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Previous work has suggested that escalation may have characterized the history of the naticid gastropod predator-prey system, based on apparent increases in drilling frequencies and the occurrence of antipredatory aptations among prey. We evaluate this hypothesis based on a comprehensive survey (over 40,000 specimens) of predation on molluscs from the Upper Cretaceous through lower Oligocene formations within the U.S. Gulf and Atlantic Coastal Plain. Patterns in drilling of both bivalve and gastropod prey are complex. Drilling frequencies were relatively low in the Cretaceous but increased sharply above the Cretaceous-Tertiary boundary, remaining high until the late Eocene. Following a significant decline near the Eocene-Oligocene boundary, drilling frequencies increased to a moderate level in the Oligocene. Contrary to our prediction based on the hypothesis of escalation, no temporal trend of increasing stereotypy of drillhole site occurred. However, significant increases in prey effectiveness (indicated by the incidence of incomplete drillholes and multiply bored shells) occurred between the Cretaceous and Oligocene. This pattern characterizes entire faunas as well as individual prey taxa that were consistently heavily drilled (turritellid gastropods and corbulid bivalves).

The fossil record is the primary source of data used to study predator-prey interactions in deep time and to evaluate key questions regarding the evolutionary and ecological importance of predation. Here, we review the types of paleontological data used to infer predation in the marine fossil record, discuss strengths and limitations of paleontological lines of evidence used to recognize and evaluate predatory activity, assess the influence of environmental gradients on predation patterns, and review fossil evidence for predator behavior and prey defense. We assembled a predation database from the literature that documents a steady increase in the number of papers on predation since the 1960s. These studies have become increasingly quantitative and have expanded in focus from reporting cases of predation documented by fossils to using the fossil record of predation to test ecological and evolutionary hypotheses. The data on the fossil record of predation amassed so far in the literature primarily come from trace fossils, mostly drill holes and, to a lesser extent, repair scars, derived predominantly from the Cenozoic of Europe and North America. Mollusks are the clade most often studied as prey and inferred predators. We discuss how to distinguish biotic from abiotic damage and predatory from parasitic traces, and how to recognize failed predation. Our data show that identifying the predator is easiest when predator and prey are preserved in the act of predation or when predators were fossilized with their gut contents preserved. However, determining the culprits responsible for bite traces, drill holes, and other types of predation traces can be more problematic.Taphonomic and other factors can distort patterns of predation, but their potential effects can be minimized by careful study design. With the correct identification and quantification of fossilized traces of predation, ecological trends in predator-prey interactions may be discerned along environmental gradients in water depth, habitat, and oxygen and nutrient availability.However, so far, these trends have not been explored adequately for the fossil record. We also

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drilling frequencies (mean = 30.3%, n = 4 samples) than did the Gulf Coast assemblages (mean = 18.0%, n = 11 samples; Mann-Whitney U-test, P < 0.01). This result is consistent with a trend found by some previous workers of an equatorward decrease in naticid drilling.

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Arms races between predators and prey may be driven by two related processes—escalation and coevolution. Escalation is enemy-driven evolution. In this top-down view of an arms race, the role of prey (with the exception of dangerous prey) is downplayed. In coevolution, two or more species change reciprocally in response to one another; prey are thought to drive the evolution of their predator, and vice versa. In the fossil record, the two processes are most reliably distinguished when the predator-prey system is viewed within the context of the other species that may influence the interaction, thus allowing for a relative ranking of the importance of selective agents. Detailed documentation of the natural history of living predator-prey systems is recommended in order to distinguish the processes in some fossil systems. A geographic view of species interactions and the processes driving their evolution may lead to a more diverse array of testable hypotheses on how predator-prey systems evolve and what constraints interactions impose on the evolution of organisms. Scale is important in evaluating the role of escalation and coevolution in the evolution of species interactions. If short-term reciprocal adaptation (via phenotypic plasticity or selection mosaics among populations) between predator and prey is a common process, then prey are likely to exert some selective pressure over their predators over the short term (on ecological time scales), but in the long run predators may still exert primary “top-down” control in directing evolution. On the scale of evolutionary time, predators of large effect likely control the overall directionality of evolution due to the inequalities of predator and prey in control of resources.