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PALEONTOLOGICAL SOCIETY PAPERS, V. 8, 2002Paleogene marine revolution in the molluscanrealm, because of the increased abundance ofdrilling neogastropods, the first real records ofdurophagous stomatopods and decapod crustaceans,and the evolution of specialized bird and mammalianpredators. However, most of these durophagousgroups are generalists, and it may be that they hada diffuse effect on their invertebrate prey.Finally, several new groups of carnivorousmarine mammals and birds originated in theMiocene. Walruses, gray whales, and humans arosein the Neogene and affected the coastal hard-shelledbiota in the areas where they foraged or settled. Thus,a Neogene phase of further predator intensificationis also suggested. However, there is no directevidence that prey were selectively affected (exceptfor the widespread decimation of species by humansand their alteration of marine habitats).The Cenozoic record seems to provide anexcellent window into predation and its effects, butfew have examined the temporal trends in predationduring this time (except for naticid molluscandrillers; e.g., Hansen et al., 1999; Kelley et al., 2001).Given that many predators leave their signature onshells and other prey, it is just a matter of reexaminingthe fossil record with the specific intentto look for predation. More work needs to be donein this area, especially on drilling records fromother gastropod groups, and on putative shell repairrecords that allow a comparison of Paleogene withNeogene localities. Additionally, in the Cenozoic,vast deep-sea (bathyal and deeper) fossil depositsof molluscs are well preserved in uplifted terracesin tectonically active regions of the world, allowingfor comparisons of predation (shell repair, shelldrilling) between shallow benthic and deep seafossil assemblages (Walker and Voight, 1994;Walker et al., 2002).Although escalation is sometimes cast as anongoing “arms race,” in actuality the predatoryrecord shows episodes of abrupt bioticreorganization during and after mass extinctions,punctuating longer interludes of relative stability(Brett et al., 1996). Some clades may retain thehistorical legacy of the Paleozoic predatoryrevolutions, as could be argued for the stalkedcrinoids in modern oceans; other clades maycontinuously evolve new predators, as Vermeij(1987) has argued based on the gastropod fossilrecord. Schneider and Carter (2001) show thatcardiid spine forms in Mesozoic and Cenozoicgroups appear to be a Paleozoic ancestral condition,and appear not to be related to the putativeMesozoic Marine Revolution. Thus, a clade-bycladeanalysis of predation would be most useful,as the different groups each have their ownevolutionary histories and ecological constraints.This review shows that not all morphology inbenthic organisms need be directly related topredation. Additionally, most durophagous predatorsdo not prey specifically on molluscs. They also preyon hard-shelled crustaceans, a major group oforganisms deemed to have caused selective pressuretoward escalated armor in gastropods (Vermeij,1987). We also must strive to examine predation inassemblages spatially across different environments,mindful of taphonomic bias, if we are to deriveevolutionarily and paleoecologically meaningfulinterpretations. The Phanerozoic record ofpredation is there, but it has not been fully explored;it is especially important to consider multipleworking hypotheses about Phanerozoic predationas we seek to interpret this record.Vermeij (1987) reviewed the record ofmolluscivorous predators, and their multifariousmethods of predation in the Phanerozoic. He madea plea for more data on the responses of prey speciesin Mesozoic and Cenozoic assemblages (Vermeij,1987, p. 239). Fifteen years later, his plea still stands.ACKNOWLEDGMENTSWe dedicate this second entrée to RichardBambach for his pioneering work concerningseafood through time. We greatly appreciateM.<strong>Kowalewski</strong> and P . Kelley for commissioningus to produce this synthesis. We thank L. Anderson,R. Feldmann, C. Hickman, A. Hoffmeister,G.Storrs, H. Greene, O. Rieppel, J. S. Pearse,R.Portell, and J. Voight for their thoughtfulcomments concerning numerous predation-relatedqueries. A. Hoffmeister, W. Miller III, and twoanonymous reviewers greatly improved the first178
