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Speciation and extinction asymmetries in paleontological phylogenies: evidence for evolutionary progress?

Published online by Cambridge University Press:  20 May 2016

Paul N. Pearson*
Affiliation:
Department of Earth Sciences, University of Bristol, Wills Memorial Building, Queens Road, Bristol, BS8 1RJ, UnitedKingdom. E-mail: [email protected]

Abstract

This paper concerns paleontological phylogenies that have a “budding” configuration, wherein “ancestral” species persist through branching events to coexist with their “descendants.” Two principal tests are proposed for detecting patterns within such trees. The first test, called the “ancestor-descendant extinction test,” compares the number of cases in which, after a split, the ancestral species became extinct before its descendant with the number of cases in which the descendant became extinct before its ancestor. The second test, called the “ancestor-descendant speciation test,” compares the number of cases in which, after a split, the ancestral species gave rise to a further species with the number of cases in which the descendant species gave rise to a further species. The null hypothesis in each case is that the frequencies are equal, as predicted by a random Markovian branching model of evolution.

Five stratophenetic species-level phylogenies of three taxonomic groups, planktonic foraminifera, nannofossils, and graptoloids, are examined using these tests, including one (Paleogene planktonic foraminifera) that is presented for the first time. In all cases, the phylogenetic trees are found to be strongly nonrandom. The general pattern, although by no means expressed perfectly in every case, corresponds to a Simpsonian “step-series,” in which ancestor taxa are simultaneously more likely to become extinct and less likely to speciate than their coexisting descendants. It is shown that this pattern cannot simply be the result of simple age-dependent factors such as an increasing extinction risk in older taxa. Instead, the very fact that a species has given rise to another appears to increase its future extinction risk and decrease its likelihood of further speciation.

Many possible biases may affect the shape of paleontological phylogenies, which are as yet poorly understood and unquantified. One potentially important effect follows from the taxonomic subdivision of gradual chronoclines into artificial morphospecies, such as might conceivably induce a step-series pattern in the phylogeny. Even if this is the partial or entire reason for the observed patterns, it would appear to imply directional evolution in phyletic gradualism. Other possible artifacts are discussed, but they are regarded as probably too weak to produce the observed patterns.

If the pattern is not artificial, the fact that three of the best known fossil groups exhibit substantial asymmetries in speciation and extinction argues against the currently popular “nonprogressive” view of evolution. Instead, the evolutionary step-series pattern is consistent with the classical Darwinian concept of the general competitive superiority of newly evolved species over their ancestors and supports the idea of evolutionary progress.

