The fundamental biodiversity number, θ, as proposed by Hubbell, should be positively correlated with province area. Because θ can be calculated from preserved relative abundance distributions, this correlation can be tested in the fossil record for regions with known provinces. Late Ordovician (443–458 Ma) strata of Laurentia are divided into four geochemically and biologically distinct regions that reflect provinces in the epicontinental sea. We use existing and newly obtained bed-level census data to test whether Hubbell’s θ is positively correlated with the area of these four regions, corresponding roughly to the Appalachian Basin, Cincinnati Arch, Upper Mississippi Valley, and western United States and Canada. Results indicate a positive relationship between province area and θ that suggests the influence of provincial area, among other factors, on diversity. This correlation highlights the inherent link between diversity and abundance structure at local and regional scales, such that changes at one scale will necessarily affect the other. Since diversity at these smaller spatial scales is an important component of global biodiversity, determining the nature of this relationship in the fossil record has implications for understanding how diversity is assembled globally throughout the Phanerozoic.
References
[1]
Hubbell, S.P. The Unified Neutral Theory of Biogeography; Princeton University Press: Princeton, NJ, USA, 2001; p. 375.
[2]
Holmden, C.; Creaser, R.A.; Muehlenbachs, K.; Leslie, S.A.; Bergstr？m, S.M. Isotopic evidence for geochemical decoupling between ancient epeiric seas and bordering oceans: implications for secular curves. Geology 1998, 26, 567–570.
[3]
Jin, J.; Harper, D.A.T.; Rasmussen, J.A.; Sheehan, P.M. Late Ordovician massive-bedded Thalassinoides ichnofacies along the paleoequator of Laurentia. Palaeogeogr. Palaeocl. 2012, 367–368, 73–88.
[4]
Panchuk, K.M.; Holmden, C.E.; Leslie, S.A. Local controls on carbon cycling in the Ordovician midcontinent region of North America, with implications for carbon isotope secular curves. J. Sediment. Res. 2006, 76, 200–211, doi:10.2110/jsr.2006.017.
[5]
Alonso, D.; Etienne, R.S.; McKane, A.J. The merits of neutral theory. Trends Ecol. Evol. 2006, 21, 451–457, doi:10.1016/j.tree.2006.03.019.
[6]
Purves, D.W.; Turnbull, L.A. Different but not equal: the implausible assumption at the heart of neutral theory. J. Anim. Ecol. 2010, 79, 1215–1225, doi:10.1111/j.1365-2656.2010.01738.x.
[7]
Rosindell, J.; Hubbell, S.P.; Etienne, R.S. The unified neutral theory of biodiversity and biogeography at age ten. Trends Ecol. Evol. 2010, 26, 340–348, doi:10.1016/j.tree.2011.03.024.
[8]
Hubbell, S.P. The neutral theory of biodiversity and biogeography and Stephen Jay Gould. Paleobiology 2005, 31, 122–132, doi:10.1666/0094-8373(2005)031[0122:TNTOBA]2.0.CO;2.
[9]
Olszewski, T.D.; Erwin, D.H. Dynamic response of Permian brachiopod communities to long-term environmental change. Nature 2004, 428, 738–741, doi:10.1038/nature02464.
[10]
Toma？ovych, A.; Kidwell, S.M. Predicting the effects of increasing temporal scale on species composition, diversity, and rank abundance distributions. Paleobiology 2010, 36, 672–695, doi:10.1666/08092.1.
[11]
Scotese, C.R.; McKerrow, W.S. Ordovician plate tectonic reconstructions. Geol. S. Can. Paper 1991, 90, 271–282.
[12]
Beaumont, C.; Quinlan, G.; Hamilton, J. Orogeny and stratigraphy: Numerical models of the Paleozoic in the eastern interior of North America. Tectonics 1988, 7, 389–416, doi:10.1029/TC007i003p00389.
[13]
Sweet, W.C.; Bergstr？m, S.M. Provincialism exhibited by Ordovician conodont faunas. SEPM Spec. P. 1974, 21, 189–202.
[14]
Barnes, C.R.; F？hr？us, L.E. Provinces, communities, and the proposed nektobenthic habit of Ordovician conodontophorids. Lethaia 1975, 8, 133–149, doi:10.1111/j.1502-3931.1975.tb01308.x.
[15]
Elias, R.J. Solitary rugose corals of the Stony Mountain Formation, Southern Manitoba, and its equivalents. J. Paleontol. 1983, 57, 924–956.
[16]
Antsey, R.L. Bryozoan provinces and patterns of generic evolution and extinction in the Late Ordovician of North America. Lethaia 1986, 19, 33–51, doi:10.1111/j.1502-3931.1986.tb01898.x.
[17]
Blakey, R. Global Paleogeography. Available online: http://jan.ucc.nau.edu/rcb7/globaltext2.html/ (accessed on 18 April 2012).
