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KOWALEWSKI—ANALYTICAL METHODSTHE FOSSIL RECORD OF PREDATION:AN OVERVIEW OF ANALYTICAL METHODSMICHAL KOWALEWSKIDepartment of Geological Sciences, <strong>Virginia</strong> Polytechnic Institute and State University,Blacksburg, <strong>Virginia</strong> 24060 USAABSTRACT—Paleontological research on predation has been expanding rapidly in scope, methods, and goals. Thegrowing assortment of research strategies and goals has led to increasing differences in sampling strategies,types of data collected, definition of variables, and even reporting style. This methodological overview serves asa starting point for erecting some general methodological guidelines for studying the fossil record of predation.I focus here on trace fossils left by predators in the skeleton of their prey, arguably one of the most powerfulsources of direct data on predator-prey interactions available in the fossil record. A critical survey of samplingprotocols (data collecting strategy, sieve size, and sample size) and analytical approaches (predation intensitymetrics, strategies for evaluating behavioral selectivity of predators, and taphonomic tests) reveals that variousapproaches can be fruitful depending on logistic circumstances and scientific goals of paleoecological projects.Despite numerous caveats and uncertainties, trace fossils left by predators on skeletons of their prey remain oneof the most promising directions of research in paleoecology and evolutionary paleobiology.INTRODUCTIONIN RECENT YEARS, paleontological researchon predation has become increasingly sophisticatedin terms of complexity of tested hypotheses,intricacy of sampling designs, and quality ofanalytical methods. Moreover, its thematic scope hasexpanded abruptly as we now collect much morediverse data for a much broader spectrum oforganisms over a much wider range of observationalscales, from individual interactions to global-scalesecular trends. Unfortunately, albeit perhapsinevitably, our data are collected in various, oftendisparate ways, so our research efforts arecontaminated with methodologically undesirableidiosyncrasies. The irreconcilable differences insampling strategies, types of collected information,definition of variables, and even reporting stylemake it difficult to compare directly manyotherwise valuable data sets, and hamper metaanalyticalattempts to explore hoards of dataamassed in the rapidly growing literature on thefossil record of predation.The methodological overview presented in thischapter and the two subsequent contributionsincluded in this volume (Chin, 2002; Haynes, 2002)bring together a diversity of methods used forstudying the fossil record of predation. Thesereviews should help us in collecting and reportingfuture data in a more congruent manner so as toavoid the confusion that we often encounter whencommunicating our research.This chapter focuses primarily on trace fossilsfound on skeletons of marine invertebrate prey. Suchfossilized traces of predation provide arguably therichest source of quantifiable data on prey-predatorinteractions available in the fossil record (seeespecially Kitchell, 1986) and have been widely usedin paleontological research to date. Other importantlines of evidence for studying predator-preyinteractions are discussed here only briefly. Thesubsequent methodological contributions includedin this volume review the methods employed toinvestigate coprolites (Chin, 2002) and the distinctstrategies used to study hominids and othervertebrate predators and prey (Haynes, 2002).Following Bambach (2002), predators aredefined here as organisms that “…hunt or trap,3

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