Hostname: page-component-cd9895bd7-8ctnn Total loading time: 0 Render date: 2024-12-27T10:27:43.220Z Has data issue: false hasContentIssue false

Mass extinctions and macroevolution

Published online by Cambridge University Press:  08 April 2016

David Jablonski*
Affiliation:
Department of Geophysical Sciences, University of Chicago, 5734 South Ellis Avenue, Chicago, Illinois 60637. E-mail: [email protected]

Abstract

Mass extinctions are important to macroevolution not only because they involve a sharp increase in extinction intensity over “background” levels, but also because they bring a change in extinction selectivity, and these quantitative and qualitative shifts set the stage for evolutionary recoveries. The set of extinction intensities for all stratigraphic stages appears to fall into a single right-skewed distribution, but this apparent continuity may derive from failure to factor out the well-known secular trend in background extinction: high early Paleozoic rates fill in the gap between later background extinction and the major mass extinctions. In any case, the failure of many organism-, species-, and clade-level traits to predict survivorship during mass extinctions is a more important challenge to the extrapolationist premise that all macroevolutionary processes are simply smooth extensions of microevolution. Although a variety of factors have been found to correlate with taxon survivorship for particular extinction events, the most pervasive effect involves geographic range at the clade level, an emergent property independent of the range sizes of constituent species. Such differential extinction would impose “nonconstructive selectivity,” in which survivorship is unrelated to many organismic traits but is not strictly random. It also implies that correlations among taxon attributes may obscure causation, and even the focal level of selection, in the survival of a trait or clade, for example when widespread taxa within a major group tend to have particular body sizes, trophic habits, or metabolic rates. Survivorship patterns will also be sensitive to the inexact correlations of taxonomic, morphological, and functional diversity, to phylogenetically nonrandom extinction, and to the topology of evolutionary trees. Evolutionary recoveries may be as important as the extinction events themselves in shaping the long-term trajectories of individual clades and permitting once-marginal groups to diversify, but we know little about sorting processes during recovery intervals. However, both empirical extrapolationism (where outcomes can be predicted from observation of pre- or post-extinction patterns) and theoretical extrapolationism (where mechanisms reside exclusively at the level of organisms within populations) evidently fail during mass extinctions and their evolutionary aftermath. This does not mean that conventional natural selection was inoperative during mass extinctions, but that many features that promoted survivorship during background times were superseded as predictive factors by higher-level attributes. Many intriguing issues remain, including the generality of survivorship rules across extinction events; the potential for gradational changes in selectivity patterns with extinction intensity or the volatility of target clades; the heritability of clade-level traits; the macroevolutionary consequences of the inexact correlations between taxonomic, morphological, and functional diversity; the factors governing the dynamics and outcome of recoveries; and the spatial fabric of extinctions and recoveries. The detection of general survivorship rules—including the disappearance of many patterns evident during background times—demonstrates that studies of mass extinctions and recovery can contribute substantially to evolutionary theory.

Type
Macroevolutionary Patterns within and among Clades
Copyright
Copyright © The Paleontological Society 

Access options

Get access to the full version of this content by using one of the access options below. (Log in options will check for institutional or personal access. Content may require purchase if you do not have access.)

References

Literature Cited

Aberhan, M., and Baumiller, T. K. 2003. Selective extinction among Early Jurassic bivalves: a consequence of anoxia. Geology 31:10771080.Google Scholar
Adrain, J. M., and Westrop, S. R. 2000. An empirical assessment of taxic paleobiology. Science 289:110112.Google Scholar
Aguirre, J., Riding, R., and Braga, J. C. 2000. Late Cretaceous incident light reduction: evidence from benthic algae. Lethaia 33:205213.Google Scholar
Anstey, R. L. 1978. Taxonomic survivorship and morphologic complexity in Paleozoic bryozoan genera. Paleobiology 4:407418.Google Scholar
Anstey, R. L. 1986. Bryozoan provinces and patterns of generic evolution and extinction in the Late Ordovician of North America. Lethaia 19:3351.Google Scholar
Anstey, R. L., and Pachut, J. F. 1995. Phylogeny, diversity history, and speciation in Paleozoic bryozoans. Pp. 239284in Erwin, D. H. and Anstey, R. L., eds. New approaches to speciation in the fossil record. Columbia University Press, New York.Google Scholar
