |       |--P. bispiralis (Savigny 1820)HG14
    |       |--P. palliata (Willey 1905)HG14
    |       `--P. tubulariaM62
          |  `--SerpulinaeRP07
          `--+--Galeolaria Lamarck 1818RP07, H62
             |    |--*G. caespitosa Lamarck 1818H62
             |    `--G. hystrixH79
             `--FicopomatusRP07 [incl. Mercierella Fauvel 1923H62]
                  |--F. enigmaticus (Fauvel 1923)HB-E05 [=*Mercierella enigmaticaH62, H79]
                  |--F. macrodonH79
                  |--F. miamiensis (Treadwell 1934)HB-E05
                  `--F. uschakovi (Pillai 1960)HB-E05
Serpulidae incertae sedis:
  Filograna Oken 1815 [incl. Filogranula Nielsen 1931 non Langerhans 1884]H62
    `--*F. filograna [=Serpula filograna]H62
  Salmacina Claparède 1870H62
    |--*S. incrustans Claparède 1870H62
    |--S. aedificatrixH62
    |--S. australisHS01
    |--S. dysteriPP64
    `--S. tribranchiata (Moore 1923)SS07
  Josephella Caullery & Mesnil 1896H62
    `--*J. marenzelleri Caullery & Mesnil 1896H62
  Filogranula gracilis [incl. Omphalopoma gracilis]H79
  Crucigera zygophoraH79
  Vermiliopsis Saint-Joseph 1906H62
    |--*V. infundibulum (Langerhans 1884) [=Vermilia infundibulum]H62
    |--V. bermudiensis (Bush 1904) [incl. Vermilia glandulata Treadwell 1936]H56
    |--V. elegantulaH62
    |--V. glandigera Gravier 1908HG14
    |--V. multiannulata (Moore 1923) (see below for synonymy)H56
    |--V. multicristataPP64
    `--V. negevensisPTV14
  Pseudovermilia multispinosaH79
  Pomatoleios kraussii (Baird 1865)HG14
  Pomatoceros Philippi 1844H62
    |--*P. triqueter (Linnaeus 1758) [=Serpula triquetra]H62
    |--P. arietinusJ64
    |--P. caeruleus (Schmarda 1861) [=Placostegus caeruleus; incl. Po. davaoensis Treadwell 1942]H56
    |--P. cariniferusL90
    |--P. lamarckiiH79
    |--P. minutus Rioja 1941HB-E05
    `--P. strigicepsB64
    |--S. baileybrockae Pillai 2009HG14
    |--S. cariniferusH79
    |--S. corniculatus (Grube 1862)HG14
    |--S. giganteusH79
    |--S. paumontanusH79
    |--S. polycerus (Schmarda 1861)HB-E05
    |--S. polytrema (Philippi 1844)HB-E05
    |--S. richardsmithii Pillai 2009HG14
    `--S. tetraceros (Schmarda 1861)HG14
  Trigonocrinus Bather 1889WR78
    `--*T. liratus Bather 1889WR78
  Ditrupa Berkeley 1835H62
    |--*D. cornea (Linné 1767) [=Dentalium corneum; incl. De. subulatum Deshayes 1826]H62
    `--D. arietinaM62
  Spirulaea Bronn 1828H62
    `--S. gregaria Etheridge 1907F71 [=Rotularia gregariaB59]
  Cycloserpula Parsch 1956H62
  Dorsoserpula Parsch 1956H62
  Tetraserpula Parsch 1956H62
  Pentaserpula Parsch 1956H62
  Hexaserpula Parsch 1956H62
  Asterosalpinx Sokolov 1948H62
    `--*A. asiaticus Sokolov 1948H62
