656
Journal of Paleontology 91(4):643–661
Figure 7. Echinoderms as benthic islands: (1) Post-mortal encrustation of the early Miocene Clypeaster latirostris from the Molasse Zone (Austria) by balanid barnacles; (2) Post-mortal encrustation of the Recent spantagoid Ova canalifera from the Northern Adriatic Sea by serpulid worm tubes (see Nebelsick, 2004); (3) Post-mortal encrustation of the Recent sea urchin Echinus sp. from Ibiza (Balearic Islands, Spain) mainly by bryozoans and serpulid worms. Scale bars are (1, 2) 1 cm; (3) 5 cm.
considered as benthic islands for many epidionts; in these cases the colonization may occur on both living and dead specimens (e.g., Brett and Eckert, 1982; Liddell and Brett, 1982; Peters and Bork, 1998; see also previous section concerning bioclaustra- tions). In addition, cystoid thecae used as benthic islands have been also documented in the fossil record (Thomka et al., 2016). From Miocene Clypeaster tests deposited in a soft and non-
consolidated biocalcarenite of northeastern Spain (i.e., benthic islands), Belaústegui et al. (2013) described two modes of occurrence for traces (cf. Gastrochaenolites dijugus) produced by gastrochaenid bivalves: (1) ‘intrastereom clavate borings’ restricted to the echinoid stereom; and (2) ‘semi-endoskeletal dwellings’, which penetrate across the test wall and extend as carbonate crypts into the sediment fill of the internal test cavity. Jagt et al. (2009) erected the new ichnospecies Petroxestes
altera to describe shallow, elongate borings with narrowly rounded ends and irregular bioglyphs (grooves and scratches) produced in a hemipneustid echinoid (late Maastrichtian, Belgium). Jagt et al. (2009) interpreted that this trace was pro- duced by a sipunculid worm, after the death of the echinoid.
Conclusions
Modern and fossil bioturbation and bioerosion structures pro- duced and interpreted as produced by echinoderms are reviewed. In the trace fossil record, 22 ichnogenera and 53 ichnospecies have been erected; among them, and following latest ichnotaxonomic discussions, 14 ichnogenera and 26 ichnospecies are considered as valid ichnotaxa. All of them correspond to bioturbation structures except four ichnogenera interpreted as bioerosion structures. Echinoid and asterozoan traces exhibit the highest ichno-
diversities both in the fossil record and today. By contrast, the ichnological record of other groups, such as crinoids or mitrates, is reduced to unique and exceptional specimens. In addition, although nowadays the different burrowing behaviors con- ducted by holothurians are really well known, their trace fossil
record is limited at this time to a single ichnogenus and two ichnospecies. This contribution highlights the importance of the combi-
nation of neo- and paleoichnological studies as a very powerful tool to interpret and to better understand the ecology and ethology of burrowing and boring invertebrates (echinoderms, in this particular case), and the trace fossil record.
Acknowledgments
The authors acknowledge the photographic material provided by S. Zamora (Instituto Geológico y Minero de España, Zaragoza, Spain) and M. Ballesteros (Universitat de Barcelona, Spain) and the comments of D. Knaust (Statoil, Norway). Many thanks to W. Gerber for excellent photographs and T. Grun (both Department of Geosciences, Tübingen) for help in sample preparation. The authors appreciate the comments provided by the editor B. Hunda (Cincinnati Museum Center, USA) and the reviews of A. Uchman (Jagiellonian University, Poland) and C. Brett (University of Cincinnati, USA); their suggestions have contributed to improve the original manuscript. This study is part of the activities of the Research Group
RNM 293 “Geomorfología Ambiental y Recursos Hídricos”, University of Huelva (FM). Z.B. is supported by a BDR post-doctoral fellowship of the University of Barcelona.
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