Anstey, R. L., Pachut, J. F., and Tuckey, M. E. 2003. Patterns of bryozoan endemism through the Ordovician–Silurian transition. Paleobiology 29:305328.2.0.CO;2>CrossRefGoogle Scholar
Balinski, A., Olempska, E., and Racki, G., eds. 2002. Biotic responses to the Late Devonian global events. Acta Palaeontologica Polonica 47:186404.Google Scholar
Bambach, R. K., and Knoll, A. H. 2001. Is there a separate class of “mass” extinctions? Geological Society of America Abstracts with Programs 33(6):A-141.Google Scholar
Bambach, R. K., Knoll, A. H., and Sepkoski, J. J. Jr. 2002. Anatomical and ecological constraints on Phanerozoic animal diversity in the marine realm. Proceedings of the National Academy of Sciences USA 99:68546859.Google Scholar
Bambach, R. K., Knoll, A. H., and Wang, S. C. 2004. Origination, extinction, and mass depletions of marine diversity. Paleobiology 30:522542.Google Scholar
Bannerjee, A., and Boyajian, G. 1996. Changing biologic selectivity in the Foraminifera over the past 150 m.y. Geology 24:607610.Google Scholar
Barron, J. A. 1985. Miocene to Holocene planktic diatoms. Pp. 763809in Bolli, H. M., Saunders, J. B., and Perch-Nielsen, K., eds. Plankton stratigraphy. Cambridge University Press, Cambridge.Google Scholar
Bennett, P. M., and Owens, I. P. F. 1997. Variation in extinction risk among birds: chance or evolutionary predisposition? Proceedings of the Royal Society of London B 264:401408.Google Scholar
Benton, M. J. 1987. Progress and competition in macroevolution. Biological Reviews 62:305338.Google Scholar
Benton, M. J. 1991. Extinction, biotic replacements, and clade interactions. Pp. 89102in Dudley, E. C., ed. The unity of evolutionary biology. Dioscorides Press, Portland, Oregon.Google Scholar
Benton, M. J., ed. 1993. The fossil record 2. Chapman and Hall, London.Google Scholar
Benton, M. J., and Twitchett, R. J. 2003. How to kill (almost) all life: the end-Permian extinction event. Trends in Ecology and Evolution 18:358365.Google Scholar
Brandon, R. N. 1982. The levels of selection. Pp. 315323in Asquith, P. and Nichols, T., eds. PSA 1982, Vol. 1. Philosophy of Science Association, East Lansing, Mich.Google Scholar
Brandon, R. N. 1988. The levels of selection: a hierarchy of interactors. Pp. 5171in Plotkin, H., ed. The role of behavior in evolution. MIT Press, Cambridge.Google Scholar
Brandon, R. N., Antonovics, J., Burian, R., Carson, S., Cooper, G., Davies, P. S., Horvath, C., Mishler, B. D., Richardson, R. C., Smith, K., and Thrall, P. 1994. Sober on Brandon on screening-off and the levels of selection. Philosophy of Science 61:475486.CrossRefGoogle Scholar
Brenchley, P. J., Marshall, J. D., and Underwood, C. J. 2001. Do all mass extinctions represent an ecological crisis? Evidence from the Late Ordovician. Geological Journal 36:329340.Google Scholar
Bretsky, P. W. 1973. Evolutionary patterns in the Paleozoic Bivalvia: documentation and some theoretical considerations. Geological Society of America Bulletin 84:20792096.2.0.CO;2>CrossRefGoogle Scholar
Brown, J. H. 1995. Macroecology. University of Chicago Press, Chicago.Google Scholar
Cardillo, M., and Bromham, L. 2001. Body size and risk of extinction in Australian mammals. Conservation Biology 15:14351440.Google Scholar
Chatterton, B. D. E., and Speyer, S. E. 1989. Larval ecology, life-history strategies, and patterns of extinction and survivorship among Ordovician trilobites. Paleobiology 15:118132.CrossRefGoogle Scholar
Copper, P. 2002. Reef development at the Frasnian/Famennian mass extinction boundary. Palaeogeography, Palaeoclimatology, Palaeoecology 181:2765.Google Scholar
D'Hondt, S., Donaghay, P., Zachos, J. C., Luttenberg, D., and Lindinger, M. 1998. Organic carbon fluxes and ecological recovery from the Cretaceous-Tertiary mass extinction. Science 282:276279.Google Scholar
Dommergues, J.-L., Laurin, B., and Meister, C. 1996. Evolution of ammonoid morphospace during the Early Jurassic radiation. Paleobiology 22:219240.Google Scholar
Dommergues, J.-L., Laurin, B., and Meister, C. 2001. The recovery and radiation of Early Jurassic ammonoids: morphologic versus palaeobiogeographical patterns. Palaeogeography, Palaeoclimatology, Palaeoecology 165:195213.CrossRefGoogle Scholar
Droser, M. L., Bottjer, D. J., Sheehan, P. M., and McGhee, G. R Jr. 2000. Decoupling of taxonomic and ecologic severity of Phanerozoic marine mass extinctions. Geology 28:675678.Google Scholar
Duncan, J. R., and Lockwood, J. L. 2001. Extinction in a field of bullets: a search for causes in the decline of the world's freshwater fishes. Biological Conservation 102:97105.Google Scholar
Eble, G. J. 1999. On the dual nature of chance in evolutionary biology and paleobiology. Paleobiology 25:7587.Google Scholar
Eble, G. J. 2000. Constrasting evolutionary flexibility in sister groups: disparity and diversity in Mesozoic atelostomate echinoids. Paleobiology 26:5679.Google Scholar
