Hostname: page-component-78c5997874-4rdpn Total loading time: 0 Render date: 2024-11-17T16:09:44.021Z Has data issue: false hasContentIssue false

Fair Sampling of Taxonomic Richness and Unbiased Estimation of Origination and Extinction Rates

Published online by Cambridge University Press:  21 July 2017

John Alroy*
Affiliation:
Department of Biological Sciences, Faculty of Science, Macquarie University, Sydney, NSW 2109 Australia
Get access

Abstract

Paleobiologists are reaching a consensus that biases in diversity curves, origination rates, and extinction rates need to be removed using statistical estimation methods. Diversity estimates are biased both by methods of counting and by variation in the amount of fossil data. Traditional counts are essentially tallies of age ranges. Because these counts are distorted by interrelated factors such as the Pull of the Recent and the Signor-Lipps effect, counts of taxa actually sampled within intervals should be used instead. Sampling intensity biases can be addressed with randomized subsampling of data records such as individual taxonomic occurrences or entire fossil collections. Fair subsampling would yield taxon counts that track changes in the species pool size, i.e., the diversity of all taxa that could ever be sampled. Most of the literature has overlooked this point, having instead focused on making sample sizes uniform through methods such as rarefaction. These methods flatten the data, undersampling when true diversity is high. A good solution to this problem involves the concept of frequency distribution coverage: a taxon's underlying frequency is said to be “covered” when it is represented by at least one fossil in a data set. A fair subsample, but not a uniform one, can be created by drawing collections until estimated coverage reaches a fixed target (i.e., until a “shareholder quorum” is attained). Origination and extinction rates present other challenges. For many years they were thought of in terms of simple counts or ratios, but they are now treated as exponential decay coefficients of the kind featuring in simple birth-death models. Unfortunately, these instantaneous rates also suffer from counting method biases (e.g., the Pull of the Recent). Such biases can be removed by only examining taxa sampled twice consecutively, three times consecutively, or in the first and third of three intervals but not the second (i.e., two timers, three timers, and part timers). Two similar equations involving these counts can be used. Alternative methods of estimating diversity and turnover through extrapolation share some of the advantages of quorum subsampling and two-timer family equations, but it remains to be shown whether they produce precise and accurate estimates when applied to fossil data.

Type
Taxonomic Data
Copyright
Copyright © 2010 by 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

