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Unveiling rare diversity by integrating museum, literature, and field data

Published online by Cambridge University Press:  08 April 2016

Paul G. Harnik
Paul G. Harnik*
Affiliation:
Committee on Evolutionary Biology, University of Chicago, Chicago, Illinois 60637. E-mail: [email protected]
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Abstract

Estimates of taxonomic richness and abundance are complicated by sampling biases. The failure to sample rare taxa is most often attributed to inadequate sampling and to removal during the process of sample-size standardization. Here I present two methods for unveiling rare diversity by integrating species presence/absence data from museum collections and the literature with quantitative estimates of species richness and abundance gathered from field-based bulk samples. Combining museum, literature, and field data can provide a more comprehensive estimate of taxonomic richness and abundance without substantial increase in current sampling effort. First, in a given bulk sample, the lowest proportional abundance value observed can be used to estimate the maximum abundance of rare species known to have occurred at the locality at least once but not recorded in the current sample. Second, a model-selection approach can be used, in which a set of relative abundance distribution models are fit to the bulk-sample abundance data and the parameter estimates for the best model used to calculate the abundance distribution for all species known from the locality. The Paleogene marine fossil record of the U.S. Gulf Coastal Plain is suitable for applying these methods, because (1) the molluscan fauna is well represented in museum collections and the literature, (2) the molluscan fauna has been taxonomically standardized, and (3) many classic localities remain accessible for standardized bulk sampling. I introduce these methods by applying them to a single locality and then, using the faunas of the Gosport, Moodys Branch, and Red Bluff Formations, I demonstrate how the model-fitting approach can be used to compare taxonomic richness among multiple localities. A substantial fraction of the molluscan richness known from each locality is not captured in bulk samples and much of this unobserved richness may be attributed to the rarity of species. The multiple-locality comparison suggests that the greatest Paleogene decline in standing richness occurred in the middle Eocene and that the recovery of richness following the Eocene-Oligocene extinction was quite rapid despite substantial loss of taxa. These analyses underscore the magnitude of veiled diversity in marine fossil assemblages and the potential of existing sources of data to unveil rare taxa, allowing them to be incorporated into quantitative diversity studies.

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Information
Paleobiology , Volume 35 , Issue 2 , Spring 2009 , pp. 190 - 208
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Copyright © The Paleontological Society 

