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PALEONTOLOGICAL SOCIETY PAPERS, V. 8, 2002borings of the type made by muricid gastropods.Drill holes of at least two distinctive typesoccur in middle to late Paleozoic shells (Ausichand Gurrola, 1979). These were termed Type A andType B boreholes and they are morphologicallysimilar to the later muricid and naticid gastropoddrill holes, respectively (see Figs. 4.3, 4.4). TypeA drill holes (Figs. 5.1, 5.2) are smaller, cylindrical,and may penetrate shells from below; they mayFIGURE 4—Traces of predation, in fossil and Recentshells. 1—Shell of Devonian gastropodPalaeozygopleura with sublethal healed fracture ofthe outer lip. 2—Permian bivalve shell with healedcrescentic fractures along valve margin; probably theresult of attempted predation by a fish. 3—Incompletebore hole of Polinices duplicata; note raised boss atcenter. 4—Complete drill hole of Natica severa; notebeveled outer margin. Redrawn from photographsin the following sources: 1, Brett and Cottrell (1982),2, Boyd and Newell (1972); 3, 4, Carriker andYochelson (1968). Figure modified from Brett (1992).record attacks by parasitic organisms (Ausich andGurrola, 1979). However, Type A drill holes occuras a single drill hole per shell and display a nonrandomstereotyped pattern characteristic ofcarnivores (Fig. 5.6) (Smith et al., 1985; Leighton,2001a, b). Cylindrical holes are known from LateOrdovician (Cincinnatian) brachiopods (Bucher,1938; Cameron, 1967). Some of these have provento be domichnial borings (Trypanites) made in deadshells (Carriker and Yochelson, 1969; Richards andShabica, 1969). But Kaplan and Baumiller (2000)argued recently that at least some of these holesshow non-random positioning, and hence wereprobably produced by predatory organisms.Rohr (1976) observed prey and site selectivityof small boreholes in Silurian orthid brachiopods.Liljedal (1985) also noted Type A borings insilicified Silurian bivalves. Similarly, Type Aborings occur in about 11% of the Early Devonianbrachiopod Discomyorthis and show evidence ofsize and site selectivity on the prey shells (Sheehanand Lespérance, 1978). Buehler (1969) reported alow frequency (2.25%) of cylindrical borings inMiddle Devonian shells, as did Rodriguez andGutschick (1970). However, the jury is still out onthe issue of whether these were predatory or merelyparasitic in nature (Leighton, 2001a, b).Type B boreholes are parabolic, 1–3 mm indiameter, and display a chamfer or bevel;incomplete boreholes possess a central raised knobor boss (see Figs. 4.3–5.5). These most closelyresemble drillings of modern naticid gastropods.These boreholes first become common in Devonianbrachiopods (Fenton and Fenton, 1931, 1932;Smith et al., 1985; <strong>Kowalewski</strong> et al., 1998); earlierpossible examples are known from Ordovicianbrachiopods (S. Felton, pers. comm.) but have notbeen documented in the literature. Brunton (1966)reports frequencies of up to 30% of brachiopodsdrilled with this type of hole in assemblages of lateCarboniferous age. Relatively few typical Type Bborings are reported from the Upper Carboniferousto Permian (see <strong>Kowalewski</strong> et al., 2000).However, a series of papers document small (< 2mm) boreholes with chamfering, which shouldperhaps be assigned to a third category; these occur100
BRETT AND WALKER—PREDATION IN PALEOZOIC MARINE ENVIRONMENTSFIGURE 5—Gastropod-like boreholes in brachiopods from the Middle Devonian Hamilton Group ofNew York State. Note scale bars. 1, 2—Type A borehole on the brachiopod Rhipidomella; 1 showsexterior view, 2, with negative reversed for comparison, shows blister of healed shell on interior of theshell. 3—Incomplete hole in Rhipidomella showing central boss. 4—Two overlapping sediment-filledboreholes on Parazyga; upper hole is complete; note chamfer (bevel) well displayed on lower hole.5—Incomplete borehole in Douvillina showing central raised boss; note truncated pseudopunctae ofshell. 6—positions of drill holes on the brachiopod Rhipidomella showing stereotypy of positioning overmain visceral mass of brachiopod. Modified from Smith et al. (1985).primarily in diminutive brachiopods belonging tothe family Cardiarinidae (Cooper, 1956; Bassett andBryant, 1993; Grant 1988; Morris, 1994). Hoffmeisteret al. (2001a) report drilling frequencies of up to33% in Cardiarina. The boreholes show stereotypywith respect to valves and preferred site on shells.Until recently, it was quite unclear whatorganisms were responsible for Type B boreholes,but the discovery of platyceratid gastropods in directassociation with this type of drill hole on LowerCarboniferous crinoids suggests that these snailswere among the culprits (Baumiller, 1990, 1996;Baumiller et al., 1999). Several studies have shownthat Type B hole-drillers display a distinct preferencefor particular prey taxa, notably athyrid and certainstrophomenid brachiopods. They also show101
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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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