Alroy, J. 1996. Constant extinction, constrained diversification, and uncoordinated stasis in North American mammals. Palaeogeography, Palaeoclimatology, Palaeoecology, 127:285311.Google Scholar
Alroy, J. 1998. Equilibrial diversity dynamics in North American mammals. p. 232287 In McKinney, M. L. and Drake, J. A. (eds.), Biodiversity Dynamics: Turnover of Populations, Taxa, and Communities. Columbia University Press, New York.Google Scholar
Alroy, J. 1999. The fossil record of North American mammals: evidence for a Paleocene evolutionary radiation. Systematic Biology, 48:107118.CrossRefGoogle ScholarPubMed
Alroy, J. 2000a. Successive approximations of diversity curves: ten more years in the library. Geology, 28:10231026.Google Scholar
Alroy, J. 2000b. New methods for quantifying macroevolutionary patterns and processes. Paleobiology, 26:707733.2.0.CO;2>CrossRefGoogle Scholar
Alroy, J. 2002. How many named species are valid? Proceedings of the National Academy of Sciences, USA, 99:37063711.CrossRefGoogle ScholarPubMed
Alroy, J. 2008. Dynamics of origination and extinction in the marine fossil record. Proceedings of the National Academy of Sciences, USA, 105:1153611542.CrossRefGoogle ScholarPubMed
Alroy, J. 2009. Speciation and extinction in the fossil record of North American mammals. p. 301323 In Butlin, R., Bridle, J., and Schluter, D. (eds.), Speciation and Patterns of Diversity. Cambridge University Press, Cambridge, 346 p.Google Scholar
Alroy, J. 2010. The shifting balance of diversity among major marine animal groups. Science, 329:11911194.Google Scholar
Alroy, J. In press. Geographic, environmental, and intrinsic biotic controls on Phanerozoic marine diversification. Palaeontology.Google Scholar
Alroy, J. et al. 2001. Effects of sampling standardization on estimates of Phanerozoic marine diversification. Proceedings of the National Academy of Sciences, USA, 98:62616266.CrossRefGoogle ScholarPubMed
Alroy, J. et al. 2008. Phanerozoic trends in the global diversity of marine invertebrates. Science, 321:97100.Google Scholar
Bambach, R. K. 1977. Species richness in marine benthic habitats through the Phanerozoic. Paleobiology, 3:152167.Google Scholar
Bambach, R. K. 1999. Energetics in the global marine fauna: a connection between terrestrial diversification and change in the marine biosphere. Geobios, 32:131144.Google Scholar
Benton, M. J. 1995. Diversification and extinction in the history of life. Science, 268:5258.Google Scholar
Benton, M. J. 2008. How to find a dinosaur, and the role of synonymy in biodiversity studies. Paleobiology, 34:516533.Google Scholar
Berger, W. H., and Parker, F. L. 1970. Diversity of planktonic Foraminifera in deep-sea sediments. Science, 168, 13451347.Google Scholar
Bush, A. M., and Bambach, R. K. 2004. Did alpha diversity increase during the Phanerozoic? Lifting the veils of taphonomic, latitudinal, and environmental biases. Journal of Geology, 112:625642.CrossRefGoogle Scholar
Bush, A. M., Markey, M. J., and Marshall, C. R. 2004. Removing bias from diversity curves: the effects of spatially organized biodiversity on sampling standardization. Paleobiology, 30:666686.2.0.CO;2>CrossRefGoogle Scholar
Chiarucci, A., Bacaro, G., Rocchini, D., and Fattorini, L. 2008. Discovering and rediscovering the sample-based rarefaction formula in the ecological literature. Community Ecology, 9, 121123.Google Scholar
Colwell, R. K., and Coddington, J. A. 1994. Estimating terrestrial biodiversity through extrapolation. Philosophical Transactions of the Royal Society (Series B), 345:101118.Google ScholarPubMed
Connolly, S. R., and Miller, A. I. 2001. Joint estimation of sampling and turnover rates from databases: capture-mark-recapture methods revisited. Paleobiology, 27:751767.Google Scholar
Cormack, R. M. 1964. Estimates of survival from the sighting of marked animals. Biometrika, 51:429438.CrossRefGoogle Scholar
Darwin, C. 1859. On the Origin of Species. John Murray, London, 502 p.Google Scholar
Foote, M. 1988. Survivorship analysis of Cambrian and Ordovician trilobites. Paleobiology, 14:258271.CrossRefGoogle Scholar
Foote, M. 1994. Temporal variation in extinction risk and temporal scaling of extinction metrics. Paleobiology, 20:424444.CrossRefGoogle Scholar
Foote, M. 1999. Morphological diversity in the evolutionary radiation of Paleozoic and post-Paleozoic crinoids. Paleobiology, 25(suppl.):1115.CrossRefGoogle Scholar
Foote, M. 2000. Origination and extinction components of taxonomic diversity: generaly problems. p. 74102 In Erwin, D. H. and Wing, S. L. (eds.), Deep Time: Paleobiology's Perspective. Paleobiology, 26(suppl.)Google Scholar
Foote, M. 2001. Inferring temporal patterns of preservation, origination, and extinction from taxonomic survivorship analysis. Paleobiology, 27:602630.Google Scholar