  Camptosalpinx Sokolov 1948H62
    `--*C. siberiensis Sokolov 1948H62
  Cementula Nielsen 1931H62
    `--*C. sphaerica Nielsen 1931H62
  Chaetosalpinx Sokolov 1948H62
    `--*C. ferganensis Sokolov 1948H62
  Diploconcha Conrad 1875H62
    `--*D. cretacea Conrad 1875H62
  Discouvermetulus Roverto 1904H62
    `--*D. pissarroi Roverto 1904H62
  Ditrupula Nielsen 1931H62
    `--*D. canteriata (von Hagenow 1840) [=Serpula canteriata]H62
  Genicularia Quenstedt 1858 non De Bary 1858 (ICBN)H62
    `--*G. ornata Quenstedt 1858H62
  Hamulus Morton 1834 [incl. Falcula Conrad 1870]H62
    `--*H. onyx Morton 1834H62
  Howellitubus Richardson 1956H62
    `--*H. whitfieldorum Richardson 1956H62
  *Jereminella pfenderaeH62
  Longitubus Howell 1943H62
    `--*L. lineatus (Weller 1907) [=Hamulus lineatus]H62
  Neomicrorbis Roverto 1903H62
    `--*N. granulata (Sowerby 1829) [=Serpula granulata]H62
  Ornatoporta Gardner 1916H62
    `--*O. marylandica Gardner 1916H62
  *Paliurus triangularisH62
  Phragmosalpinx Sokolov 1948H62
    `--*P. australiensis Sokolov 1948H62
  Placostegus Philippi 1844H62
    `--*P. tridentatus (Fabricius 1779) [=Serpula tridentatus]H62
  Proterula Nielsen 1931H62
    `--*P. costata Nielsen 1931H62
  Protulites Jaskó 1940H62
    `--*P. segmentata Jaskó 1940H62
  Pyrgopolon Montfort 1808 [incl. Entalium Defrance 1819, Pharetrium König 1825]H62
    `--*P. mosae Montfort 1808H62
  Sclerostyla Mørch 1863H62
    `--*S. ctenactis (Mørch 1863)H62 (see below for synonymy)
  Semiserpula Wetzel 1957H62
    `--*S. chilensis Wetzel 1957H62
  Serpentula Nielsen 1931H62
    `--*S. ampullacea (Sowerby 1829) [=Serpula ampullacea]H62Serpularia’ Münster 1840 non Fries 1829H62
    `--*S. crenata Münster 1840H62
  Sinuocornu Howell 1959H62
    `--*S. curtum (Salter 1848) [=Serpulites curtus]H62
  Spirorbula Nielsen 1931H62
    `--*S. aspera (von Hagenow 1840) [=Serpula aspera]H62
  Streptindytes Calvin 1888H62
    `--*S. acervulariae Calvin 1888H62
  Torlessia Bather 1905H62
    `--*T. mackayi Bather 1905H62
  Turbinia Michelin 1845H62
    |--*T. graciosa Michelin 1845H62
    `--T. abbreviataH62
  Vermilia Lamarck 1818H62
    `--*V. triquetra Lamarck 1818H62
    |--P. actinoceros (Mörch 1863)HG14
    `--P. polytremaPP64
  ‘Dentalium’ nigrumD26
  ‘Dentalium’ deformeD26
  ‘Dentalium’ pellucidumD26
    |--E. similis Treadwell 1929H56
    `--E. uncinatus Philippi 1844 [incl. E. operculata Treadwell 1929]H56
  Sphaeropomatus Treadwell 1934H56
    `--S. miamiensis Treadwell 1934H56
  ‘Hipponyx’ inexpectata Mestayer 1929D56
  Kimberleya hutchingsae Pillai 2009HG14
  Pseudoprotula kimberleyensis Pillai 2009HG14
  Spiraserpula snellii Pillai & ten Hove 1994HG14

