Self-regulation of trilobite diversity in Murero (middle Cambrian, Spain) due to compensatory extinction

Authors

  • J.S. LÓPEZ-VILLALTA Dpto. Biología y Geología, IES Pedro Simón Abril. Pº San Francisco 89, 02300 Alcaraz, Spain

DOI:

https://doi.org/10.1344/GeologicaActa2016.14.1.6

Keywords:

Extinction rate, Species turnover, Diversification, Macroevolution, Origination rate

Abstract

As species accumulate in a community, competition for available ecospace is expected to prevent the addition of new species and to facilitate species extinction, thus producing a dynamic equilibrium of diversity. This mechanism remains under debate since its empirical support comes mainly from indirect or partial evidence, with very few direct tests at the species level. Here a new method is described to detect the presence of selfregulation feedbacks between species richness and turnover rates. It consists of Monte Carlo simulations which randomize the distribution of species ranges among stratigraphic intervals, providing predictions which allow the detection of genuine self-regulation feedbacks in the real data. Since the simulations include any potential bias due to preservation, sampling, or change in depositional environment, and these biases would also affect the real dataset, they are thus ruled out as explanations for any difference found. This method is applied to one of the best known fossiliferous sequences worldwide, the Rambla de Valdemiedes in Murero (RV1 section, middle Cambrian, Spain), a classic locality that has been studied for more than 150 years and which stands out due to its excellent sampling density, continuous deposition, and homogeneous fossil preservation. The results show that trilobite species richness was self-regulated due to positive feedback with extinction rate, which implies that compensatory extinction regulated this fauna in spite of the on-going Cambrian radiation. The lack of evidence of any origination feedback suggests ecological opportunities were not limiting for new species to colonize this Cambrian community.

References

Alroy, J., 2008. Dynamics of origination and extinction in the marine fossil record. Proceedings of the National Academy of Sciences of the USA, 105, 11536–11542.

Alroy, J., 2014. Accurate and precise estimates of origination and extinction rates. Paleobiology, 40, 374-397.

Bambach, R.K., 1983. Ecospace utilization and guilds in marine communities through the Phanerozoic. In: Tevesz, M.J.S., McCall, P.L. (eds.). Biotic interactions in recent and fossil benthic communities. New York, Springer Science and Business Media, 719-746.

Bambach, R.K., Bush, A.M., Erwin, D.H., 2007. Autoecology and the filling of ecospace: key metazoan radiations. Palaeontology, 50, 1-22.

Brown, J.H., Ernest, S.M., Parody, J.M., Haskell, J.P., 2001. Regulation of diversity: maintenance of species richness in changing environments. Oecologia, 126, 321-332.

Cox, G.W., Ricklefs, R.E., 1977. Species diversity and ecological release in Caribbean land bird faunas. Oikos, 113-122.

Darwin, C., 1859. On the origins of species by means of natural selection. London, John Murray, 558pp.

De Verneuil, E., 1862. Descubrimiento de la fauna primordial en la provincia de Zaragoza. Revista Minera , 13, 479.

Dereims, A., 1898. Recherches géologiques dans le Sud de l’Aragon. Lille, Le Bigot Frères, 199pp.

Elton, C.S., 1958. The ecology of invasions by animals and plants. Chicago, University of Chicago Press, 181pp.

Foote, M., 2000. Origination and extinction components of taxonomic diversity: general problems. Paleobiology, 26 (4), 74-102.

Gaston, K.J., Spicer, J.I., 2009. Biodiversity: an introduction. Oxford, John Wiley & Sons, 208pp.

Gause, G.F., 1934. The struggle for existence. Baltimore, The Williams & Wilkins Company, 192pp.

Harmon, L.J., Harrison, S., 2015. Species diversity is dynamic and unbounded at local and continental scales. The American

Naturalist, 185, 584-593.

Lack, D., 1969. The numbers of bird species on islands. Bird Study, 16, 193-209.

Levine, J.M., Adler, P.B., Yelenik, S.G., 2004. A meta-analysis of biotic resistance to exotic plant invasions. Ecology Letters,

, 975-989.

Liñán, E., 23/6/2015. The trilobite fossil record of Murero. Personal communication.

Liñán, E., Gozalo, R., 1986. Trilobites del Cámbrico inferior y medio de Murero (Cordillera Ibérica). Memorias del Museo Paleontológico de la Universidad de Zaragoza, 2, 1-104.

Liñán, E., Gozalo, R., Dies Álvarez, M.E., Gámez Vintaned, J.A., Mayoral, E., Chirivella Martorell, J.B., Esteve, J., Zamora, S., Zhuravlev, A.Y., Andrés, J.A., 2008. Fourth International Trilobite Conference Trilo 08 Toledo, Spain, 2008. PostConference Field Trip. Lower and Middle Cambrian trilobites of selected localities in Cadenas Ibéricas (NE Spain). Universidad Zaragoza, 52pp.

