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Friday, 13 June 2025

Malformations in Trilobites from the Silurian and Devonian of Europe.

Malformations in fossils, such as pathologies caused by infections, scars left by recovered injuries, or teratologies caused by developmental problems, can tell us a lot about how extinct organisms grew and interacted with their environments, although when analysing these is clearly preferable to have access to non-malformed examples of the same species, or at least a close relative.

Trilobites were a diverse and abundant group of Arthropods which appeared early in the Cambrian, and survived until the End Permian Extinction. Their dorsal exoskeleton was heavily biomineralized, and was shed periodically to allow the animals to grow. This has lead to an extensive fossil record with Trilobites being extremely common in many Palaeozoic marine deposits. This abundant fossil record makes Trilobites an excellent candidate group for the study of malformations.

In a paper published in the journal Acta Palaeontologica Polonica on 22 April 2025, Russell Bicknell of the Division of Paleontology (Invertebrates) at the American Museum of Natural History, and the Palaeoscience Research Centre at the University of New EnglandPatrick Smith of the Palaeontology Department at the Australian Museum Research Institute, and the Department of Biological Sciences at Macquarie UniversityLisa Amati of Paleontology at the New York State Museum, and Melanie Hopkins, also of the Division of Paleontology (Invertebrates) at the American Museum of Natural History, describe malformations in European Silurian and Devonian Trilobite specimens from the collections of the Natural History Museum in London and the New York State Museum.

The first specimen examined by Bicknell et al., NYSM 19739, is an isolated cephalon (head part) from a Harpetid Trilobite, Lioharpes venulosus, from the Early Devonian Koněprusy Limestone of the Czech Republic, in the collection of the New York State Museum. The cephalon is 26.2 mm long and 22.3 mm wide with a u-shaped indentation on its right marginal rim. This indentation is 5.6 mm long and extends 1.9 mm towards the midline. The marginal rim is covered in small circular pits, which around the indentation are irregular, ovate, and occasionally fused into larger pits.

Malformed Harpetid Trilobite Lioharpes venulosus, NYSM 19739 from the Koněprusy Limestone, Pragian, Lower Devonian, Koněprusy, Czech Republic. (A₁) complete cephalon; (A₂) close up showing U-shaped indentation (arrow). Specimen coated in ammonium chloride sublimate. Bicknell et al. (2025).

Bicknell et al. note that malformations to the cephalic fringes of Harpetid Trilobites have been recorded before, and that these are usealy attributed to injuries, an analysis with which they concur. However, they also observe that injuries can happen in a variety of ways, with fringe injuries having previously attributed to problems during moulting, failed predation attempts, or unknown causes. They suggest that a moulting injury is the most likely explanation for the injury to the Koněprusy specimen, with the delicate fringe likely torn during moulting, and the enlarged and fused pits being a result of fusion of the torn margin during healing. Various purposes have been suggested for the cephalic fringes of Harpetid Trilobites, including filtering for food, sensory roles, sediment ploughing, hydrostatic support, cephalic reinforcement, burrowing, and enhancing hydrodynamic efficiency. Whatever the purpose of this organ, an injury to it is likely to have been detrimental to the living Trilobite, and presumably repairing this injury would have been a priority during subsequent moults.

The second specimen, NHMUK PI In 65061, looked at is a Phacopid Trilobite, Calymene blumenbachii, from the Early Silurian Much Wenlock Limestone Formation of Shropshire, England, in the collection of the Natural History Museum. This specimen comprises a partial cephalon, thorax, and pygidium, with a total length of 92.9 mm and a width of 48.6 mm. The second thoracic axial ring (middle part of the second segment of the thorax) of this specimen is covered by a structure with closely spaced openings, which has an elevated round crater at its right extremity, with an opening 1.7 mm across.

Bicknell et al. interpret this as an encrusting Trepostome Bryozoan covering the 3rd thoracic tergite, with the larger opening being an ovate zoarium (specially modified zooid which produced eggs). The restriction of the encrustation to one tergite strongly suggests that this happened while the Trilobite was alive, and that the Bryozoan colony was therefore unable to overgrow the articulations between tergites.

Abnormal Calymenid Trilobites Calymene blumenbachii from the Much Wenlock Limestone Formation, Homerian, Wenlock, Silurian, England, UK. (A) NHMUK PI In 65061, (A₁) complete specimen; (A₂) close up showing the large bryozoan growth. (B) NHMUK PI In 19857 showing pygidial ribs that terminate early (white arrows) and are fused proximal to the medial lobe (black arrow).  Bicknell et al. (2025).

The next specimen examined, NHMUK PI In 19857, is another example of Calymene blumenbachii from the Much Wenlock Limestone. This specimen is a partial pygidium (tail part) 13.2 mm long and 18.2 mm wide. On the right side of this specimen the pygidial ribs are disrupted and irregular, with two ribs terminating 1.6 mm short of the pygidial margin, while another two fuse 1.2 mm from the pygidial axis.

Bicknel et al. observe that similar deformations to the pygidia have been observed in a wide range of other Trilobites, including Dalmanities pleuroptyxDechenella macrocephalusNiobina sp., and Prionopeltis archiaci. They attribute these deformations to genetic or developmental issues, but suggest that the limited disruption to the pygidium they caused did not represent a major handicap.

Specimen NYSM 19740 is an Acastid Trilobite, Treveropyge sp., from the early Devonian Saint Céneré Formation of Mayenne in northwest France. this specimen is another isolated pygidium, s 11.6 mm long and 17.9 mm wide. It has a deformation to the axial lobe, which is asymmetric, with two of the axial rings malformed and curving to the right.

Malformed Acastid Trilobite Treveropyge sp., NYSM 19740 from the Saint Céneré Formation, Lochkovian, Lower Devonian, Mayenne, France. (A₁) Complete pygidium; (A₂) close up showing asymmetrical axial lobe and incomplete axial ring (arrows). Specimen coated in ammonium chloride sublimate. Bicknell et al. (2025).

Again, Bicknell et al. note that similar deformations have been seen in other Trilobites, such as Calliops marginatusDolicholeptus licticallis, and Sceptaspis lincolnensis. They suggest that these malformations are caused by genetic deformations, leading to incomplete development or non-functional somites.

The final specimen examined, NHMUK PI I 1108, is an external impression of a partial pygidium belonging to the Styginid Trilobite Scutellum (Scutellum) pardalios, from the Middle Devonin Barton Limestone Member of Devon, England. This impression is t is 59.5 mm long and 44.0 mm wide. On the right side of this impression (i.e. on the left side of the Trilobite) two ribs fuse into a single rib 29.1 mm from the pygidial axis. This single rib then terminates 4.8 mm from the pygidial margin.

Malformed Styginid Trilobite Scutellum (Scutellum) pardalios, NHMUK PI I 1108 from the Barton Limestone Member, Torquay Limestone Formation, Givetian, Middle Devonian, England, UK. (A₁) Pygidium preserved as external impression; (A₂) close up showing fused pygidial pleurae (arrows). Bicknell et al. (2025).

Bicknell et al. note that the surface of the pygidium was covered with ornamentation, with no visible break in this, which appears to  rule out the malformation having been formed by an unsuccessful predation attempt, or similar injury. Recovery from injury is the most commonly sited reason for malformations seen in Styginid Trilobites, followed by parasitic infections during early development, which seems equally unlikely. Bicknell et al. suggest instead that this deformity might be the result of a difficult moult, or possibly a genetic aberration. They do not believe this minor deformity would have significantly affected the living Trilobite.

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