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References

Literature Cited

Adrain, J. M., and Westrop, S. R. 2003. Paleobiodiversity: we need new data. Paleobiology 29:22–25.Google Scholar
Akaike, H. 1974. A new look at the statistical model identification. IEEE Transactions on Automatic Control 19:716–723.Google Scholar
Allmon, W. D. 1996. Systematics and evolution of Cenozoic American Turritellidae (Mollusca: Gastropoda) I: Paleocene and Eocene Coastal Plain species related to “Turritella mortoni Conrad” and “Turritella humerosa Conrad.” Palaeontographica Americana 59:1–134.Google Scholar
Allmon, W. D. 2005. The importance of museum collections in paleobiology. Paleobiology 31:1–5.Google Scholar
Alroy, J. 2002. How many named species are valid? Proceedings of the National Academy of Sciences USA 99:3706–3711.Google Scholar
Alroy, J., Aberhan, M., Bottjer, D. J., Foote, M., Fursich, F. T., Harries, P. J., Hendy, A. J. W., Holland, S. M., Ivany, L. C., Kiessling, W., Kosnik, M. A., Marshall, C. R., McGowan, A. J., Miller, A. I., Olszewski, T. D., Patzkowsky, M. E., Peters, S. E., Villier, L., Wagner, P. J., Bonuso, N., Borkow, P. S., Brenneis, B., Clapham, M. E., Fall, L. M., Ferguson, C. A., Hanson, V. L., Krug, A. Z., Layou, K. M., Leckey, E. H., Nurnberg, S., Powers, C. M., Sessa, J. A., Simpson, C., Tomasovych, A., and Visaggi, C. C. 2008. Phanerozoic trends in the global diversity of marine invertebrates. Science 321:97–100.Google Scholar
Bennington, J. B. 2003. Transcending patchiness in the comparative analysis of paleocommunities: a test case from the Upper Cretaceous of New Jersey. Palaios 18:22–33.Google Scholar
Bieler, R., and Mikkelsen, P. M. 2004. Marine bivalves of the Florida Keys: a qualitative faunal analysis based on original collections, museum holdings and literature data. Malacologia 46:503–544.Google Scholar
Bouchet, P., Lozouet, P., Maestrati, P., and Heros, V. 2002. Assessing the magnitude of species richness in tropical marine environments: exceptionally high numbers of molluscs at a New Caledonia site. Biological Journal of the Linnean Society 75:421–436.Google Scholar
Brown, J. H. 1984. On the relationship between abundance and distribution of species. American Naturalist 124:255–279.Google Scholar
Brown, J. H. Macroecology. University of Chicago Press, Chicago.Google Scholar
Budd, A. F., and Johnson, K. G. 2001. Contrasting patterns in rare and abundant species during evolutionary turnover. Pp. 295–325 in Jackson, J. B. C., Lidgard, S., and McKinney, F. K., eds. Evolutionary patterns: growth, form, and tempo in the fossil record. University of Chicago Press, Chicago.Google Scholar
Bulinski, K. V. 2007. Analysis of sample-level properties along a paleoenvironmental gradient: the behavior of evenness as a function of sample size. Palaeogeography, Palaeoclimatology, Palaeoecology 253:490–508.Google Scholar
Burnham, K. P., and Anderson, D. R. 2002. Model selection and multimodel inference: a practical information-theoretic approach. Springer, New York.Google 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 30:666–686.Google Scholar
Bush, A. M., Bambach, R. K., and Daley, G. M. 2007. Changes in theoretical ecospace utilization in marine fossil assemblages between the mid-Paleozoic and late Cenozoic. Paleobiology 33:76–97.CrossRefGoogle Scholar
Chisholm, R. A. 2007. Sampling species abundance distributions: resolving the veil-line debate. Journal of Theoretical Biology 247:600–607.Google Scholar
CoBabe, E. A., and Allmon, W. D. 1994. Effects of sampling on paleoecological and taphonomic analyses in high-diversity fossil accumulations: an example from the Eocene Gosport Sand, Alabama. Lethaia 27:167–178.Google Scholar
Crampton, J. S., Foote, M., Beu, A. G., Maxwell, P. A., Cooper, R. A., Matcham, L., Marshall, B. A., and Jones, C. M. 2006. The ark was full! Constant to declining Cenozoic shallow marine biodiversity on an isolated midlatitude continent. Paleobiology 32:509–532.Google Scholar
Davidoff, A. J., and Yancey, T. E. 1993. Eustatic cyclicity in the Paleocene and Eocene: data from the Brazos River Valley, Texas. Tectonophysics 222:371–395.Google Scholar
