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14th Swiss Geoscience Meeting, Geneva 2016

Detailed moulting behaviour preserved in the , South Australia

Harriet B Drage*,**, James D Holmes***, Diego C García-Bellido****, Allison C Daley*,**,*****

*Department of Zoology, University of Oxford, Tinbergen Building, South Parks Road, Oxford, UK, OX1 3PS ([email protected]) **Oxford University Museum of Natural History, Parks Road, Oxford, UK, OX1 3PW ***Department of Biological Sciences, University of Adelaide, South Australia, SA 5005 ****South Australian Museum, North Terrace, Adelaide, South Australia, SA 5000 *****Faculty of Geosciences and Environment, University of Lausanne, Sorge Géopolis, CH-1015 Lausanne, Switzerland

The Emu Bay Shale (EBS; , South Australia) is an early (c. 514 Ma) locality with exceptional preservation of soft tissues (Konservat-Lagerstätte). The locality is important for expanding our knowledge of the earliest evolved animals, and is particularly interesting for its unique preservation. This resulted from rapid burial at an active tectonic margin and sediment anoxia, facilitated by low levels of abiotic (water currents) and biotic (bioturbation, predation) disruptive processes (Gehling et al., 2011).

The EBS displays a diverse -type fauna, however the composition is dominated by both preserved carcasses and moulted exoskeletons of two trilobite species, Estaingia bilobata in the majority, and Redlichia takooensis (Paterson et al., 2016). The excellent preservational conditions at the EBS have produced a great number of virtually undisturbed moulted exoskeleton assemblages for these taxa. These assemblages capture the moment of exuviation, describing the sequence of movement and pattern of disarticulation during moulting. This has allowed for an unprecendented interpretation of detailed behavioural information from trilobite moults.

Extensive field collections of E. bilobata and R. takooensis housed in the South Australian Museum (Adelaide) were surveyed. Moult assemblages of both species displaying the total observable range of variation in moulting behaviour were chosen for closer examination. Moulting behaviour and movement during exuviation were described and interpreted for each specimen. These data were then compared to the observable moulting behaviour for the extremely numerous species Ogygopsis klotzi (Burgess Shale) and kingii (, Utah) from other localities with exceptional preservation.

The resulting observations and inferences made for movement during moulting are much more detailed for the EBS than those from the other localities. Very rare moulting events requiring unusual patterns of movement are discernable for E. bilobata and R. takooensis, because of the style of preservation at the EBS. For example, both species usually disarticulate the free cheeks (these are often associated with the cranidium and form a lower cephalic unit in E. bilobata, and laterally inverted for R. takooensis). However, they also rarely demonstrate moulting through disarticulation of the entire cephalon (=Salter’s configuration). The latter assemblages are not observed at the other localities, which often show non-moulting related disruption in the placement of disarticulated . 14th Swiss Geoscience Meeting, Geneva 2016

The EBS assemblages suggest that trilobite moulting behaviour, even within a single species, was more variable than expected. As a group, trilobites displayed flexibility in moulting, presumably to adapt to different conditions and circumstances.This only becomes obvious through the observation of large collections of moult assemblages from localities with exceptional preservation and almost no transportation of exoskeletal sclerites prior to preservation.

Figure 1. Moulted exoskeleton assemblages of Estaingia bilobata (top left) and Redlichia takooensis (top right) from the Emu Bay Shale. The bottom two specimens of R. takooensis display the rare Salterian moult configuration (disarticulation and inversion of entire cephalon). The bottom panel interprets exuvial movement (grey) for E. bilobata, leaving the moult assemblage (white). Scale bars 5mm for E. bilobata, 10mm for R. takooensis. Photographs: HBD.

REFERENCES Paterson, J.R., García-Bellido, D.C., Jago, J.B., Gehling, J.G., Lee, M.S.Y. & Edgecombe, G.D. 2016: The Emu Bay Shale Konservat-Lagerstätte: a view of Cambrian life from East Gondwana. J. Geol. Soc, 173, 1-11. Gehling, J.G., Jago, J.B., Paterson, J.R., García-Bellido, D.C. & Edgecombe, G.D. 2011: The geological context of the Lower Cambrian (Series 2) Emu Bay Shale Lagerstätte and adjacent stratigraphic units, Kangaroo Island, South Australia. Aust. J. Earth Sci., 58, 243-257.