Eldredge, N. 1997. Extinction and the evolutionary process. Pp. 6073in Abe, T., Levin, S. A., and Higashi, M., eds. Biodiversity: an ecological perspective. Springer, Berlin.Google Scholar
Eldredge, N. 2003. The sloshing bucket: how the physical realm controls evolution. Pp. 332in Crutchfield, J. P. and Schuster, P., eds. Evolutionary dynamics. Oxford University Press, New York.Google Scholar
Erwin, D. H. 1989. Regional paleoecology of Permian gastropod genera, southwestern United States and the end-Permian mass extinction. Palaios 4:424438.Google Scholar
Erwin, D. H. 1993. The great Paleozoic crisis: life and death in the Permian. Columbia University Press, New York.Google Scholar
Erwin, D. H. 1996. Understanding biotic recoveries: extinction, survival, and preservation during the end-Permian mass extinction. Pp. 398418in Jablonski, , et al. 1996.Google Scholar
Erwin, D. H. 1998. The end and the beginning: recoveries from mass extinctions. Trends in Ecology and Evolution 13:344349.Google Scholar
Erwin, D. H. 2001. Lessons from the past: evolutionary impacts of mass extinctions. Proceedings of the National Academy of Sciences USA 98:53995403.Google Scholar
Erwin, D. H. 2003. Impact at the Permo-Triassic boundary: a critical evaluation. Astrobiology 3:6774.CrossRefGoogle ScholarPubMed
Erwin, D. H. 2004. Mass extinctions and evolutionary radiations. Pp. 218228in Moya, A. and Font, E., eds. Evolution from molecules to ecosystems. Oxford University Press, New York.Google Scholar
Erwin, D. H., Valentine, J. W., and Sepkoski, J. J. Jr. 1987. A comparative study of diversification events: the early Paleozoic versus the Mesozoic. Evolution 41:11771186.Google Scholar
Erwin, D. H., Bowring, S. A., and Jin, Y. G. 2002. End-Permian mass extinctions: a review. Geological Society of America Special Paper 356:363384.Google Scholar
Fara, E. 2000. Diversity of Callovian-Ypresian (Middle Jurassic-Eocene) tetrapod families and selectivity of extinctions at the K/T boundary. Geobios 33:387396.Google Scholar
Fara, E. 2001. What are Lazarus taxa? Geological Journal 36:291303.CrossRefGoogle Scholar
Fisher, D. O., Blomberg, S. P., and Owens, I. P. F. 2003. Extrinsic versus intrinsic factors in the decline and extinction of Australian marsupials. Proceedings of the Royal Society of London B 270:18011808.CrossRefGoogle ScholarPubMed
Flügel, E. 2002. Triassic reef patterns. In Kiessling, W., Flügel, E., and Golonka, J., eds. Phanerozoic reef patterns. SEPM Special Publication 72:391463.Google Scholar
Foote, M. 1993. Discordance and concordance between morphological and taxonomic diversity. Paleobiology 19:185204.Google Scholar
Foote, M. 1996. Models of morphological diversification. Pp. 6286in Jablonski, et al. 1996.Google Scholar
Foote, M. 1997. The evolution of morphological diversity. Annual Review of Ecology and Systematics 28:129152.Google Scholar
Foote, M. 1999. Morphological diversity in the evolutionary radiation of Paleozoic and post-Paleozoic crinoids. Paleobiology Memoirs No. 1. Paleobiology 25(Suppl. to No. 2).Google Scholar
Foote, M. 2003. Origination and extinction through the Phanerozoic: a new approach. Journal of Geology 111:125148.Google Scholar
Foote, M. 2005. Pulsed origination and extinction in the marine realm. Paleobiology 31:620.Google Scholar
Gaston, K. J. 2003. The structure and dynamics of geographic ranges. Oxford University Press, Oxford.Google Scholar
Gaston, K. J., and Blackburn, T. M. 1995. Birds, body size, and the threat of extinction. Philosophical Transactions of the Royal Society of London B 347:205212.Google Scholar
Gaston, K. J., and Blackburn, T. M. 1997. Age, area and avian diversification. Biological Journal of the Linnean Society 62:239253.Google Scholar
Gaston, K. J., and Blackburn, T. M. 2000. Pattern and process in macroecology. Blackwell Science, Oxford.CrossRefGoogle Scholar
Gaston, K. J., Quinn, R. M., Wood, S., and Arnold, H. R. 1996. Measures of geographic range size: the effects of sample size. Ecography 19:259268.Google Scholar
Gilinsky, N. L. 1994. Volatility and the Phanerozoic decline of background extinction intensity. Paleobiology 20:445458.Google Scholar
Gilinsky, N. L., and Bambach, R. K. 1987. Asymmetrical patterns of origination and extinction in higher taxa. Paleobiology 13:427445.CrossRefGoogle Scholar
Gould, S. J. 1985. The paradox of the first tier: an agenda for paleobiology. Paleobiology 11:212.Google Scholar
Gould, S. J. 1989. Wonderful life. W. W. Norton, New York.Google Scholar
Gould, S. J. 2002. The structure of evolutionary theory. Harvard University Press, Cambridge.Google Scholar
Grantham, T. A. 2004. Constaints and spandrels in Gould's Structure of Evolutionary Theory. Biology and Philosophy 19:2943.Google Scholar
Griffis, K., and Chapman, D. J. 1988. Survival of phytoplankton under prolonged darkness: implications for the Cretaceous-Tertiary darkness hypothesis. Palaeogeography, Palaeoclimatology, Palaeoecology 67:305314.Google Scholar
Hallam, A. 1981. The end-Triassic bivalve extinction event. Palaeogeography, Palaeoclimatology, Palaeoecology 35:144.Google Scholar
Hallam, A., and Wignall, P. B. 1997. Mass extinctions and their aftermath. Oxford University Press, Oxford.Google Scholar