Davis, E. B., and Pyenson, N. D. 2007. Diversity biases in terrestrial mammalian assemblages and quantifying the differences between museum collections and published accounts: a case study from the Miocene of Nevada. Palaeogeography, Palaeoclimatology, Palaeoecology 250:139–149.Google Scholar
Dewdney, A. K. 1998. A general theory of the sampling process with applications to the “veil line.” Theoretical Population Biology 54:294–302.Google Scholar
Dockery, D. T., III. 1977. Mollusca of the Moodys Branch Formation, Mississippi. Mississippi Geological Survey Bulletin 120:1–212.Google Scholar
Dockery, D. T., III. 1982. Lower Oligocene Bivalvia of the Vicksburg Group in Mississippi. Mississippi Bureau of Geology Bulletin 123:1–261.Google Scholar
Dockery, D. T., III. 1986a. The Cockfield (Claiborne Group), Moodys Branch and Yazoo (Jackson Group) Formations at the Riverside Park locality in Jackson, Mississippi. Geological Society of America centennial field guide, pp. 401–403. Geological Society of America, Boulder, Colo. Google Scholar
Dockery, D. T., III. 1986b. Punctuated succession of Paleogene mollusks in the Northern Gulf Coastal Plain. Palaios 1:582–589.Google Scholar
Dockery, D. T. III., and Lozouet, P. 2003. Molluscan faunas across the Eocene/Oligocene boundary in the North American Gulf Coastal Plain, with comparisons to those of the Eocene and Oligocene of France. Pp. 303–340 in Prothero, D. R., Ivany, L. C., and Nesbitt, E. A., eds. From greenhouse to icehouse: the marine Eocene-Oligocene transition. Columbia University Press, New York.Google Scholar
Elder, S. R. 1981. Fossil assemblages of a marine transgressive sand, Moodys Branch Formation (Upper Eocene), Louisiana and Mississippi. . University of Texas, Austin.Google Scholar
Ellingsen, K. E. 2001. Biodiversity of a continental shelf soft-sediment macrobenthos community. Marine Ecology Progress Series 218:1–15.Google Scholar
Foote, M. 2000. Origination and extinction components of taxonomic diversity: general problems. In Erwin, D. H., and Wing, S. L., eds. Deep time: paleobiology's perspective. Paleobiology 26 (Suppl. to No. 4):74–102.Google Scholar
Foote, M. 2007. Symmetric waxing and waning of marine invertebrate genera. Paleobiology 33:517–529.Google Scholar
Foote, M., Crampton, J. S., Beu, A. G., Marshall, B. A., Cooper, R. A., Maxwell, P. A., and Matcham, I. 2007. Rise and fall of species occupancy in Cenozoic fossil mollusks. Science 318:1131–1134.Google Scholar
Frontier, S. 1985. Diversity and structure in aquatic ecosystems. Oceanography and Marine Biology: An Annual Review 23:253–312.Google Scholar
Garvie, C. L. 1996. The molluscan macrofauna of the Reklaw Formation, Marquez Member (Eocene: Lower Claibornian), in Texas. Bulletins of American Paleontology 111:1–177.Google Scholar
Gaston, K. J. 1994. Rarity. Chapman and Hall, London.Google Scholar
Gaston, K. J., and Blackburn, T. M. 2000. Pattern and process in macroecology. Blackwell Science, Oxford.Google Scholar
Gaston, K. J., Blackburn, T. M., and Lawton, J. H. 1997. Interspecific abundance-range size relationships: an appraisal of mechanisms. Journal of Animal Ecology 66:579–601.Google Scholar
Gotelli, N. J., and Colwell, R. K. 2001. Quantifying biodiversity: procedures and pitfalls in the measurement and comparison of species richness. Ecology Letters 4:379–391.Google Scholar
Gregory, R. D. 2000. Abundance patterns of European breeding birds. Ecography 23:201–208.Google Scholar
Guralnick, R., and Van Cleve, J. 2005. Strengths and weaknesses of museum and national survey data sets for predicting regional species richness: comparative and combined approaches. Diversity and Distributions 11:349–359.Google Scholar
Hansen, T. A., Kelley, P. H., and Haasl, D. M. 2004. Paleoecological patterns in molluscan extinctions and recoveries: comparison of the Cretaceous-Paleogene and Eocene-Oligocene extinctions in North America. Palaeogeography, Palaeoclimatology, Palaeoecology 214:233–242.Google Scholar
