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PALEONTOLOGICAL SOCIETY PAPERS, V. 8, 2002TABLE 1—Fossil record of parasites on marine invertebrates.Host ParasiteCategoryParasitebenefitHostdetrimentEvidence for durationof relationshipFirstoccurrenceLastoccurrenceReferencesarthropod fungi 3 nutrients ? fungal mass in coprolites Silurian Tertiary Taylor & Osborn, 1996belemnite acrothoracid barnacle 3 ? ? teardrop-shaped boring Jurassic E. Cretaceous Seilacher, 1968bivalve capulid gastropod 1 ? ? attachment, scars, borings L. Cretaceous Tertiary Hayami & Kanie, 1980bivalve cestode or trematode 3 ? ? pearls Triassic Tertiary Herdman, 1906bivalve Clionid sponge 3 ? ? borings Cretaceous Tertiary Barry et al., 1972bivalve polychaete or sipunculid 2 ? ? tube-like borings Tertiary Tertiary Savazzi, 1995bivalve trematode 3 ? ? pits TertiaryTertiary Boucot, 1990(venerid)(Miocene)blastoid platyceratid gastropod 1 nutrients nutrients attached, scars, borings Devonian Mississippian Baumiller, 1996brachiopod acrothoracid barnacle 2 ? ? elongate borings L. Devonian Pennsylvanian Rodriguez &Gutschick, 1977brachiopod polychaete orgastropod3 ? ? borings M. Ordovician Tertiary Baumiller et al., 1999;Leighton, 2001brachiopod uncertain 3 ? ? tube secretions Devonian Devonian MacKinnon & Biernat,1970; Chatterton, 1975brachiopod uncertain 3 ? ? calcareous partitions L. Silurian L. Silurian Boucot &McCutcheon, 1986bryozoan algae and fungi 3 ? ? microstructure Ordovician Tertiary Elias, 1966bryozoan hydroid/3 ? ? embedment structures LateLatePalmer & Wilson, 1988colonian ascidianOrdovician Ordovicianbryozoan serpulid polychaete 3 ? ? tube secretions UpperCretaceousU. Cretaceous Ehrhard Voight, 1955chitinozoa bacteria and fungi 3 ? ? borings Ordovician Mississippian Grahn, 1981;Sutherland, 1994conularid uncertain 3 ? ? pearls Pennsylvanian Pennsylvanian Babcock, 1990crinoid myzostomid annelid 2 ? ? galls, pits Ordovician Tertiary Welch, 1976;Brett, 1978crinoid platyceratid gastropod 1 nutrients nutrients attached, scars, borings M. Ordovician Permian Rollins &Brezinski, 1988crinoid uncertain 3 ? ? circular pits M. Ordovician Permian Brett, 1985cystoid platyceratid gastropod 1 nutrients nutrients attached, scars, borings M. Silurian M. Silurian Boucot, 1990decapod bopyrid isopod 2 nutrients soft tissues carapace deformation L. Jurassic Tertiary Glaessner, 1969decapod rhizocephalan 2 nutrients sex organs claw feminization/castration L. Jurassic? Tertiary Boucot, 1990echinoid barnacle 2 ? ? elongate borings L. Cretaceous Tertiary Madsen & Wolff, 1965;Marouf, 1999echinoid copepod 2 ? ? internal galls Jurassic Tertiary Solovjev, 1961; Mercier,1937; Marouf, 1999echinoid eumelid gastropod 2 nutrients soft tissues borings Cretaceous Tertiary Kier, 1981; Alekseev &Endelman, 1989echinoid gastropod 1 ? ? deformed spines Cretaceous Tertiary Tasnádi-Kubacska, 1962echinoid myzostomid annelid 2 ? ? galls Tertiary (Miocene) Tertiary Roman, 1952echinoid prosobranch gastropod 2 ? ? skeletal malformations E. Cretaceous Tertiary Boucot, 1990gastropod acrothoracid barnacle 2 ? ? elongate borings E. Devonian Pennsylvanian Baird et al, 1990(platyceratid)graptolite uncertain 3 ? ? closed blisters E. Ordovician L. Silurian Bates and Loydell, 2000graptolite uncertain 3 ? ? open-ended tubes E. Ordovician L. Silurian Bates and Loydell, 2000graptolite uncertain 3 ? ? tubothecae E. Ordovician E. Ordovician Conway Morris, 1981196
BAUMILLER AND GAHN—PARASITISM ON MARINE INVERTEBRATESheavily on two important reviews of parasitism:Conway Morris’ (1981) “Parasites and the fossilrecord,” and Boucot’s (1990) “Evolutionarypaleobiology of behavior and coevolution.”We evaluated the published examples in termsof several criteria (Table 1). In considering theidentity of the parasite, if the inferred parasite wasdirectly observable in association with the host, itwas assigned to “category 1.” If the identity of theparasite was inferred from a well-constrainedproxy, namely the trace fossil that it produced, weassigned it to “category 2.” In all other instances,the identity of the parasite was considered morespeculative and assigned to “category 3.”We also considered the evidence presented forthe benefit that was gained by the parasite and forthe detrimental effects on the host. Finally, dataused to infer a long-term duration of the associationwere noted.Results.—Our search produced a substantialnumber of inferred cases of parasitism (Table 1),and, as Figure 1 illustrates, reports of parasiticassociation are quite evenly distributed through thepost-Cambrian Phanerozoic. Although slightlylower frequencies characterize the Triassic andJurassic and higher frequencies the Cretaceous andTertiary, the coarse temporal and taxonomicresolution prevents us from assigning muchsignificance to these differences, though it is worthnoting that the pattern roughly mirrors Sepkoski’s(1984) marine diversity curve. The total numberof parasitic associations for any geologic periodrarely exceeds a dozen, and if we consider thoseassociations for which the identity of the parasiteis well-constrained, based on co-occurrence orcharacteristic trace fossils (categories 1 and 2 inFig. 1), the number is lower still.To illustrate our scheme, we consider thebivalve-capulid gastropod association reported byHayami and Kanie (1980). These authors reportedspecimens of Cretaceous capulid gastropodsattached to the valves of inoceramids. Since theidentity of the interacting organisms is known, thisrepresents “category 1” in our scheme. Hayami andKanie (1980) inferred that this interactionrepresented parasitism because it was ecologicallylong-lived, as evidenced by the presence ofattached specimens, and was analogous to themodern capulids that are found in association withbivalves, gastropods, brachiopods, and annelids.Extant capulids are suspension feeders and can befacultative semi-parasites or parasites: they canFIGURE 1—Temporal distribution of reported parasites on marine invertebrates. Black, stippled, andwhite bars represent categories 1, 2, and 3, respectively. Categories 1, 2, and 3 explained in text. (Datafrom Table 1.)197
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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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