Hansen, T. A. 1982. Modes of larval development in early Tertiary neogastropods. Paleobiology 8:367377.Google Scholar
Hansen, T. A. 1988. Early Tertiary radiation of marine molluscs and the long-term effects of the Cretaceous-Tertiary extinction. Paleobiology 14:3751.CrossRefGoogle Scholar
Hansen, T. A., Upshaw, B., Kauffman, E. G., and Gose, W. 1993. Patterns of molluscan extinction and recovery across the Cretaceous-Tertiary boundary in east Texas: report on new outcrops. Cretaceous Research 14:685706.Google Scholar
Hargraves, P. E., and French, F. W. 1983. Diatom resting spores: significance and strategies. Pp. 4968in Fryxell, G. A., ed. Survival strategies of the algae. Cambridge University Press, Cambridge.Google Scholar
Harper, D. A. T., and Rong, J.-Y. 2001. Palaeozoic brachiopod extinctions, survival and recovery: patterns within the rhynchonelliformeans. Geological Journal 36:317328.Google Scholar
Harries, P. J. 1999. Repopulations from Cretaceous mass extinctions: environmental and/or evolutionary controls? Geological Society of America Special Paper 332:345364.Google Scholar
Heard, S. B., and Mooers, A. Ø. 2000. Phylogenetically patterned speciation rates and extinction risks change the loss of evolutionary history during extinctions. Proceedings of the Royal Society of London B 267:613620.Google Scholar
Heard, S. B., and Mooers, A. Ø. 2002. Signatures of random and selective extinctions in phylogenetic tree balance. Systematic Biology 51:889898.Google Scholar
Heinberg, C. 1999. Lower Danian bivalves, Stevns Klint, Denmark: continuity across the K/T boundary. Palaeogeography, Palaeoclimatology, Palaeoecology 154:87106.Google Scholar
Hoffman, A. 1989. Arguments on evolution. Oxford University Press, New York.Google Scholar
Hotton, C. L. 2002. Palynology of the Cretaceous-Tertiary boundary in central Montana: evidence for extraterrestrial impact as a cause of the terminal Cretaceous extinctions. Geological Society of America Special Paper 361:473502.Google Scholar
Hubbard, A. E., and Gilinsky, N. L. 1992. Mass extinctions as statistical phenomena: an examination of the evidence using X2 tests and bootstrapping. Paleobiology 18:148160.Google Scholar
Hulsey, C. D., and Wainwright, P. C. 2002. Projecting mechanics into morphospace: disparity in the feeding system of labrid fishes. Proceedings of the Royal Society of London B 269:317326.Google Scholar
Hunt, G., Roy, K., and Jablonski, D. 2005. Heritability of geographic range sizes revisited. American Naturalist (in press).Google Scholar
Ives, I. R., and Cardinale, E. J. 2004. Food-web interactions govern the resistance of communities after non-random extinctions. Nature. 429:174177.Google Scholar
Jablonski, D. 1986a. Causes and consequences of mass extinctions: a comparative approach. Pp. 183229in Elliott, D. K., ed. Dynamics of extinction. Wiley, New York.Google Scholar
Jablonski, D. 1986b. Background and mass extinctions: the alternation of macroevolutionary regimes. Science 231:129133.Google Scholar
Jablonski, D. 1986c. Larval ecology and macroevolution of marine invertebrates. Bulletin of Marine Science 39:565587.Google Scholar
Jablonski, D. 1986d. Evolutionary consequences of mass extinctions. Pp. 313329in Raup, D. M. and Jablonski, D., eds. Patterns and processes in the history of life. Springer, Berlin.Google Scholar
Jablonski, D. 1987. Heritability at the species level: analysis of geographic ranges of Cretaceous mollusks. Science 238:360363.Google Scholar
Jablonski, D. 1989. The biology of mass extinction: a paleontological view. Philosophical Transactions of the Royal Society of London B 325:357368.Google Scholar
Jablonski, D. 1995. Extinction in the fossil record. Pp. 2544in May, R. M. and Lawton, J. H., eds. Extinction rates. Oxford University Press, Oxford.Google Scholar
Jablonski, D. 1996. Body size and macroevolution. Pp. 256289in Jablonski, et al. 1996.Google Scholar
Jablonski, D. 1998. Geographic variation in the molluscan recovery from the end-Cretaceous extinction. Science 279:13271330.Google Scholar
Jablonski, D. 2000. Micro- and macroevolution: scale and hierarchy in evolutionary biology and paleobiology. In Erwin, D. H., and Wings, S. L., eds. Deep time: Paleobiology's perspective. Paleobiology 26(Supplement to No. 4):1552.Google Scholar
Jablonski, D. 2001. Lessons from the past: evolutionary impacts of mass extinctions. Proceedings of the National Academy of Sciences USA 98:53935398.Google Scholar
Jablonski, D. 2002. Survival without recovery after mass extinctions. Proceedings of the National Academy of Sciences USA 99:81398144.Google Scholar
Jablonski, D. 2003. The interplay of physical and biotic factors in macroevolution. Pp. 235252in Rothschild, L. J. and Lister, A. M., eds. Evolution on planet Earth. Elsevier/Academic Press, Amsterdam / London.Google Scholar
Jablonski, D., and Raup, D. M. 1995. Selectivity of end-Cretaceous marine bivalve extinctions. Science 268:389391.Google Scholar
Jablonski, D., and Sepkoski, J. J. Jr. 1996. Paleobiology, community ecology, and scales of ecological pattern. Ecology 77:13671378.Google Scholar
