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PALEONTOLOGICAL SOCIETY PAPERS, V. 8, 2002FIGURE 8—Traces of predation in fossil and Recent shells. 1, Shell of Cretaceous ammonitePlacenticeras with rows of punctures, probably made by a mosasaur. 2, Recent gastropod shell(Fasciolaria) exhibiting peeling damage inflicted by a callapid crab, ×1. 3, 4, Profile views of incompleteand complete gastropod drill holes in cross sections of bivalve shell: 3, cylindrical muricid drill holes;4, typical naticid holes; note parabolic cross section and central boss in incomplete borehole. 5, Locationsof most frequent drill holes in Recent bivalves from the Niger Delta. Redrawn from photographs in thefollowing sources: 1, Kauffman and Kesling (1960); 2, Bishop (1975); 3, Reyment (1971); 4, Sliter(1971). Figure modified from Brett (1990).with shark teeth embedded in bone, tooth marksslashed into the bone, or evidence of gastric-acidetching on putative prey items. Rothchild and Martin(1990) report on a shark tooth embedded in mosasaur(Clidastes) bone, which subsquently was repairedand ultimately caused spondylitus. Bite marks onmosasaurs without evidence of healing are alsoreported (Hawkins, 1990). Shark bite marks are alsoknown from elasmosaurid plesiosaur bones(Williston and Moodie, 1917; Welles, 1943).Late Cretaceous galeomorph selachian sharks(Squalicorax) are thought to have been scavengers142
WALKER AND BRETT—POST-PALEOZOIC PATTERNS IN MARINE PREDATIONpar excellence in the eastern Gulf Coastal Plainand Western Interior of the United States(Schwimmer et al., 1997). All living neoselachiansharks are carnivores, while galeomorph sharksmay prey on mollusks, crustaceans, andvertebrates; Squalicorax is also thought to scavengemany types of prey (see table 1 of Schwimmer, etal., 1997). Evidence of scavenging may includeembedded teeth which do not show evidence ofwound healing or tissue necrosis, and shark teethassociated with putatively scavenged remains(Schwimmer et al., 1997). Squalicorax, amoderately sized shark at 3.5 m, had serrateddentition of the “cutting type,” which may indicaterelatively diverse feeding strategies. IsolatedSqualicorax teeth, sometimes with associated bitemarks, were reported embedded in a decayedmosasaur vertebra and a juvenile hadrosaurmetatarsal (Schwimmer et al., 1997). Putative gutcontents from Squalicorax include mosasaur limbbones (Druckenmiller et al., 1993).Even fewer examples are known of sharkattacks on benthic invertebrates. Molluscan shells,such as those of inoceramid bivalves, are known tohave marginal edge fragmentation, and frequentlyare preserved as fragmented remains sometimesassociated with the putative shell-crushing shark,Ptychodus (Kauffman, 1972). One inoceramidspecimen, Inoceramus tenuis, is described as havingshell injuries on its left valve perhaps directlystemming from Ptychodus predation (Kauffman,1972). The right valve is uninjured, and Kauffman(1972) explains that this lack of injury is compatiblewith the life habits of the inoceramid, as the leftvalve was more exposed. Speden (1971) interpretsaggregations of fragmented inoceramid shells fromLate Cretaceous sites in New Zealand as evidenceof regurgitated or fecal material from vertebratepredators. Inoceramids are usually found in quiet,deep-water settings, either in chalks or black shales(Kauffman, 1972)—thus, any information on theirpotential predators would illuminate the littleknownpaleoecology of deep-water fauna in theLate Cretaceous.The batoid rays and skates first appeared in theEarly Jurassic and diversified during the LateJurassic and Cretaceous (Benton 1997; see alsoVermeij, 1987). These specialized elasmobranchswere adapted in large part for durophagous benthicpredation. Rays evolved stout hypermineralizedpavement plates for crushing hard-shelled prey, suchas crustaceans and molluscs (Fig. 9). Ray dentitionis thus similar to the pavement teeth of Devonianptyctodonts and rhenanids, and late Paleozoicholocephalans, Triassic–Jurassic semionotid fish,and Triassic placodonts. In all cases, crushing ofhard-shelled prey is inferred, but of these groupscertainly the batoid rays have been most successful.Many rays, exemplified by the cow nose rays, arecapable of excavating shallow pits in sandysubstrates in pursuit of infaunal bivalve, gastropod,polychaete, and other prey (see Fig. 12). Possibleancient ray pits have been reported from depositsas old as Late Cretaceous (Howard et al., 1977).Osteichthyans.—The neoselachian radiationsaw its counterpart in the Cretaceous osteichthyanteleost fishes (Fig. 5.1–5.3). The achievement ofimproved buoyancy via swim bladders,development of deep bodies, and anterior placementof pectoral and pelvic fins, represent coordinatedadaptations for increased swimming efficiency andmaneuverability during the Jurassic Period. Duringthe Late Cretaceous, the development of hingedmaxillae-premaxillae and highly protrusible mouthsfurther gave rise to a new mode of suctorial predatoryfeeding. These adaptations in turn fostered a majoradaptive radiation of neoteleost predators in the seaand in fresh water.In the Cretaceous, large basal teleosts clearlydominated in the intermediate- to large-sized fisheatingpredator guild. Many specimens of the largeXiphactinus (Fig. 5.1) from the Cretaceous of NorthAmerica have been found with ingested fish in thebody cavity. These include specimens from Kansaswith as many as ten fish in the stomach and a 4.5-meter specimen with a 1.6-meter relatedichthyodectid fish inside (Benton, 1997)! Specimensof the pavement-toothed Tribodus from theCretaceous Santana Formation of Brazil hadstomach contents that included shrimp andfragmentary molluscan shells (Maisey, 1996).Advanced acanthomorph teleosts evolved143
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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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