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View - Kowalewski, M. - Virginia Tech

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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

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