Type
Articles
Copyright
Copyright © The Paleontological Society 

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References

Literature Cited

Abel, O. 1912. Grundzüge der Palaeobiologie der Wirbeltiere. E. Schweizerbart'sche verlagsbuchhandlung Nägele und Dr. Sproesser. Stuttgart.Google Scholar
Arnold, A. J. 1983. Phyletic evolution in the Globorotalia crassaformis (Galloway and Wissler) lineage: a preliminary report. Paleobiology 9: 390397.Google Scholar
Aubry, M-P. 1989. Phylogenetically based calcareous nannofossil taxonomy: implications for the interpretation of geological events. Pp. 2140. in Crux, J. A., van Heck, A. eds. Nannofossils and their applications Ellis Horwood, London.Google Scholar
Barlow, C. 1994. Evolution extended: biological debates on the meaning of life. MIT Press Cambridge.Google Scholar
Bock, W. J. 1979. The synthetic explanation of macroevolutionary change—a reductionistic approach. Bulletin of the Carnegie Museum of Natural History 13: 5463.Google Scholar
Darwin, C. 1859. On the origin of species. John Murray London.Google Scholar
Eldredge, N. and Gould, S. J. 1972. Punctuated equilibrium: an alternative to phyletic gradualism. Pp. 82115. in Schopf, T. ed. Models in paleobiology Freeman, Cooper, San Francisco.Google Scholar
Foote, M. 1996. On the probability of ancestors in the fossil record. Paleobiology 22: 141151.Google Scholar
Fortey, R. A. 1985. Gradualism and punctuated equilibrium as competing and complimentary theories. Special Papers in Palaeontology 33: 1728.Google Scholar
Gingerich, P. D. 1976. Cranial anatomy and evolution of North American Plesiadapidae (Mammalia, Primates). University of Michigan Museum of Paleontology Papers in Paleontology. 15: 140.Google Scholar
Gingerich, P. D. 1990. Stratophenetics. Pp. 437442. in Briggs, D. E. G., Crowther, P. R. eds. Palaeobiology: a synthesis. Blackwell Scientific, Oxford.Google Scholar
Gould, S. J. 1988. Trends as changes in variance: a new slant on progress and directionality in evolution (Presidential Address). Journal of Paleontology 62: 319329.Google Scholar
Gould, S. J. 1990. Speciation and sorting as the source of evolutionary trends, or “things are seldom what them seem”. Pp. 327. in McNamara, K. J. ed. Evolutionary trends. Belhaven, London.Google Scholar
Gould, S. J. 1996. Life's grandeur: the spread of excellence from Plato to Darwin. Cape, London.Google Scholar
Grant, V. 1963. The origin of adaptations. Columbia University Press, New York.Google Scholar
Guyer, C. and Slowinski, J. B. 1991. Comparisons of observed phylogenetic topologies with null expectations among three monophyletic lineages. Evolution 45: 340350.Google Scholar
Guyer, C. and Slowinski, J. B. 1993. Adaptive radiation and the topology of large phylogenies. Evolution 47: 253263.Google Scholar
Haeckel, E. 1866. Generelle Morphologie der Organismen. Reimer, Berlin.Google Scholar
Haeckel, E. 1874. Anthropogenie; oder Entwickelungsgeschichte des Menschen. Engelmann, Leipzig.Google Scholar
Heard, S. B. 1992. Patterns in tree balance among cladistic phenetic and randomly generated phylogenetic trees. Evolution 46: 18181826.Google Scholar
Hennig, W. 1966. Phylogenetic systematics. University of Illinois Press, Urbana.Google Scholar
Huelsenbeck, J. P. and Kirkpatrick, M. 1996. Do phylogenetic methods produce trees with biased shapes? Evolution 50: 14181424.Google Scholar
Keith, A. 1934. Man's family tree. Watts, London.Google Scholar
Kellogg, D. E. 1975. The role of phyletic change in the evolution of Pseudocubus vema (Radiolaria). Paleobiology 1: 359370.Google Scholar
Kellogg, D. E. 1983. Phenology of morphometric change in radiolarian lineages from deep sea cores: implications for macroevolution. Paleobiology 9: 355362.Google Scholar
Kennett, J. P. and Srinivasan, M. S. 1983. Neogene planktonic foraminifera. Hutchinson Ross Stroudsberg Penn.Google Scholar
Kirkpatrick, M. and Slatkin, M. 1993. Searching for evolutionary patterns in the shape of a phylogenetic tree. Evolution 47: 11711181.Google Scholar
Malmgren, B. A. and Kennett, J. P. 1983. Phyletic gradualism in a late Cenozoic planktonic foraminiferal lineage DSDP Site 284 Southwest Pacific. Paleobiology 7: 230240>.Google Scholar
Mayr, E. 1974. Cladistic analysis or cladistic classification. Zeitschrift für Zoologische Systematik und Evolutionsforschung 12: 94128.Google Scholar
McKinney, M. L. 1990. Classifying and analysing evolutionary trends. Pp. 2858. in McNamara, K. J. ed. Evolutionary trends. Belhaven, London.Google Scholar
Mooers, A. O. and Heard, S. B. 1997. Inferring evolutionary processes from phylogenetic tree shape. Quarterly Review of Biology 72: 3154.Google Scholar
Mooers, A. O., Page, R. D. M., Purvis, A., and Harvey, P. H. 1995. Phylogenetic noise leads to unbalanced tree constructions. Systematic Biology 44: 332342.CrossRefGoogle Scholar