Jablonski, D., and Valentine, J. W. 1990. From regional to total geographic ranges: testing the relationship in Recent bivalves. Paleobiology 16:126142.Google Scholar
Jablonski, D., Erwin, D. H., and Lipps, J. H. 1996. Evolutionary paleobiology. University of Chicago Press, Chicago.Google Scholar
Jeffery, C. H. 1997. Dawn of echinoid nonplankototrophy: coordinated shifts in development indicate environmental instability prior to the K-T boundary. Geology 25:991994.Google Scholar
Jeffery, C. H. 2001. Heart urchins at the Cretaceous/Tertiary boundary: a tale of two clades. Paleobiology 27:140158.Google Scholar
Jernvall, J., and Wright, P. C. 1998. Diversity components of impending primate extinctions. Proceedings of the National Academy of Sciences USA 95:1127911283.Google Scholar
Joachimski, M. M., Pancost, R. D., Freeman, K. H., Ostertag-Henning, C., and Buggisch, W. 2002. Carbon isotope geochemistry of the Frasnian-Famennian transition. Palaeogeography, Palaeoclimatology, Palaeoecology 181:91109.Google Scholar
Kidwell, S. M. 2005. Shell composition has no net impact on large-scale evolutionary patterns in mollusks. Science 307:914917.Google Scholar
Kidwell, S. M., and Holland, S. M. 2002. The quality of the fossil record: implications for evolutionary analyses. Annual Review of Ecology and Systematics 33:561588.Google Scholar
Kiessling, W., and Baron-Szabo, R. C. 2004. Extinction and recovery patterns of scleractinian corals at the Cretaceous-Tertiary boundary. Palaeogeography, Palaeoclimatology, Palaeoecology 214:195223.Google Scholar
Kitchell, J. A., Clark, D. L., and Gombos, A. M. 1986. Biological selectivity of extinction: a link between background and mass extinction. Palaios 1:504511.Google Scholar
Koch, C. F. 1987. Prediction of sample-size effects on the measured temporal and geographic distribution patterns of species. Paleobiology 13:100107.Google Scholar
Lee, M. S. Y., and Doughty, P. 2003. The geometric meaning of macroevolution. Trends in Ecology and Evolution 18:263266.Google Scholar
Lerosey-Aubril, R., and Feist, R. 2003. Early ontogeny of trilobites: implications for selectivity of survivorship at the end-Devonian crisis. Geological Society of America Abstracts with Programs 34(7):385.Google Scholar
Levinton, J. S. 1996. Trophic group and the end-Cretaceous extinction: did deposit feeders have it made in the shade? Paleobiology 22:104112.Google Scholar
Lewis, J., Harris, A. S. D., Jones, K. J., and Edmonds, R. L. 1999. Long-term survival of marine planktonic diatoms and dinoflagellates in stored sediment samples. Journal of Plankton Research 21:343354.Google Scholar
Lockwood, J. L., Russell, G. J., Gittleman, J. L., Daehler, C. C., McKinney, M. L., and Purvis, A. 2002. A metric for analyzing taxonomic patterns of extinction risk. Conservation Biology 16:11371142.Google Scholar
Lockwood, R. 2003. Abundance not linked to survival across the end-Cretaceous mass extinction: patterns in North American bivalves. Proceedings of the National Academy of Sciences USA 100:24782482.Google Scholar
Lockwood, R. 2004. The K/T event and infaunality: morphological and ecological patterns of extinction and recovery in veneroid bivalves. Paleobiology 30:507521.Google Scholar
Lupia, R. 1999. Discordant morphological disparity and taxonomic diversity during the Cretaceous angiosperm radiation: North American pollen record. Paleobiology 25:128.Google Scholar
MacLeod, N. 2002. Testing evolutionary hypotheses with adaptive landscapes: use of random phylogenetic-morphological studies. Mathematische Geologie 6:4555.Google Scholar
MacLeod, N. 2003a. The causes of Phanerozoic extinctions. Pp. 253277in Rothschild, L. J. and Lister, A. M., eds. Evolution on planet Earth. Elsevier/Academic Press, Amsterdam/London.Google Scholar
MacLeod, N. 2003b. [Review of] The structure of evolutionary theory. Palaeontological Association Newsletter 50:4046.Google Scholar
MacLeod, N., and 21 others. 1997. The Cretaceous-Tertiary biotic transition. Journal of the Geological Society, London 154:265292.Google Scholar
Marshall, C. R. 1991. Estimation of taxonomic ranges from the fossil record. In Gilinsky, N. L. and Signor, P. W., eds. Analytical paleontology. Short Courses in Paleontology 4:1938. Paleontological Society, Knoxville, Tenn.Google Scholar
McGhee, G. R. Jr. 1996. The Late Devonian mass extinction. Columbia University Press, New York.Google Scholar
McGhee, G. R. Jr. 1999. Theoretical morphology: the concept and its application. Columbia University Press, New York.Google Scholar
McGowan, A. J. 2002. A morphometric study of the effect of diversity crises on Triassic ammonoid evolution. Geological Society of America Abstracts with Programs 34(6):361.Google Scholar
McGowan, A. J. 2004a. Ammonoid taxonomic and morphologic recovery patterns after the Permian-Traissic. Geology 32:665668.Google Scholar
McGowan, A. J. 2004b. The effect of the Permo-Triassic bottleneck on Triassic ammonoid morphological evolution. Paleobiology 30:369395.2.0.CO;2>CrossRefGoogle Scholar