[18]
Piepgras, D.J.; Wasserburg, G.J. Neodymium isotopic variations in seawater. Earth Planet. Sc. Lett. 1980, 50, 128–138, doi:10.1016/0012-821X(80)90124-7.
[19]
Patterson, W.P.; Walter, L.M. Depletion of 13C in seawater ΣCO2 on modern carbonate platforms: significance for the carbon isotopic record of carbonates. Geology 1994, 22, 885–888, doi:10.1130/0091-7613(1994)022<0885:DOCISC>2.3.CO;2.
[20]
Bergstr？m, S.M. Biogeography, evolutionary relationships, and biostratigraphic significance of Ordovician platform conodonts. Foss. Strata 1983, 15, 35–58.
[21]
Riva, J. A revision of some Ordovician graptolites of eastern North America. Palaeontology 1974, 17, 1–40.
[22]
Bergstr？m, S.M.; Mitchell, C.E. The graptolite correlation of the North American Upper Ordovician standard. Lethaia 1986, 19, 247–266, doi:10.1111/j.1502-3931.1986.tb00739.x.
[23]
Berry, W.B.N. Review of late Middle Ordovician graptolites in eastern New York and Pennsylvania. Am. J. Sci. 1970, 269, 304–313, doi:10.2475/ajs.269.3.304.
[24]
Goldman, D.; Bergstr？m, S.M.; Mitchell, C.E. Revision of the Zone 13 graptolite biostratigraphy in the Marathon, Texas, standard succession and its bearing on Upper Ordovician graptolite biostratigraphy. Lethaia 1995, 28, 115–128, doi:10.1111/j.1502-3931.1995.tb01601.x.
[25]
Mohibullah, M.; Williams, M.; Vandenbroucke, T.R.A.; Sabbe, K.; Zalasiewicz, J.A. Marine ostracod provinciality in the Late Ordovician of palaeocontinental Laurentia and its environmental and geographical expression. PLOS one 2012, 7, e41682.
[26]
Bretsky, P.W. Central Appalachian Late Ordovician communities. Geol. Soc. Am. Bull. 1969, 80, 193–212, doi:10.1130/0016-7606(1969)80[193:CALOC]2.0.CO;2.
[27]
Elias, R.J.; Young, G.A. Coral diversity, ecology, and provincial structure during a time of crisis: the latest Ordovician to earliest Silurian Edgewood Province in Laurentia. Palaios 1998, 13, 98–112, doi:10.2307/3515483.
[28]
Elias, R.J. Latest Ordovician solitary rugose corals of eastern North America. Bull. Am. Paleontol. 1982, 81, 1–85.
[29]
Flower, R.H. Ordovician cephalopods of the Cincinnati region, part 1. Bull. Am. Paleontol. 1946, 29, 1–656.
[30]
Nelson, S.J. Arctic Ordovician fauna: an equatorial assemblage. J. Alberta Soc. Petr. Geol. 1959, 7, 45–47.
[31]
Jin, J.; Zhan, R.B. Late Ordovician Articulate Brachiopods from the Red River and Stony Mountain Formations, Southern Manitoba; NRC Research Press: Ottawa, Canada, 2001; p. 117.
[32]
Jin, J.; Caldwell, W.G.E.; Norford, B.S. Late Ordovician brachiopods Rafinesquina lata Whiteaves, 1896 and Kjaerina hartae n. sp. from southern Manitoba and the Hudson Bay lowlands. Can. J. Earth Sci. 1995, 32, 1255–1266, doi:10.1139/e95-102.
Holland, S.M.; Patzkowsky, M.E. Sequence stratigraphy and long-term paleoceanographic change in the Middle and Upper Ordovician of the eastern United States. Geol. Soc. Am. S. 1996, 306, 117–129.
[35]
Holland, S.M.; Patzkowsky, M.E. Ecosystem structure and stability: Middle Upper Ordovician of central Kentucky, USA. Palaios 2004, 19, 316–331, doi:10.1669/0883-1351(2004)019<0316:ESASMU>2.0.CO;2.
[36]
Holland, S.M.; Patzkowsky, M.E. Gradient ecology of a biotic invasion: biofacies of the type Cincinnatian Series (Upper Ordovician), Cincinnati, Ohio region, USA. Palaios 2007, 22, 392–407, doi:10.2110/palo.2006.p06-066r.
[37]
Holland, S.M.; Patzkowsky, M.E. Stratigraphic architecture of a tropical carbonate platform and its effect on the distribution of fossils: Ordovician Bighorn Dolomite, Wyoming, USA. Palaios 2009, 24, 303–317, doi:10.2110/palo.2008.p08-095r.
[38]
Paleobiology Database Home Page. Available online: http://paleodb.org/ (accessed on 29 September 2012).