Liñán, E., Gámez Vintaned, J.A., Dies Álvarez, M.E., Chirivella Martorell, J., Mayoral, E., Zhuravlev, A.Y., Andrés, J.A., Gozalo, R., 2013. 150 años del descubrimiento del yacimiento cámbrico de Murero (Cadenas Ibéricas, NE España). Geogaceta, 53, 25-28.

Lockwood, J.L., Hoopes, M.F., Marchetti, M.P., 2013. Invasion ecology. New York, John Wiley & Sons, 466pp.

Lodge, D.M., 1993. Biological invasions: lessons for ecology. Trends in Ecology and Evolution, 8, 133-137.

Lotze, F., 1969. El Cámbrico de España. Madrid, Memorias del Instituto Geológico y Minero de España, 256pp.

MacArthur, R.H., Wilson, E.O., 1967. The theory of island biogeography. Princeton University Press, 215pp.

Moen, D., Morlon, H., 2014. Why does diversification slow down? Trends in Ecology and Evolution, 29, 190-197.

Nürnberg, S., Aberhan, M., 2015. Interdependence of specialization and biodiversity in Phanerozoic marine invertebrates. Nature Communications, 6, 6602.

Pimm, S.L., Jones, H.L., Diamond, J., 1988. On the risk of extinction. The American Naturalist, 132, 757-785.

Price, T.D., 2015. The debate on determinants of species richness. The American Naturalist, 185, 571.

Price, T.D., Hooper, D.M., Buchanan, C.D., Johansson, U.S., Tietze, D.T., Alström, P., Olsson, U., Ghosh-Harihar, M., Ishtiaq, F., Gupta, S.K., Martens, J., Harr, B., Singh, P., Mohan, D., 2014. Niche filling slows the diversification of Himalayan songbirds. Nature, 509, 222-225.

Rabosky, D.L., Glor, R.E., 2010. Equilibrium speciation dynamics in a model adaptive radiation of island lizards. Proceedings of the National Academy of Sciences of the USA, 107, 22178-22183.

Rabosky, D.L., Adams, D.C., 2012. Rates of morphological evolution are correlated with species richness in salamanders. Evolution, 66, 1807-1818.

Rabosky, D.L., Hurlbert, A.H., 2015. Species richness at continental scale is dominated by ecological limits. The American Naturalist, 185, 572-583.

Ricklefs, R.E., 2010. Evolutionary diversification, coevolution between populations and their antagonists, and the filling of niche space. Proceedings of the National Academy of Sciences of the USA, 107, 1265-1272.

Rosenzweig, M.L., 1995. Species diversity in space and time. Cambridge University Press, 436pp.

Sepkoski, J.J., 1978. A kinetic model of Phanerozoic taxonomic diversity I. Analysis of marine orders. Paleobiology, 4, 223-

Stanley, S.M., 1979. Macroevolution, pattern and process. San Francisco, W.H. Freeman and Company, 343pp.

Warren, B.H., Simberloff, D., Ricklefs, R.E., Aguilée, R., Condamine, F.L., Gravel, D., Morlon, H., Mouquet, N., Rosindell, J., Casquet, J., Conti, E., Cornuault, J., FernándezPalacios, J.M., Hengl, T., Norder, S.J., Rijdsdijk, K.F., Sanmartín, I., Strasberg, D., Triantis, K.A., Valente, L.M., Whittaker, R.J., Gillespie, R.G., Emerson, B.C., Thébaud, D., 2015. Islands as model systems in ecology and evolution: prospects fifty years after MacArthur-Wilson. Ecology Letters, 18, 200-217.

Westrop, S.R., Adrain, J.M., 1998. Trilobite alpha diversity and the reorganization of Ordovician benthic marine communities. Paleobiology, 24, 1-16.

Whittaker, R.H., 1972. Evolution and measurement of species diversity. Taxon, 213-251.

Wilson, E.O., 1992. The diversity of life. Harvard University Press, 432pp.

Wunderle, J.M., 1985. An ecological comparison of the avifaunas of Grenada and Tobago, West Indies. Wilson Bulletin, 97,

-365.

Yoder, J.B., Clancey, E., Des Roches, S., Eastman, J.M., Gentry, L., Godsoe, W., Hagey, T.J., Jochimsen, D., Oswald, B.P., Robertson, J., Sarver, B.A.J., Schenk, J.J., Spear, S.F., Harmon, L.J., 2010. Ecological opportunity and the origin of adaptive radiations. Journal of Evolutionary Biology, 23, 1581-1596.

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2016-02-25

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