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PALEONTOLOGICAL SOCIETY PAPERS, V. 8, 2002end of the Permian Period, in which the PaleozoicInsect Fauna was replaced by the Modern InsectFauna through the loss of a high percentage of highrankedclades (Labandeira and Sepkoski, 1993).Among carnivores, this event affected onlypredators, as the other carnivore guilds presumablywere absent. The second event, at the Cretaceous-Paleogene boundary, is more surprising, givenprevious studies that failed to detect any family-levelsignal beyond background rates (Ponomarenko,1988; Jarzembowski, 1989; Labandeira andSepkoski, 1993). A recent examination, however,has taken the explicitly ecological approach ofevaluating insect-mediated leaf damage patterns inwell-preserved and diverse floras across the K-Tboundary in North Dakota (Labandeira et al., 2002).This study demonstrated a pronounced qualitativeand quantitative diminution of insect herbivory(especially among specialists) at the boundary,indicating a significant deterioration of the herbivoreportion of terrestrial food webs. Such a major trophicalteration would have induced extinctions, based onour present knowledge of food-web perturbationfrom both theoretical modeling and empirical fieldobservations (DeAngelis, 1992; Dunne et al., 2002).After the expansion of parasitoidism and parasitismduring the later Mesozoic, this sudden reduction inhost resource probably trophically affected majorcarnivore guilds in continental food chains.SUMMARYThe transfer of energy in terrestrial ecosystems,both in fresh water and on land, culminates in theprocess of carnivory or the consumption of animalsby animals. This consumption may be rapid as inthe case of predators and their prey, or it may beprolonged as in the case of parasitoids with theirrelatively small hosts, and parasites with theircomparatively large hosts. While these three typesof carnivory are essential to the structure of foodwebs in modern ecosystems, their detection in thefossil record presents challenges. Six types ofpaleobiological evidence can reveal the presenceof carnivory: taxonomic affiliation, structural andfunctional attributes (especially mouthparts andovipositors), organismic damage, gut contents,coprolites, and mechanisms of predationavoidance. This evidence typically occurs in wellpreserveddeposits.Twelve invertebrate phyla have adopted acontinental carnivorous existence, representingpredatory, parasitoid, and parasitic life-habits. Ofthese phyla, the most speciose, ecologically varied,and abundant are nematodes and arthropods, bothof which have representative members practicingeach of the three modes of carnivory. Thishyperdiversity is related, particularly in arthropods,to such features as small size, ecdysis, ovipositors,and mouthpart morphology. This latter can involveeither a buccal stylet apparatus or the varied, oftenco-opted multielement mouthpart structures thathave been modified into several functional modesfor obtaining nutrition—including chewing,piercing-and-sucking, and boring. In the terrestrialinvertebrate world, accommodationist strategiessuch as forms of carnivory that result in theprolonged survival of a host, are probably morecommon than the rapid dispatching of prey.Members of all twelve continental invertebratephyla are documented in the fossil record, andarthropods are first represented by predatorscoexisting with detritivores and herbivores. Theyhave their first appearances in the earliest terrestrialbiotas from the latest Silurian to Middle Devonian,but it is not until the Middle Jurassic to mid-Cretaceous, approximately 250 million years later,that parasitoids and parasites are recorded as fossils,even though their phylogenetic relationships mayindicate greater antiquity. The diversification ofthese two functional groups of carnivores addedconsiderable complexity to food webs and may havebeen associated with the origin of insect sociality asa means of warding off attack. Although insect massextinctions at the end of the Permian had a profoundeffect on terrestrial invertebrate predation, recentevidence indicates that an end-Cretaceous extinctionmay have had less profound but more integrativeconsequences on food webs by affecting dependentpredators, parasitoids, and parasites. Thus anintriguing aim for further study is to understand the238