[39]
R Development Core Team. R: A Language and Environment for Statistical Computing; R Foundation for Statistical Computing: Vienna, Austria, 2011. ISBN 3-900051-07-0. Available online: http://www.R-project.org/ (accessed on 19 August 2011).
[40]
Etienne, R.S. A neutral sampling formula for multiple samples and an ‘exact’ test of neutrality. Ecol. Lett. 2007, 10, 608–618, doi:10.1111/j.1461-0248.2007.01052.x.
[41]
Etienne, R.S. Maximum likelihood estimation of neutral model parameters for multiple samples with different degrees of dispersal limitation. J. Theor. Biol. 2009, 257, 510–514, doi:10.1016/j.jtbi.2008.12.016.
[42]
PARI/GP Home Page. Available online: http://pari.math.u-bordeaux.fr/ (accessed on 3 April 2012).
[43]
ESRI, ArcGIS Desktop: Release 10, Environmental Systems Research Institute, Redlands, CA, USA, 2011.
[44]
Holland, S.M.; Patzkowsky, M.E. Sequence architecture of the Bighorn Dolomite, Wyoming, USA: Transition to the Late Ordovician icehouse. J. Sediment. Res. 2012, 82, 599–615, doi:10.2110/jsr.2012.52.
[45]
Lobdell, F.K. Arthrostylidae (Bryozoa: Cryptostomata) from the Gunn Member, Stony Mountain Formation (Upper Ordovician), North Dakota and Manitoba. N. Dak. Geol. Survey Misc. Series 1992, 76, 99–115.
[46]
Richards, P.W.; Nieschmidt, C.L. The Bighorn Dolomite and Correlative Formations in Southern Montana and Northern Wyoming (Oil and Gas Investigations Map, OM-202) [Map]; U.S. Geological Survey: Reston, Virginia, VA, USA, 1961.
[47]
Longman, M.W.; Fertal, T.G.; Glennie, J.S. Origin and geometry of Red River Dolomite reservoirs, western Williston Basin. AAPG Bull. 1983, 67, 744–771.
Preston, F.W. The commonness and rarity of species. Ecology 1948, 29, 254–283, doi:10.2307/1930989.
[50]
Rex, M.A. Community structure in the deep-sea benthos. Annu. Rev. Ecol. Syst. 1981, 12, 331–353.
[51]
Patzkowsky, M.E.; Holland, S.M. Extinction, invasion, and sequence stratigraphy: patterns of faunal change in the Middle and Upper Ordovician of the eastern United States. Geol. S. Am. S. 1996, 306, 131–142.
[52]
Allen, A.P.; Gillooly, J.F.; Savage, V.M.; Brown, J.H. Kinetic effects of temperature on rates of genetic divergence and speciation. Proc. Natl. Acad. Sci. USA 2006, 103, 9130–9135.
[53]
Jablonski, D.; Roy, K.; Valentine, J.W. Out of the tropics: evolutionary dynamics of the latitudinal diversity gradient. Science 2006, 314, 102–106, doi:10.1126/science.1130880.
[54]
Crame, J.A. Evolution of taxonomic diversity gradients in the marine realm: a comparison of Late Jurassic and Recent bivalve faunas. Paleobiology 2002, 28, 184–207, doi:10.1666/0094-8373(2002)028<0184:EOTDGI>2.0.CO;2.
[55]
Rosenzweig, M.L. Species Diversity in Space and Time; Cambridge University Press: Cambridge, UK, 1995; p. 436.
[56]
Palmer, M.W.; White, P.S. Scale dependence and the species-area relationship. Am. Nat. 1994, 144, 717–740.
[57]
Shmida, A.; Wilson, M.V. Biological Determinants of species diversity. J. Biogeogr. 1985, 12, 1–20, doi:10.2307/2845026.
[58]
Preston, F.W. Time and space variation of species. Ecology 1960, 41, 611–627, doi:10.2307/1931793.
[59]
Hu, X.-S.; He, F.; Hubbell, S.P. Species diversity in local neutral communities. Am. Nat. 2007, 170, 844–853.
[60]
Bush, A.M.; Bambach, R.K. Does alpha diversity increase during the Phanerozoic? Lifting the veils of taphonomic, latitudinal, and environmental biases. J. Geol. 2004, 112, 625–642, doi:10.1086/424576.
[61]
Sepkoski, J.J., Jr. Alpha, beta, or gamma: where does all the diversity go? Paleobiology 1988, 14, 221–234.
[62]
Valentine, J.W.; Foin, T.C.; Peart, D. A provincial model of Phanerozoic marine diversity. Paleobiology 1978, 4, 55–66.
[63]
Alroy, J.; Aberhan, M.; Bottjer, D.J.; Foote, M.; Fürsich, F.T.; Harries, P.J.; Hendy, A.J.W.; Holland, S.M.; Ivany, L.C.; Kiessling, W.; et al. Phanerozoic trends in the global diversity of marine invertebrates. Science 2008, 321, 97–100.