Foote, M. 2005. Pulsed origination and extinction in the marine realm. Paleobiology, 31:620.2.0.CO;2>CrossRefGoogle Scholar
Foote, M. 2007. Extinction and quiescence in marine animal genera. Paleobiology, 33:262273.Google Scholar
Foote, M., and Raup, D. M. 1996. Fossil preservation and the stratigraphic ranges of taxa. Paleobiology, 22:121140.CrossRefGoogle ScholarPubMed
Gilinsky, N. J., and Good, I. J. 1991. Probabilities of origination, persistence, and extinction of families of marine invertebrate life. Paleobiology, 17:145166.Google Scholar
Good, I. J. 1953. The population frequencies of species and the estimation of population. Biometrika, 40:237264.Google Scholar
Gotelli, N. J., and Colwell, R. K. 2001. Quantifying biodiversity: procedures and pitfalls in the measurement and comparison of species diversity. Ecology Letters, 4:379391.CrossRefGoogle Scholar
Gould, S. J., Raup, D. M., Sepkoski, J. J. Jr., Schopf, T. J. M., and Simberloff, D. S. 1977. The shape of evolution: a comparison of real and random clades. Paleobiology, 3:2340.Google Scholar
Harper, C. W. Jr. 1975. Standing diversity of fossil groups in successive intervals of geologic time: a new measure. Journal of Paleontology, 49:752757.Google Scholar
Hendy, A. J. W. 2009. The influence of lithification on Cenozoic marine biodiversity trends. Paleobiology, 35:5162.CrossRefGoogle Scholar
Kowalewski, M., Kiessling, W., Aberhan, M., Fürsich, F. T., Scarponi, D., Barbour Wood, S. L., and Hoffmeister, A. P. 2006. Ecological, taxonomic, and taphonomic components of the post-Paleozoic increase in sample-level species diversity of marine benthos. Paleobiology, 32:533561.Google Scholar
Kurtén, B. 1960. Chronology and faunal evolution of the earlier European glaciations. Commentationes Biologicae, Societas Scientarium Fennici, 21:4062.Google Scholar
Lyell, C. 1830. Principles of Geology. John Murray, London.Google Scholar
MacArthur, R. H., and Wilson, E. O. 1967. The Theory of Island Biogeography. Princeton University Press, New Jersey, 203 p.Google Scholar
Marshall, C. R., and Ward, P. D. 1996. Sudden and gradual molluscan extinctions in the latest Cretaceous of Western European Tethys. Science, 274:13601363.CrossRefGoogle ScholarPubMed
May, R. M. 1975. Patterns of species abundance and diversity. p. 81120 In Cody, M. L. and Diamond, J. E. (eds.), Ecology and Evolution of Communities. Belknap Press of Harvard University Press, Cambridge, Massachusetts, 545 p.Google Scholar
Miller, A. I. 2000. Conversations about Phanerozoic diversity. p. 5373 In Erwin, D. H. and Wing, S. L. (eds.), Deep Time: Paleobiology's Perspective. Paleobiology, 26(suppl.)Google Scholar
Miller, A. I. and Foote, M. 1996. Calibrating the Ordovician radiation of marine life: implications for Phanerozoic diversity trends. Paleobiology, 22:304309.Google Scholar
Newell, N. D. 1952. Periodicity in invertebrate evolution. Journal of Paleontology, 26:371385.Google Scholar
Newell, N. D. 1959. Adequacy of the fossil record. Journal of Paleontology, 33:488499.Google Scholar
Nichols, J. D., and Pollock, K. H. 1983. Estimating taxonomic diversity, extinction rates, and speciation rates from fossil data using capture-recapture models. Paleobiology, 9:150163.Google Scholar
Niklas, K. J., Tiffney, B. H., and Knoll, A. H. 1983. Patterns in vascular plant diversification. Nature, 303:614616.Google Scholar
Paul, C. R. C. 1982. The adequacy of the fossil record. p. 75117 In Joysey, K. A. and Friday, A. E. (eds.), Problems of Phylogenetic Reconstruction. Academic Press, New York, 442 p.Google Scholar
Phillips, J. 1860. Life on Earth: Its Origin and Succession. Macmillan, London, 224 p.Google Scholar
Pocock, M. J. O., Frantz, A. C., Cowan, D. P., White, P. C. L., and Searle, J. B. 2004. Tapering bias inherent in minimum number alive (MNA) population indices. Journal of Mammalogy, 85: 959962.Google Scholar
Preston, F. W. 1948. The commonness, and rarity, of species. Ecology, 29:254283.Google Scholar
Quental, T. B., and Marshall, C. R. 2009. Extinction during evolutionary radiations: reconciling the fossil record with molecular phylogenies. Evolution, 63:31583167.Google Scholar
Raup, D. M. 1972. Taxonomic diversity during the Phanerozoic. Science, 177:10651071.Google Scholar
Raup, D. M. 1975. Taxonomic diversity estimation using rarefaction. Paleobiology, 1:333342.Google Scholar
Raup, D. M. 1976. Species diversity in the Phanerozoic: an interpretation. Paleobiology, 2:289–279.Google Scholar
Raup, D. M. 1978. Cohort analysis of generic survivorship. Paleobiology, 4:115.Google Scholar
Raup, D. M. 1979. Biases in the fossil record of species and genera. Bulletin of the Carnegie Museum of Natural History, 13:8591.Google Scholar
