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AUTECOLOGY AND THE FILLING OF ECOSPACE: KEY METAZOAN RADIATIONS
Authors:RICHARD K BAMBACH  REW M BUSH†  DOUGLAS H ERWIN
Institution:Department of Paleobiology, MRC-121, National Museum of Natural History, Smithsonian Institution, PO Box 37012, Washington, DC 20013-7012, USA;e-mails:;; Department of Ecology and Evolutionary Biology and Center for Integrative Geosciences, University of Connecticut, 75 North Eagleville Road, Unit 3043, Storrs, CT 06269, USA;e-mail:
Abstract:Abstract:  All possible combinations of six tiering positions in relation to the substratum/water interface, six motility levels and six feeding strategies define a complete theoretical ecospace of 216 potential modes of life for marine animals. The number of modes of life actually utilized specifies realized ecospace. Owing to constraints of effectiveness and efficiency the modern marine fauna utilizes only about half the potential number of modes of life, two-thirds of which (62 of 92) are utilized by animals with readily preserved, mineralized hard parts. Realized ecospace has increased markedly since the early evolution of animal ecosystems. The Ediacaran fauna utilized at most 12 modes of life, with just two practised by skeletal organisms. A total of 30 modes of life are recorded in the Early and Middle Cambrian, 19 of which were utilized by skeletal organisms. The other 11 are documented from soft-bodied animals preserved in the Chengjiang and Burgess Shale Konservat-Lagerstätten. The number of modes of life utilized by skeletal organisms increased by more than 50 per cent during the Ordovician radiation to a Late Ordovician total of 30. Between the Late Ordovician and the Recent the number of utilized modes of life has doubled again. The autecological and taxonomic diversity histories of the marine metazoa appear to be broadly parallel, and future studies of theoretical ecospace utilization should provide more detailed tests of pattern and process in the ecological history of the metazoa.
Keywords:theoretical ecospace  evolutionary constraint  mode of life  ecological complexity  tiering  motility  feeding
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