McKinney, M. L. 1995. Extinction selectivity among lower taxa: gradational patterns and rarefaction error in extinction estimates. Paleobiology 21:300313.Google Scholar
McKinney, M. L. 1997. Extinction vulnerability and selectivity: combining ecological and paleontological views. Annual Review of Ecology and Systematics 28:495516.Google Scholar
McRoberts, C. A. 2001. Triassic bivalves and the initial marine Mesozoic revolution: a role for predators? Geology 29:359362.Google Scholar
McRoberts, C. A., and Newton, C. R. 1995. Selective extinction among end-Triassic European bivalves. Geology 23:102104.Google Scholar
Melchin, M. J., and Mitchell, C. E. 1991. Late Ordovician extinction of the Graptoloidea. In Barnes, C. R. and Williams, S. H., eds. Advances in Ordovician geology. Geological Survey of Canada Paper 90–9:143156.Google Scholar
Miller, A. I. 1997a. Dissecting global diversity patterns: examples from the Ordovician Radiation. Annual Review of Ecology and Systematics 28:85104.Google Scholar
Miller, A. I. 1997b. A new look at age and area: the geographic and environmental expansion of genera during the Ordovician Radiation. Paleobiology 23:410419.Google Scholar
Miller, A. I. 1998. Biotic transitions in global marine diversity. Science 281:11571160.Google Scholar
Miller, A. I., and Foote, M. 2003. Increased longevities of post-Paleozoic marine genera after mass extinctions. Science 302:10301032.Google Scholar
Miller, A. I., and Mao, S. 1995. Association of orogenic activity with the Ordovician radiation of marine life. Geology 23:305308.Google Scholar
Miller, A. I., and Mao, S. 1998. Scales of diversification and the Ordovician Radiation. Pp. 288310in McKinney, M. L. and Drake, J. A., eds. Biodiversity dynamics. Columbia University Press, New York.Google Scholar
Miller, A. I., and Sepkoski, J. J. Jr. 1988. Modeling bivalve diversification: the effect of interaction on a macroevolutionary system. Paleobiology 14:364369.Google Scholar
Mitchell, C. E. 1990. Directional macroevolution of the diplograptacean graptolites: a product of astogenetic heterochrony and directed speciation. In Taylor, P. D. and Larwood, G. P., eds. Major evolutionary radiations. Systematics Association Special Volume 42:235264. Clarendon, Oxford.Google Scholar
Nee, S., and May, R. M. 1997. Extinction and the loss of evolutionary history. Science 278:692694.Google Scholar
Neige, P. 2003. Spatial patterns of disparity and diversity of the Recent cuttlefishes (Cephalopoda) across the Old World. Journal of Biogeography 30:11251137.Google Scholar
Newman, M. E. J., and Eble, G. J. 1999. Decline in extinction rates and scale invariance in the fossil record. Paleobiology 25:434439.Google Scholar
Novack-Gottshall, P. M., and Miller, A. I. 2003. Comparative geographic and environmental diversity dynamics of gastropods and bivalves. Paleobiology 29:576604.Google Scholar
Page, K. N., 1996. Mesozoic ammonoids in space and time. Pp. 755794in Landman, N. H., Tanabe, K., and Davis, R. A., eds. Ammonoid paleobiology. Plenum, New York.Google Scholar
Patzkowsky, M. E. 1995. A hierarchical branching model of evolutionary radiations. Paleobiology 21:440460.Google Scholar
Petchey, O. L., and Gaston, K. J. 2002. Extinction and the loss of functional diversity. Proceedings of the Royal Society of London B 269:17211727.Google Scholar
Peters, E. 1996. Prolonged darkness and diatom mortality. 2. Marine temperate species. Journal of Experimental Marine Biology and Ecology 207:4358.Google Scholar
Peters, S. E., and Foote, M. 2001. Biodiversity in the Phanerozoic: a reinterpretation. Paleobiology 27:583601.Google Scholar
Purvis, A., Jones, K. E., and Mace, G. M. 2000a. Extinction. BioEssays 22:11231133.Google Scholar
Purvis, A., Agapow, P.-M., Gittleman, J. L., and Mace, G. M. 2000b. Nonrandom extinction and the loss of evolutionary history. Science 288:328330.Google Scholar
Racki, G. 1999. Silica-secreting biota and mass extinctions: survival patterns and processes. Palaeogeography, Palaeoclimatology, Palaeoecology 154:107132.Google Scholar
Racki, G., and House, M. R., eds. 2002. Late Devonian biotic crisis: ecological, depositional and geochemical records. Palaeogeography, Palaeoclimatology, Palaeoecology 181:1374.Google Scholar
Raup, D. M. 1984. Evolutionary radiations and extinctions. Pp. 514in Holland, H. D. and Trendall, A. F., eds. Patterns of change in Earth evolution. Springer, Berlin.Google Scholar
Raup, D. M. 1985. Mathematical models of cladogenesis. Paleobiology 11:4252.Google Scholar
Raup, D. M. 1991a. A kill curve for Phanerozoic marine species. Paleobiology 17:3748.Google Scholar
Raup, D. M. 1991b. Extinction: bad genes or bad luck? Norton, New York.Google Scholar
Raup, D. M. 1991c. The future of analytical paleontology. In Gilinsky, N. L. and Signor, P. W., eds. Analytical paleontology. Short Courses in Paleontology 4:207216. Paleontological Society, Knoxville, Tenn.Google Scholar
Raup, D. M. 1994. The role of extinction in evolution. Proceedings of the National Academy of Sciences USA 91:67586763.Google Scholar