*Sclerostyla ctenactis (Mørch 1863)H62 [=Serpula (*Sclerostyla) ctenactisH62; incl. Placostegus calciferus Treadwell 1929H56, Spirodiscus calciferusH56, Sclerostyla differens Augener 1922H56]

Vermiliopsis multiannulata (Moore 1923) [=Metavermilia multiannulata; incl. V. hawaiiensis Treadwell 1943, V. torquata Treadwell 1943]H56

*Type species of generic name indicated


[B59] Boreham, A. U. E. 1959. Cretaceous fossils from the Chatham Islands. Transactions of the Royal Society of New Zealand 86 (1): 119–125.

[D56] Dell, R. K. 1956. The archibenthal Mollusca of New Zealand. Dominion Museum Bulletin 18: 1–235.

[D26] Deshayes, G.-P. 1826. Anatomie et monographe du genre Dentale. Mémoires de la Société d’Histoire Naturelle de Paris, ser. 2, 2 (2): 321–378, 4 pls.

[F71] Fletcher, H. O. 1971. Catalogue of type specimens of fossils in the Australian Museum, Sydney. Australian Museum Memoir 13: 1–167.

[H56] Hartman, O. 1956. Polychaetous annelids erected by Treadwell, 1891 to 1948, together with a brief chronology. Bulletin of the American Museum of Natural History 109 (2): 239–310.

[HS01] Hayward, B. W., A. B. Stephenson, M. S. Morley, W. M. Blom, H. R. Grenfell, F. J. Brook, J. L. Riley, F. Thompson & J. J. Hayward. 2001. Marine biota of Parengarenga Harbour, Northland, New Zealand. Records of the Auckland Museum 37: 45–80.

[H79] Hove, H. A. ten. 1979. Different causes of mass occurrence in serpulids. In: Larwood, G., & B. R. Rosen (eds) Biology and Systematics of Colonial Organisms pp. 281–298. Academic Press: London.

[HB-E05] Hove, H. A. ten, & M. N. Ben-Eliahu. 2005. On the identity of Hydroides priscus Pillai 1971—taxonomic confusion due to ontogeny in some serpulid genera (Annelida: Polychaeta: Serpulidae). Senckenbergiana Biologica 85 (2): 127–145.

[H62] Howell, B. F. 1962. Worms. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt W. Miscellanea: Conodonts, Conoidal Shells of Uncertain Affinities, Worms, Trace Fossils and Problematica pp. W144–W177. Geological Society of America, and University of Kansas Press.

[HG14] Hutchings, P., C. Glasby, M. Capa & A. Sampey. 2014. Kimberley marine biota. Historical data: polychaetes (Annelida). Records of the Western Australian Museum Supplement 84: 133–159.

[J64] Jeffreys, J. G. 1864. Report of the committee appointed for exploring the coasts of Shetland by means of the dredge. Report of the British Association for the Advancement of Science 33: 70–81.

[L90] Luxton, M. 1990. The marine littoral mites of the New Zealand region. Journal of the Royal Society of New Zealand 20 (4): 367–418.

[M62] Monniot, F. 1962. Recherches sur les graviers a Amphioxus de la région de Banyuls-sur-Mer. Vie et Milieu 13: 231–322.

Münster, G. 1840. Beiträge zur Petrefacten-Kunde von Herm. v. Meyer und Georg Graf zu Münster vol. 3. In Commission der Buchner’schen Buchhandlung: Bayreuth.

[PTV14] Parry, L., A. Tanner & J. Vinther. 2014. The origin of annelids. Palaeontology 57 (6): 1091–1103.

[PP64] Peres, J. M., & J. Picard. 1964. Nouveau manuel de bionomie benthique de la mer Mediterranee. Recueil des Travaux de la Station Marine d’Endoume, Bulletin 31 (27): 5–137.

[RP07] Rousset, V., F. Pleijel, G. W. Rouse, C. Erséus & M. E. Siddall. 2007. A molecular phylogeny of annelids. Cladistics 23: 41–63.

[SS07] Struck, T. H., N. Schutt, T. Kusen, E. Hickman, C. Bleidorn, D. McHugh & K. M. Halanych. 2007. Annelid phylogeny and the status of Sipuncula and Echiura. BMC Evolutionary Biology 7: 57.

Vinn, O., & H. Mutvei. 2009. Calcareous tubeworms of the Phanerozoic. Estonian Journal of Earth Sciences 58 (4): 286–296.

Weedon, M. J. 1994. Tube microstructure of Recent and Jurassic serpulid polychaets and the question of the Palaeozoic ‘spirorbids’. Acta Palaeontologica Polonica 39 (1): 1–15.

[WR78] Wienberg Rasmussen, H. 1978. Articulata. In: Moore, R. C., & C. Teichert (eds) Treatise on Invertebrate Paleontology pt T. Echinodermata 2. Crinoidea vol. 3 pp. T813–T927. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).

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