Raup, D. M. 1985. Mathematical models of cladogenesis. Paleobiology, 11:4252.Google Scholar
Raup, D. M. 1991. A kill curve for Phanerozoic marine species. Paleobiology, 17:3748.Google Scholar
Raup, D. M., Gould, S. J., Schopf, T. J. M., and Simberloff, D. 1973. Stochastic models of phylogeny and the effect of diversity. Journal of Geology, 81:525542.CrossRefGoogle Scholar
Raup, D. M., and Sepkoski, J. J. Jr. 1982. Mass extinctions in the marine fossil record. Science, 215:15011503.CrossRefGoogle ScholarPubMed
Raup, D. M., and Sepkoski, J. J. Jr. 1984. Periodicity of extinctions in the geologic past. Proceedings of the National Academy of Sciences, USA, 81:801805.Google Scholar
Raup, D. M., and Stanley, S. M. 1971. Principles of Paleontology. W. H. Freeman, San Francisco, 388 p.Google Scholar
Rudwick, M. J. S. 1998. George Cuvier, Fossil Bones, and Geological Catastrophes: New Translations and Interpretations of the Primary Texts. University of Chicago Press, Chicago, 318 p.Google Scholar
Sanders, H. L. 1968. Marine benthic diversity: a comparative study. American Naturalist, 102:243282.CrossRefGoogle Scholar
Sepkoski, D. 2005. Stephen Jay Gould, Jack Sepkoski, and the ‘quantitative revolution’ in American paleobiology. Journal of the History of Biology, 38:209237.Google Scholar
Sepkoski, J. J. Jr. 1975. Stratigraphic biases in the analysis of taxonomic survivorship. Paleobiology, 1:343355.Google Scholar
Sepkoski, J. J. Jr. 1978. A kinetic model of Phanerozoic taxonomic diversity. I. Analysis of marine orders. Paleobiology, 4:223251.Google Scholar
Sepkoski, J. J. Jr. 1979. A kinetic model of Phanerozoic taxonomic diversity. II. Early Phanerozoic families and multiple equilibria. Paleobiology, 5:222251.Google Scholar
Sepkoski, J. J. Jr. 1982. A compendium of fossil marine families. Milwaukee Public Museum Contributions in Biology and Geology, 51:1125.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. 1990. The taxonomic structure of periodic extinction. Geological Society of America Special Paper, 247:3344.Google Scholar
Sepkoski, J. J. Jr. 1997. Biodiversity: past, present, and future. Journal of Paleontology, 71:533539.CrossRefGoogle ScholarPubMed
Sepkoski, J. J. Jr. 2002. A compendium of fossil marine animal genera. Bulletins of American Paleontology, 363:1560.Google Scholar
Sepkoski, J. J. Jr., Bambach, R. K., Raup, D. M., and Valentine, J. W. 1981. Phanerozoic marine diversity and the fossil record. Nature, 293:435437.Google Scholar
Sessa, J. A., Patzkowsky, M. E., and Bralower, T. J. 2009. The impact of lithification on the diversity, size distribution, and recovery dynamics of marine invertebrate assemblages. Geology, 37:115118.Google Scholar
Shinozaki, K. 1963. Note on the species area curve. Proceedings of the 10th Annual Meeting of the Ecological Society of Japan, 5.Google Scholar
Signor, P. W. III, and Lipps, J. H. 1982. Sampling bias, gradual extinction patterns, and catastrophes in the fossil record. Geological Society of America Special Publication, 190:291296.Google Scholar
Simpson, G. G. 1944. Tempo and Mode in Evolution. Columbia University Press, New York, 237 p.Google Scholar
Simpson, G. G. 1949. The Meaning of Evolution. Yale University Press, New Haven, Connecticut, 364 p.Google Scholar
Simpson, G. G. 1952. Periodicity in vertebrate evolution. Journal of Paleontology, 26:359370.Google Scholar
Simpson, G. G. 1960. The history of life. p. 117180 In Tax, S. (ed.), Evolution After Darwin. Volume 1: The Evolution of Life. University of Chicago Press, Chicago, 629 p.Google Scholar
Smith, E. P., Stewart, P. M., and Cairns, J. Jr. 1985. Similarities between rarefaction methods. Hydrobiologia, 120:167170.Google Scholar
Stanley, S. M. 1973. Effects of competition on rates of evolution, with special reference to bivalve molusks and mammals. Systematic Zoology, 22:486506.Google Scholar
Stanley, S. M. 1975. A theory of evolution above the species level. Proceedings of the National Academy of Sciences, USA, 72:646650.Google Scholar
Tarver, J. E., Braddy, S. J., and Benton, M. J. 2007. The effects of sampling bias on Paleozoic faunas and implications for macroevolutionary studies. Palaeontology, 50:177184.Google Scholar
Tipper, J. C. 1979. Rarefaction and rarefiction: the use and abuse of a method in paleoecology. Paleobiology, 5:423434.CrossRefGoogle Scholar
Valentine, J. W. 1970. How many marine invertebrate fossil species? A new approximation. Journal of Paleontology, 44:410415.Google Scholar
Van Valen, L. 1973. A new evolutionary law; Evolutionary Theory, 1:130.Google Scholar
Webb, S. D. 1969. Extinction-origination equilibria in late Cenozoic land mammals of North America. Evolution, 23:688702.Google Scholar