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PALEONTOLOGICAL SOCIETY PAPERS, V. 8, 2002spatial and environmental variation in drillingpredation in the middle Miocene of Central Europe.The sampling methodology allowed forcomparisons within provinces, between provinces(Boreal vs. Paratethys), and between facies (finegrainedvs. coarse-grained siliciclastics). Theyfound that unsuccessful and multiple drillholesoccurred more frequently in the Boreal provincethan the Paratethys province; the same facies alsoincluded molluscs with different drillingfrequencies—with as much as a three-folddifference between samples collected in adjacentsites from the same facies! They concludedunequivocally that spatial variation should beevaluated independently before any large-scaletemporal trends are inferred for predation. Clearlymultiple collections with emphasis on facies needto be included in the temporal analysis of escalatorypredation hypotheses.Shell ornamentation (spines).—Spinedevelopment requires extra amounts of calciumcarbonate in seawater, and shallow tropical marinewaters meet this requirement (Nichol, 1965;Stanley, 1970). Spines have inspired variedhypotheses concerning their function asantipredatory architecture (Table 5). Few studies,however, have focused on alternative hypothesessuch as whether spines are phylogenetic legaciesof shell building (as in cardiid bivalves), non-aptiveconstructional artifacts, exaptations, or adaptations.Few workers have endeavored to apply suchphilosophical rigor, and many have created adaptivescenarios. Another important possible function ofspines could be to increase surface area for thesettlement of epibionts, and for trapping debris thatcamouflages the shells (Vance, 1978; Feifarek, 1987;Stone, 1998). Stone (1998) has shown that spineson epifaunal bivalves deter the attack of muricidshell-drilling predators, but muricids can still borein areas where spines are absent. In the same study,spines were found not to deter predatory attacks bysea stars that engulf their prey. The rise of spinoseornamentation in bivalves predates the radiation ofthe predatory Muricidae in the Albian, and actuallyextends back to the late Paleozoic in thesuperfamily Pectinoidea (Stone, 1998).Molluscan conchiolin layers: Are theyantipredatory?—Conchiolin is the organiccomponent of molluscan shells composed ofproteins, polysaccharids, and glycosaminoglycans(Table 6) (Wilber and Simkiss, 1968; Gregoire,1972; Wainwright et al., 1982). The periostracumand non-calcareous operculae are composedchiefly of conchiolin, while the nacreous layersand other shell microstructures contain variousquantities of conchiolin. Thus, conchiolin has a veryold history, putatively stemming from the oldestshelled mollusc in the Cambrian Period. What ispuzzling, however, is that only a few groups ofmolluscs—chiefly the freshwater bivalves (e.g.,Margaritiferidae, Unionidae, Mutelidae), estuarineto marine bivalves (Corbiculidae and Solenidae),and a few marine species—have conchiolinrepresented as separate sheets within their shells(Taylor et al., 1969; Anderson, 1992; Harper, 1994).Conchiolin, as a protein, is thought to form at a highmetabolic cost to the organism—and, perhapsbecause of this, there appears to be an evolutionarytendency to lose conchiolin layers (references inKardon, 1998). Thus, there must be someevolutionary reason for maintaining conchiolin inmolluscan shells despite its high metabolic cost ofproduction (Table 6). It has long been hypothesizedthat the conchiolin sheets deterred predation bydrilling molluscan predators, such as naticid (Lewyand Samtleben, 1979) or muricid gastropods (Taylor,1970,1981), and most of the work done to test thishypothesis has focused on the corbulid bivalves(Fischer, 1963; Lewy and Samtleben, 1979; DeCauwer, 1985; Anderson et al., 1991; Anderson,1992; Harper, 1994). The corbulids (FamilyCorbulidae) are small, inequivalved bivalves with aglobose shell form, a single byssus thread, andshallow burrowing habits (e.g., Stanley, 1970). Theyfirst appeared in the Middle Jurassic, with thegreatest diversification taking place in theCretaceous and Eocene (e.g., Hallam, 1976).There appear to be three contrasting temporal“trends” related to whether conchiolin reducespredation. The first is that conchiolin doeseffectively reduce predation on corbulids throughtheir evolutionary history. Fischer (1963), for172
WALKER AND BRETT—POST-PALEOZOIC PATTERNS IN MARINE PREDATIONTABLE 5—Hypotheses for the origin of skeletal spines in marine invertebrates.Hypothesis Evidence Examples ReferenceSpines develop in calciumcarbonate supersaturated seawaters; most common in tropicsLess energetically costly to makespines in tropical watersSpondylus americanus Stanley, 1970Spines are antipredatoryPrimary spines that projectoutward may protect mantle edgeSpondylus americanus Logan, 1974Hollow spines and keels are forpelagic/planktonic existencesEconomy of mass Ammonoids Birkelund,1981Spines have no function Constructional artifact? Alternative hypothesisfor any invertebrateSpines function in filter-feeding Spines cover opening to animal PoricthophenidbrachiopodsCarter, 1967;Kauffman,Rudwick, 1970Spines are an ancestralcondition; phylogeneticconstraintSpines form in various ways, evenwithin closely related familiesCardiid bivalves;anomalodesmatansSchneider andCarter, 2001Spines vary withenvironmental conditionsof the substrateVarious spine types depending onsubstrate the larvae attach toSpondylus americanus(Jurassic–Recent)Logan, 1974Attachment to substrateSpines discourage epibiontsSpines acts as supports forsensory mantle tissue; "mantleoutposts" to give early warningsignals of dangerSpines serve a camouflagicfunction, breaking updistinctive outline of shellSpines stabilize the shell on ashifting substrateSpines act as attachmentmechanismsSpines and pedicellaria in someechinoderms discourage biontsettlement; perhaps barbedsecondary spines of Spondylusamericanus reduce biontsettlementCemented bivalves likeSpondylus whose rightvalve is attached tosubstrate; thishypothesis does notfunction for the left valveSea urchins; the bivalveSpondylus americanus-- Brachiopods (JurassicAcanthothirus )Hair-like barbed spines typical ofthe neanic stage of the left valve ofSpondylus which get covered withalgae and sediment; spines arethickly encrusted with epibointsLate Paleozoicproductoid brachiopod,Waagenoconcha;Spondylus americanusThe CretaceousSpondylus spinosusLogan, 1974Logan, 1974Rudwick, 1965;Logan, 1974Grant, 1966;Logan, 1974Logan, 1974;Carter, 1972Spines keep the feedingmargins of the shell abovethe substrate-- -- Logan, 1974173
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THE PALEONTOLOGICAL SOCIETYPAPERSVo
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The Paleontological Society Special
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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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