WALKER AND BRETT—POST-PALEOZOIC PATTERNS IN MARINE PREDATIONdraft of this paper. E. Reitz graciously allowedaccess to her zooarchaeology collections at theGeorgia Museum of Natural History, Athens,Georgia. We also wish to thank Ruth Mawson andPeter Cockle, of Macquarie University, Australia,for facilitating the international editing of thismanuscript. This is not an exhaustive treatment,and any predatory comments regarding this paperneed to be addressed to the first author who willbristle with delight!REFERENCESADNET, S., AND H. CAPPETTA. 2001. A paleontological and phylogenetical analysis of squaliform sharks(Chondrichthyes: Squaliformes) based on dental characters. Lethaia, 34:234–248.AHYONG, S. T., AND C. HARLING. 2000. The phylogeny of the stomatopod Crustacea. Australian Journal of Zoology,48:607–642.ALLMON, W. D., J. C. NIEH, AND R. D. NORRIS. 1990. Drilling and peeling of turritelline gastropods since the LateCretaceous. Palaeontology, 33:595–611.ANDERSON, L. 1992. Naticid gastropod predation on corbulid bivalves: effects of physical factors, morphologicalfeatures, and statistical artifacts. Palaios, 7:602–620.ANDERSON, L. C., D. H. GEARY, R. H. NEHM, AND W. D. ALLMON. 1991. A comparative study of naticid gastropodpredation on Variborbula caloosae and Chione cancellata, Plio-Pleistocene of Florida, USA. Palaeogeography,Palaeoclimatology, Palaeoecology, 85:29–46.ARATA, A. A., AND J. H. HUTCHINSON. 1964. The Raccoon (Procyon) in the Pleistocene of North America. TulaneStudies in Geology, 2:21–27.ARONSON, R. B. 1987. Predation on fossil and Recent ophiuroids. Paleobiology, 13:187–192.ARONSON, R. B. 1988. Palatability of five Caribbean ophiuroids. Bulletin of Marine Science, 43:93–97.ARONSON, R. B. 1991. Predation, physical disturbance, and sublethal arm damage in ophiuroids: a Jurassic-Recentcomparison. Marine Ecology Progress Series, 74:91–97.ARONSON, R. B., D. B. BLAKE, AND T. OJI. 1997. Retrograde community structure in the late Eocene of Antarctica.Geology, 25:903–906.AUSICH, W. I. 1997. Regional encrinites; a vanished lithofacies, p. 509–519. In C. E. Brett and G. C. Baird, PaleontologicalEvents: Stratigraphic, Ecological and Evolutionary Implications. Columbia University Press, New York.AVERY, G., AND W. R. SIEGFRIED. 1980. Food gatherers along South Africa’s seashores. Oceans, 13:33–37.BALUK, W., AND A. RADWANSKI. 1996. Stomatopod predation upon gastropods from the Korytnica Basin, and fromother classical Miocene localities in Europe. Acta Geologica Polonica, 46:279–304BAMBACH, R. K., AND M. KOWALEWSKI. 1999. Diversity of predators compared to the records of prey-predator escalation:Two tales of the history of predation. Geological Society of America Abstracts with Programs, 31(7):A336.BARDET, N. 1992. Stratigraphic evidence for the extinction of ichthyosaurs. Terra Nova, 4:649–656.BARDET, N. 1994. Extinction events among Mesozoic marine mammals. Historical Biology, 7:313–324.BARNES, L. G. 1984. Whales, dolphins and porpoises: evolution of the Cetacea, p. 139–154. In P. D. Gingerichand C. E. Badgely (eds.), Mammals: Notes for a Short Course. University of Tennessee Department ofGeological Sciences, Studies in Geology, 8.BARNES, L. G., AND S. A. MCLEOD. 1984. The fossil record and phyletic relationships of gray whales, p. 3–32. In M. L.Jones, S. L. Swartz, and S. Leatherwood (eds.), The Gray Whale Escrichtius robustus. Academic Press, Orlando, FL.BAUMILLER, T. K., R. MOOI, AND C. G. MESSING. 1999. Predator-prey interactions between cidaroids and crinoids and theirecological and evolutionary implications. Geological Society of America Abstracts with Programs, 31:172–173.BELLWOOD, D. R. 1996. The Eocene fishes of Monte Bolca: the earliest coral reef assemblage. Coral Reefs, 15:11–19.BENTON, M. J. (ed.) 1993. The Fossil Record 2. Chapman and Hall, London, 845 p.BENTON, M. J. 1997. Vertebrate Palaeontology, 2 nd edition. Chapman and Hall, London, New York, 452 p.BERRY, S. S. 1956. A new west Mexican prosobranch mollusk parasitic on echinoids. American Midland Naturalist,56:355–357.179
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AuthorsRICHARD K. BAMBACHBotanical
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THE FOSSIL RECORD OF PREDATIONMicha
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INTRODUCTIONTHE FOSSIL RECORD OF PR
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KOWALEWSKI—ANALYTICAL METHODSTHE
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