LABANDEIRA—PREDATORS, PARASITOIDS, AND PARASITESfabric of terrestrial food webs in a world in whichpredators were the only carnivores, preceding theemergence of parasitoids and parasites.ACKNOWLEDGMENTSAppreciation is extended to Finnegan Marsh,who formatted the figures. I thank the followingpeople for reviews: Steven Hasiotis, EdJarzembowski, Elena Lukashevich, MihailMostovski, George Poinar, Jr., Alex Rasnitsyn, andDena Smith. This is contribution 92 of the Evolutionof Terrestrial Ecosystems consortium at the NationalMuseum of Natural History. This article is dedicatedto the deceased paleoentomologist VladimirZherikhin of the Paleontological Institute of theRussian Academy of Sciences (Moscow), asystematist and a pioneer in the use of phylogenetic,taxic, and associational data in insect paleoecology.REFERENCESABEL, O. 1935. Vorzeitliche Lebensspuren. Gustav Fischer, Jena, 644 p.ABELE, L. G., W. KIM, AND B. E. FELGENHAUER. 1989. Molecular evidence for inclusion of the phylum Pentastomidain the Crustacea. Molecular Biology and Evolution, 6:685–691.ABRAHAMSON, W. G., AND A. E. WEIS. 1997. The Evolutionary Ecology of a Tritrophic-level Interaction: Goldenrod,the Stem Gall maker and its Natural Enemies. Princeton University Press, Princeton, NJ.AGUINALDO, A. M. A, J. M. TURBEVILLE, L. S. LINFORD, M. C. RIVERA, J. R. GAREY, R. A. RAFF, AND J. A. LAKE.1997. Evidence for a clade of nematodes, arthropods and other moulting animals. Nature, 387:489–493.ALLISON, C. W. 1975. Primitive fossil flatworm from Alaska: new evidence bearing on ancestry of the Metazoa.Geology, 3:649–652.ALMOND, J. E. 1985. The Silurian-Devonian fossil record of the Myriapoda. Philosophical Transactions of theRoyal Society of London, B, 309:227–237.ANDRES, D. 1989. Phosphatisierte Fossilien aus dem unteren Ordoviz von Südschweden. BerlinerGeowissenschaftliche Abhandlungen A, 106:9–19.ARNETT, R. H., JR. 2000. American Insects, Second Edition. CRC Press, Boca Raton, FL, 1003 p.ASKEW, R. W. 1979. Parasitic Insects. Heinemann, London, 316 p.BACHOFEN-ECHT, A. 1934. Beobachtungen über im Bernstein vorkommende Spinnengewebe. Biologia Generalis,10:179–184.BALASHOV, Y. S. 1999. Evolution of haematophagy in insects and ticks. Entomological Review, 79:943–954.BANKS, H. P., AND B. J. COLTHART. 1993. Plant-animal-fungal interactions in Early Devonian trimerophytes fromGaspé, Canada. American Journal of Botany, 80:992–1001.BÄNZIGER, H. 1975. Skin-piercing blood-sucking moths I: ecological and ethological studies on Calpe eustrigata(Lepid., Noctuidae). Acta Tropica, 32:125–144.BARNES, R. S. K., P. CALOW, AND P. J. W. OLIVE. 1993. The Invertebrates: A New Synthesis, Second Edition.Blackwell, London, 488 p.BARTHEL, K. W., N. H. M. SWINBURNE, AND S. CONWAY MORRIS. 1994. Solnhofen: A Study in Mesozoic Palaeontology.Cambridge University Press, Cambridge, UK, 236 p.BECHLEY, G. 1996. Morphologische Untersuchungen am Flügelgeäder der Rezenten Libellen und derenStammgruppenvertreter (Insecta; Pterygota; Odonata). Petalura, 2:1–402.BECHLEY, G., A. NEL, X. MARTÍNEZ-DELCLÒS, E. JARZEMBOWSI, R. CORAM, D. MARTILL, G. FLECK, F. ESCUILLIÉ, M. M.WISSHA, AND M. MAISCH. 2001. A revision and phylogenetic study of Mesozoic Aeschnoptera, with descriptionof numerous new taxa (Insecta: Odonata: Anisoptera). Neue Paläontologische Abhandlungen, 4:1–219.BERTOLANI, R., AND D. GRIMALDI. 2000. A new eutardigrade (Tardigrada: Milnesiidae) in amber from the UpperCretaceous (Turonian) of New Jersey, p. 103–110. In D. A. Grimaldi (ed.), Studies on Fossils in Amber, withParticular Reference to the Cretaceous of New Jersey. Backhuys, Leiden.239
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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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