Raup, D. M. 1996. Extinction models. Pp. 419433in Jablonski, et al. 1996.Google Scholar
Raup, D. M., and Boyajian, G. E. 1988. Patterns of generic extinction in the fossil record. Paleobiology 14:109125.Google Scholar
Raup, D. M., and Sepkoski, J. J. Jr. 1982. Mass extinctions in the marine fossil record. Science 215:15011503.Google Scholar
Rickards, R. B., and Wright, A. J. 2002. Lazarus taxa, refugia and relict faunas: evidence from graptolites. Journal of the Geological Society, London 159:14.Google Scholar
Robeck, H. E., Maley, C. C., and Donoghue, M. J. 2000. Taxonomy and temporal diversity patterns. Paleobiology 26:171187.Google Scholar
Robertson, D. B. R., Brenchley, P. J., and Owen, A. W. 1991. Ecological disruption close to the Ordovician-Silurian boundary. Historical Biology 5:131144.Google Scholar
Rode, A. L., and Lieberman, B. S. 2004. Using GIS to unlock the interactions between biogeography, environment, and evolution in Middle and Late Devonian brachiopods and bivalves. Palaeogeography, Palaeoclimatology, Palaeoecology 211:345359.Google Scholar
Roy, K. 1996. The roles of mass extinction and biotic interaction in large-scale replacements: a reexamination using the fossil record of stromboidean gastropods. Paleobiology 22:436452.Google Scholar
Roy, K., and Foote, M. 1997. Morphological approaches to measuring biodiversity. Trends in Ecology and Evolution 12:277281.Google Scholar
Roy, K., Balch, D. P., and Hellberg, M. E. 2001. Spatial patterns of morphological diversity across the Indo-Pacific: analyses using strombid gastropods. Proceedings of the Royal Society of London B 268:25032508.Google Scholar
Roy, K., Jablonski, D., and Valentine, J. W. 2004. Beyond species richness: biogeographic patterns and biodiversity dynamics using other metrics of diversity. Pp. 151170in Lomolino, M. V. and Heaney, L. R., eds. Frontiers of biogeography. Sinauer, Sunderland, Mass.Google Scholar
Russell, G. J., Brooks, T. M., McKinney, M. L., and Anderson, G. C. 1998. Present and future taxonomic selectivity in bird and mammal extinctions. Conservation Biology 12:13651376.Google Scholar
Saunders, W. B., Work, D. M., and Nikolaeva, S. V. 1999. Evolution of complexity in Paleozoic ammonoid sutures. Science 286:760763.Google Scholar
Sepkoski, J. J. Jr. 1984. A kinetic model of Phanerozoic taxonomic diversity. III. Post-Paleozoic families and mass extinctions. Paleobiology 10:246267.Google Scholar
Sepkoski, J. J. Jr. 1991a. Diversity in the Phanerozoic oceans: a partisan review. Pp. 210236in Dudley, E. C., ed. The unity of evolutionary biology. Dioscorides Press, Portland, Ore.Google Scholar
Sepkoski, J. J. Jr. 1991b. A model of onshore-offshore change in faunal diversity. Paleobiology 17:5877.Google Scholar
Sepkoski, J. J. Jr. 1996. Patterns of Phanerozoic extinction: a perspective from global data bases. Pp. 3551in Walliser, O. H., ed. Global events and event stratigraphy. Springer, Berlin.Google Scholar
Sepkoski, J. J. Jr. 1999. Rates of speciation in the fossil record. Pp. 260282in Magurran, A. E. and May, R. M., eds. Evolution of biological diversity. Oxford University Press, Oxford.Google Scholar
Sepkoski, J. J. Jr. 2002. A compendium of fossil marine animal genera. Bulletins of American Paleontology 363:1560.Google Scholar
Sepkoski, J. J. Jr., and Kendrick, D. C. 1993. Numerical experiments with model monophyletic and paraphyletic taxa. Paleobiology 19:168184.Google Scholar
Sepkoski, J. J. Jr., and Koch, C. F. 1996. Evaluating paleontologic data relating to bio-events. Pp. 2134in Walliser, O. H., ed. Global events and event stratigraphy. Springer, Berlin.Google Scholar
Sheehan, P. M. 2001. History of marine biodiversity. Geological Journal 36:231249.Google Scholar
Sheehan, P. M., and Coorough, P. J. 1990. Brachiopod zoogeography across the Ordovician-Silurian boundary. Geological Society of London Memoir 12:181187.Google Scholar
Sheehan, P. M., Coorough, P. J., and Fastovsky, D. E. 1996. Biotic selectivity during the K/T and Late Ordovician extinction events. Geological Society of America Special Paper 307477–489.Google Scholar
Shipley, B. 2000. Cause and correlation in biology. Cambridge University Press, Cambridge.Google Scholar
Smith, A. B. 2001. Large-scale heterogeneity of the fossil record: implications for Phanerozoic biodiversity studies. Philosophical Transactions of the Royal Society of London B 356:117.Google Scholar
Smith, A. B., and Jeffery, C. H. 1998. Selectivity of extinction among sea urchins at the end of the Cretaceous Period. Nature 392:6971.Google Scholar
Smith, A. B., and Jeffery, C. H. 2000a. Changes in the diversity, taxic composition and life-history patterns of echinoids over the past 145 million years. Pp. 181194in Culver, S. J. and Rawson, P. F., eds. Biotic response to global change: the last 145 million years. Cambridge University Press, Cambridge.Google Scholar
Smith, A. B., and Jeffery, C. H. 2000b. Maastrichtian and Palaeocene echinoids: a key to world faunas. Special Papers in Palaeontology 63.Google Scholar