Norris, R. D. 1996. What is gradualism? Cryptic speciation in globorotaliid foraminifera. Paleobiology 22: 386405.Google Scholar
Olsson, R. H., Hemleben, C., Berggren, W. A., Huber, B. T. eds. In press. Atlas of Paleocene planktonic foraminifera Smithsonian Contributions to Paleobiology.Google Scholar
Page, R. D. M. 1991. Random dendrograms and null hypotheses in cladistic biogeography. Systematic Zoology 40: 5462.Google Scholar
Parker, W. C. and Arnold, A. J. 1997. Species survivorship in the Cenozoic planktonic foraminifera: a test of exponential and Weibull models. Palaios 12: 311.Google Scholar
Pearson, P. N. 1992. Survivorship analysis when extinction rates vary: the Paleogene planktonic foraminifera. Paleobiology 18: 115131.Google Scholar
Pearson, P. N. 1993. A lineage phylogeny for the Paleogene planktonic foraminifera. Micropaleontology 39: 193232.Google Scholar
Pearson, P. N. 1995. Investigating age-dependency of species extinction rates using dynamic survivorship analysis. Historical Biology 10: 119136.Google Scholar
Pearson, P. N., Shackleton, N. J., and Hall, M. A. 1997. Stable isotope evidence for the sympatric divergence of Globigerinoides trilobus and Orbulina universa (planktonic foraminifera). Journal of the Geological Society of London 154: 295302.Google Scholar
Perch-Nielsen, K. 1985a. Mesozoic calcareous nannofossils. Pp. 329426. in Bolli, H. M., Saunders, J. B., Perch-Nielsen, K. eds. Plankton stratigraphy. Cambridge University Press, Cambridge.Google Scholar
Perch-Nielsen, K. 1985b. Cenozoic calcareous nannofossils. Pp. 427554. in Bolli, H. M., Saunders, J. B., Perch-Nielsen, K. eds. Plankton stratigraphy. Cambridge University Press, Cambridge.Google Scholar
Raup, D. M. 1985. Mathematical models of cladogenesis. Paleobiology 11: 4252.Google Scholar
Raup, D. M. and Crick, R. E. 1981. Evolution of single characters in the Jurassic ammonite Kosmoceras. Paleobiology 7: 200215.Google Scholar
Raup, D. M. and Stanley, S. M. 1978. Principles of paleontology 2d edition. W. H. Freeman San Francisco.Google Scholar
Raup, D. M., Gould, S. J., Schopf, T. J. M., and Simberloff, D. S. 1973. Stochastic models of phylogeny and the evolution of diversity. Journal of Geology 81: 525542.Google Scholar
Rickards, R. B. 1977. Patterns of evolution in the graptolites. Pp. 333358. in Hallam, A. ed. Patterns of evolution as illustrated by the fossil record. Elsevier Amsterdam.Google Scholar
Rickards, R. B., Hutt, J. E., and Berry, W. B. N. 1977. Evolution of the Silurian and Devonian graptoloids. Bulletin of the British Museum of Natural History (Geology) 28: 1132.Google Scholar
Romein, A. J. T. 1979. Lineages in early Paleogene calcareous nannoplankton. Utrecht Micropaleontological Bulletin 22: 1231.Google Scholar
Ruse, M. 1996. Monad to Man: the concept of progress in evolutionary biology. Harvard University Press Cambridge.Google Scholar
Savage, H. M. 1983. The shape of evolution: systematic tree topology. Biological Journal of the Linnaean Society 22: 225244.CrossRefGoogle Scholar
Schluter, D. and McPhail, J. D. 1992. Ecologic character displacement and speciation in sticklebacks. American Naturalist 140: 85108.Google Scholar
Shao, K. and Sokal, R. R. 1990. Tree balance. Systematic Zoology 39: 266276.Google Scholar
Simberloff, D., Hecht, K. L., McCoy, E. D., and Connor, E. F. 1981. There have been no statistical tests of cladistic biogeographical hypotheses. Pp. 4063. in Nelson, G., Rosen, D. E. eds. Vicariance biogeography: a critique. Columbia University Press New York.Google Scholar
Simpson, G. G. 1953. The major features of evolution. Columbia University Press New York.Google Scholar
Simpson, G. G. 1974. The concept of progress in organic evolution. Social Research (spring): 2851.Google Scholar
Sorhannus, U., Fenster, E. J., Burckle, L. H., and Hoffman, A. 1988. Cladogenetic and anagenetic changes in the morphology of Rhizosolenia praebergonii Mukhina. Historical Biology 1: 185205.Google Scholar
Stanley, S. M. 1973. An explanation for Cope's Rule. Evolution 27: 126.Google Scholar
Stanley, S. M. 1979. Macroevolution: patterns and process. W. H. Freeman San Francisco.Google Scholar
Van Valen, L. 1973. A new evolutionary law. Evolutionary Theory 1: 130.Google Scholar
Wei, K-Y. and Kennett, J. P. 1983. Nonconstant extinction rates of Neogene planktonic foraminifera. Nature 305: 218220.CrossRefGoogle Scholar
Wei, K-Y. and Kennett, J. P. 1986. Taxonomic evolution of Neogene planktonic foraminifera and palaeoceanographic relations. Palaeoceanography 1: 6784.Google Scholar
Williams, G. C. 1992. Natural selection: domains levels and challenges. Oxford University Press New York.Google Scholar
Wood, H. E. 1941. Trends in Rhinoceros evolution. Transactions of the New York Academy of Science 383.Google Scholar