Grun et al.—Drilling Predation on Miocene Echinocyamus from Malta Conclusions
The two distinct samples of phosphatized and nonphosphatized Echinocyamus stellatus from the Miocene Maltese Island Gozo can be statistically distinguished with respect to test length and drill hole frequency, but they cannot be distinguished on the basis of drill hole length alone. Drilling patterns are quite similar, while minor differences in site selectivity may be size specific. The holes in the Miocene Maltese echinoid tests are inter-
preted as Oichnus simplex (Bromley, 1981) trace fossils. The drill holes are circular to subcircular in outline. Drill hole margins are smooth or ragged depending on the microstructures of the drilling area. The drill hole wall is concave in section and perpendicular to the test surface. Comparisons of the morphological characteristic and sizes
of the drill holes from Malta with descriptions of other drill holes in Echinocyamus species indicate that the drillers of the Miocene Echinocyamus stellatus are most likely cassid gastropods. Drilling frequencies vary from 8.1% in nonphosphatized
echinoids to 20.5% in phosphatized echinoids. Variations in drilling frequencies may be due to variations of predator densities or a number of environmental differences and mirror the high variation of reported drilling frequencies within various Recent depositional environments. The fact that predators drill more frequently into the aboral
test side may be due to handling effects and the time and energy saved by drilling the thinner, highly porous areas of the test. The position of internal organs and internal supports may also be important factors. Drilling predation traces can be used for analyzing direct
predation patterns on well-preserved fossil echinoid tests. The drill holes can also provide indirect evidence for the existence of predators and allow for the analyses of specific drill hole morphologies, sizes, and drill site preferences.
Acknowledgments
We thank G. Zammit-Maempel (formerly Malta National Museum of Natural History) for the collection and export permit. Thanks to the Natural History Museum Vienna, Austria, for providing lodging during a research stay. Thanks also to T. Baumiller (Department of Earth and Environmental Sciences, University of Michigan, USA), G. Dietl (Department of Earth and Atmospheric Sciences, Cornell University, USA) and S. Zamora (Instituto Geológico y Minero de España, Zaragoza, Spain) for reviewing, editing, and constructive suggestions.
References
Abels, H.A., Hilgen, F.J., Krijgsman, W., Kruk, R.W., Raffi, I., Turco, E., and Zachariasse, W.J., 2005, Long-period orbital control on middle Miocene global cooling: Integrated stratigraphy and astronomical tuning of the Blue Clay Formation on Malta: Paleoceanography, v. 20, p. 1–17.
Baumiller, T.K., 2003, Evaluating the interaction between platyceratid gastropods and crinoids: A cost-benefit approach: Palaeogeography, Palaeoclimatology, Palaeoecology, v. 201, p. 199–209.
Baumiller, T.K., Leighton, L.R., and Thompson, D.L., 1999, Boreholes in Mississippian spiriferide brachiopods and their implications for Paleozoic
641
gastropod drilling: Palaeogeography, Palaeoclimatology, Palaeoecology, v. 147, p. 283–289.
Baumiller, T.K., Bitner, M.A., and Emig, C.C., 2006, High frequency of drill holes in brachiopods from Pliocene of Algeria and its ecological implica- tions: Lethaia, v. 39, p. 313–320.
Bellanca, A., Sgarrella, F., Neri, R., Russo, B., Sprovieri, M., Bonaduce, G., and Rocca, D., 2002, Evolution of the Mediterranean Basin during the late Langhian–early Serravallian: An integrated paleoceanographic approach: Rivista Italiana di Paleontologia e Stratigrafia, v. 108, p. 223–239.
Bengtson, S., and Zhao, Y., 1992, Predatorial borings in late Precambrian mineralized exoskeletons: Science, v. 257, p. 367.
Bennett, S.M., 1980, Palaeoenvironmental studies in Maltese mid-Tertiary carbonates [Ph.D. dissertation]: London, University of London, 347 p.
Bianucci, G., Gatt, M., Catanzariti, R., Sorbi, S., Bonavia, C.G., Curmi, R., and Varola, A., 2011, Systematics, biostratigraphy and evolutionary pattern of the Oligo-Miocene marine mammals from the Maltese Islands: Geobios, v. 44, p. 549–585.
Boggild (née Challis), G.R., and Rose, E.P.F., 1984, Mid-Tertiary echinoid biofacies as palaeonvironmental indices: Annales Géologiques des pays Helléniques, v. 32, p. 57–67.