Smith, J. T., and Roy, K. 1999. Late Neogene extinctions and modern regional species diversity: analyses using the Pectinidae of California. Geological Society of America Abstracts with Programs 31(7):473.Google Scholar
Sokal, R. R., and Rohlf, F. J. 1995. Biometry, 3d ed.W. H. Freeman, San Francisco.Google Scholar
Solan, M., Cardinale, B. J., Dowling, A. L., Engelhardt, K. A. M., Rusenick, J. L., and Srivastava, D. S. 2004. Extinction and ecosystem function in the marine benthos. Science 306:11771180.Google Scholar
Sole, R. V., Montoya, J. M., and Erwin, D. H. 2002. Recovery after mass extinction: evolutionary assembly in large-scale biosphere dynamics. Philosophical Transactions of the Royal Society of London B 357:697707.Google Scholar
Stanley, G. D. 2003. The evolution of modern corals and their early history. Earth-Science Reviews 60:195225.Google Scholar
Stanley, S. M. 1990. Delayed recovery and the spacing of major extinctions. Paleobiology 16:401414.Google Scholar
Sterelny, K., and Griffiths, P. E. 1999. Sex and death: an introduction to the philosophy of biology. University of Chicago Press, Chicago.CrossRefGoogle Scholar
Steuber, T., Mitchell, S. F., Buhl, D., Gunter, G., and Kasper, H. U. 2002. Catastrophic extinction of Caribbean rudist bivalves at the Cretaceous-Tertiary boundary. Geology 30:9991002.Google Scholar
Stilwell, J. D. 2003. Patterns of biodiversity and faunal rebound following the K-T boundary extinction event in Austral Palaeocene molluscan faunas. Palaeogeography, Palaeoclimatology, Palaeoecology 195:319356.Google Scholar
Twitchett, R. J. 2001. Incompleteness of the Permian-Triassic fossil record: a consequence of productivity decline? Geological Journal 36:341353.Google Scholar
Valentine, J. W. 1980. Determinants of diversity in higher taxonomic categories. Paleobiology 6:397407.Google Scholar
Valentine, J. W. 1990. The macroevolution of clade shape. Pp. 128150in Ross, R. M. and Allmon, W. D., eds. Causes of evolution: a paleontological perspective. University of Chicago Press, Chicago.Google Scholar
Valentine, J. W. 1992. Lessons from the history of life. Pp. 1732in Grant, P. R. and Horn, H. S., eds. Molds, molecules and Metazoa: growing points in evolutionary biology. Princeton University Press, Princeton, N.J.Google Scholar
Valentine, J. W., and Jablonski, D. 1986. Mass extinctions: sensitivity of marine larval types. Proceedings of the National Academy of Sciences USA 83:69126914.Google Scholar
Valentine, J. W., and Walker, T. D. 1987. Extinctions in a model taxonomic hierarchy. Paleobiology 13:193207.Google Scholar
Valentine, J. W., Roy, K., and Jablonski, D. 2002. Carnivore / non-carnivore ratios in northeastern Pacific marine gastropods. Marine Ecology Progress Series 228:153163.Google Scholar
Van Valen, L. M. 1973. A new evolutionary law. Evolutionary Theory 1:130.Google Scholar
Van Valen, L. M. 1984. A resetting of Phanerozoic community evolution. Nature 307:5052.Google Scholar
Van Valen, L. M. 1985a. A theory of origination and extinction. Evolutionary Theory 7:133142.Google Scholar
Van Valen, L. M. 1985b. How constant is extinction? Evolutionary Theory 7:93106.Google Scholar
Van Valen, L. M. 1987. Comment [on “Phanerozoic trends in background extinction: Consequence of an aging fauna”]. Geology 15:875876.Google Scholar
Vermeij, G. J. 1995. Economics, volcanoes, and Phanerozoic revolutions. Paleobiology 21:125152.Google Scholar
von Euler, F. 2001. Selective extinction and rapid loss of evolutionary history in the bird fauna. Proceedings of the Royal Society of London B 268:127130.Google Scholar
Wagner, P. J. 1995. Testing evolutionary constraint hypotheses: examples with early Paleozoic gastropods. Paleobiology 21:248272.Google Scholar
Wagner, P. J. 1996. Testing the underlying patterns of active trends. Evolution 50:9901017.Google Scholar
Wagner, P. J. 1997. Patterns of morphological diversification among the Rostroconchia. Paleobiology 23:115150.Google Scholar
Wang, S. C. 2003. On the continuity of background and mass extinction. Paleobiology 29:455467.Google Scholar
Wignall, P. B., and Benton, M. J. 1999. Lazarus taxa and fossil abundance at times of biotic crisis. Journal of the Geological Society, London 156:453456.Google Scholar
Willis, J. C. 1922. Age and area. Cambridge University Press, Cambridge.Google Scholar
Wills, M. A. 2001. Morphological disparity: a primer. Pp. 55144in Adrain, J. M., Edegcombe, G. D., and Lieberman, B. S., eds. Fossils, phylogeny, and form. Kluwer Academic/Plenum, New York.Google Scholar
Wimsatt, W. C. 2000. Emergence as non-aggregativity and the biases of reductionisms. Foundations of Science 5:269297.Google Scholar
Wing, S. L. 2004. Mass extinctions in plant evolution. Pp. 6197in Taylor, P. D., ed. Mass extinctions in the history of life. Cambridge University Press, Cambridge.Google Scholar
Wootton, J. T. 2004. Markov chain models predict the consequences of experimental extinctions. Ecology Letters 7:653660.Google Scholar
Zavaleta, E. S., and Hulvey, K. B. 2004. Realistic species losses disproportionately reduce grassland resistance to biological invaders. Science 306:11751177.Google Scholar