Bromley, R.G., 1981, Concepts in ichnology illustrated by small round holes in shells: Acta Geológica Hispánica, v. 16, p. 55–64.
Capeder, G., 1906, Fibularidi del Miocenemedio di S. Gavino amare (Portotorres) Sardegna: Bollettino della Società Geologica Italiana, v. 25, p. 195–534.
Carbone, S., Grasso, M., Lentini, F., and Pedley, H.H., 1987, The distribution and palaeoenvironment of early Miocene phosporites of southeast Sicily and their relationships with the Maltese phosphorites: Palaeogeography, Palaeoclimatology, Palaeoecology, v. 58, p. 35–53.
Carriker, M.R., 1981, Shell penetration and feeding by naticacean and muricacean predatory gastropods: A synthesis: Malacologia, v. 20, p. 403–422.
Carriker, M.R., and Yochelson, E.L., 1968, Recent gastropod boreholes and Ordovician cylindrical borings: U.S. Geological Survey Professional Paper, v. 953, p. 1–26.
Ceranka, T., and Złotnik, M., 2003, Traces of cassid snails predation upon the echinoids from the middle Miocene of Poland: Acta Palaeontologica Polonica, v. 48, p. 491–496.
Challis, G.R., 1979, Miocene echinoid biofacies of the Maltese Islands: Annales Géologiques des Pays Helléniques, tome hors série, v. 1, p. 253–262.
Challis, G.R., 1980, Palaeoecology and Taxonomy of mid-Tertiary Maltese echinoids [Ph.D. dissertation]: London, University of London, 401 p.
Conway Morris, S., and Bengtson, S., 1994, Cambrian predators; Possible evidence from boreholes: Journal of Paleontology, v. 68, p. 1–23.
Cotteau, G.H., 1895, Echinides recueillis par M. Lovisato dans le Miocène de Sardaigne. Mémoires de la Societe Geologique de France: Palaeontologie, Mémoire, v. 13, p. 1–56.
Cottreau, J., 1914, Les Échinides néogènes du Bassin Méditerranéen: Annales de l’Institut Océanographique Monaco, v. 6, p. 1–192.
Deline, B., 2008, The first evidence of predatory or parasitic drilling in stylo- phoran echinoderms: Acta Palaeontologica Polonica, v. 53, p. 739–743.
Föllmi, K.B., Gertsch, B., Renevey, J.-P., de Kaenel, E., and Stille, P., 2008, Stratigraphy and sedimentology of phosphate-rich sediments in Malta and south-eastern Sicily (latest Oligocene to early late Miocene): Sedimentol- ogy, v. 55, p. 1029–1051.
Foresi, L.M., Verducci, M., Baldassini, N., Lirer, F., Mazzei, R., Salvatorini, G., Ferraro, L., and Da Prato, S., 2011, Integrated stratigraphy of St. Peter’s Pool section (Malta): New age for the Upper Globigerina Limestone Member and progress towards the Langhian GSSP: Stratigraphy, v. 8, p. 125–143.
Ghiold, J., 1982, Observations on the clypeasteroid Echinocyamus pusillus (O.F. Müller): Journal of Experimental Marine Biology and Ecology, v. 61, p. 57–74.
Giannelli, L., and Salvatorini, G., 1972, I foraminiferi planctonici dei sedimenti terziari dell’Arcipelago maltese: Biostratigrafia del “Globigerina Lime- stone” I, Atti della Società Toscana di Scienza Naturali, Memorie, Serie A, v. 82, p. 1–24.
Gregory, J.W., 1891, The Maltese fossil Echinoidea and their evidence on the correlation of the Maltese rocks: Transactions of the Royal Society of Edinburgh, v. 36, p. 565–639.
Grun, T.B., and Nebelsick, J.H., 2015, Sneaky snails: How drill holes can affect paleontological analyses of the minute clypeasteroid echinoid Echinocya- mus, in Zamora, D., and Rábano, I., eds., Progress in Echinoderm Palaeo- biology: Madrid, Cuadernos del Museo Geominero, 19. Instituto Geológico y Minero de España, p. 71–74.
Grun, T., Sievers, D., and Nebelsick, J.H., 2014, Drilling predation on the clypeasteroid echinoid Echinocyamus pusillus from the Mediterranean Sea (Giglio, Italy): Historical Biology, v. 26, p. 745–757.
Gruszczyński, M., Marshall, J.D., Goldring, R., Coleman, M.L., Małkowski, K., Gaździcka, E., Semil, J., and Gatt, P., 2008, Hiatal surfaces from the
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