Harrison, H. C. 1994. Effects of environmental changes on molluscan evolutionary patterns, Gosport Sand (Middle Eocene), Southwest Alabama. . University of South Florida, Tampa.Google Scholar
Hayek, L.-A. C., and Buzas, M. A. 1997. Surveying natural populations. Columbia University Press, New York.Google Scholar
Heaslip, W. G. 1968. Cenozoic evolution of the alticostate venericards in Gulf and East Coastal North America. Palaeontographica Americana 6:51–135.Google Scholar
Holland, S. M., and Patzkowsky, M. E. 2002. Stratigraphic variation in the timing of first and last occurrences. Palaios 17:134–146.Google Scholar
Hubbell, S. P. 2001. The unified neutral theory of biodiversity and biogeography. Princeton University Press, Princeton, N.J. Google Scholar
Hunter, A. W., and Donovan, S. K. 2005. Field sampling bias, museum collections and completeness of the fossil record. Lethaia 38:305–314.Google Scholar
Hurlburt, S. H. 1971. The nonconcept of species diversity: a critique and alternative parameters. Ecology 52: 577–86.Google Scholar
Ivany, L. C. 1998. Sequence stratigraphy of the Middle Eocene Claiborne Stage, US Gulf Coastal Plain. Southeastern Geology 38:1–20.Google Scholar
Jackson, J. B. C., and Erwin, D. H. 2006. What can we learn about ecology and evolution from the fossil record? Trends in Ecology and Evolution 21:322–328.Google Scholar
Jackson, J. B. C., and Johnson, K. G. 2001. Measuring past biodiversity. Science 293:2401, 2403–2404.Google Scholar
Jaramillo, C. A., and Oboh-Ikuenobe, F. E. 1999. Sequence stratigraphic interpretations from palynofacies, dinocyst and lithological data of Upper Eocene-Lower Oligocene strata in southern Mississippi and Alabama, US Gulf Coast. Palaeogeography, Palaeoclimatology, Palaeoecology 145:259–302.Google Scholar
Johnson, K. G. 2003. New data for old questions. Paleobiology 29:19–21.Google Scholar
Kidwell, S. M. 2001. Preservation of species abundance in marine death assemblages. Science 294:1091–1094.Google Scholar
Kidwell, S. M. 2002. Time-averaged molluscan death assemblages: palimpsests of richness, snapshots of abundance. Geology 30:803–806.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 214:195–223.Google Scholar
Kosnik, M. A., and Wagner, P. J. 2006. Effects of taxon abundance distributions on expected numbers of sampled taxa. Evolutionary Ecology Research 8:195–211.Google Scholar
Kowalewski, M., Carroll, M., Casazza, L., Gupta, N., Hannisdal, B., Hendy, A., Krause, R. A. Jr., Labarbera, M., Lazo, D. G., Messina, C., Puchalski, S., Rothfus, T. A., Sälgeback, J., Stempien, J., Terry, R. C., and Tomašových, A. 2003. Quantitative fidelity of brachiopod-mollusk assemblages from modern subtidal environments of San Juan Islands, USA. Journal of Taphonomy 1:43–65.Google Scholar
Kowalewski, M., Kiessling, W., Aberhan, M., Fürsich, F. T., Scarponi, D., Wood, S. L. B., and Hoffmeister, A. P. 2006. Ecological, taxonomic, and taphonomic components of the post-Paleozoic increase in sample-level species diversity of marine benthos. Paleobiology 32:533–561.CrossRefGoogle Scholar
Krause, R. A. Jr., Stempien, J. A., Kowalewski, M., and Miller, A. I. 2006. Body size estimates from the literature: utility and potential for macroevolutionary studies. Palaios 22:60–73.CrossRefGoogle Scholar
Kunin, W. E., and Gaston, K. J., eds. 1997. The biology of rarity. Chapman and Hall, London.Google Scholar
Liow, L. H., and Stenseth, N. C. 2007. The rise and fall of species: implications for macroevolutionary and macroecological studies. Proceedings of the Royal Society of London B 274:2745–2752.Google ScholarPubMed
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 100:2478–2482.Google Scholar
Lockwood, R., and Chastant, L. R. 2006. Quantifying taphonomic bias of compositional fidelity, species richness, and rank abundance in molluscan death assemblages from the upper Chesapeake Bay. Palaios 21:376–383.CrossRefGoogle Scholar
MacNeil, F. S., and Dockery, D. T. 1984. Lower Oligocene Gastropoda, Scaphopoda, and Cephalopoda of the Vicksburg Group in Mississippi. Mississippi Bureau of Geology Bulletin 124:1–415.Google Scholar
Magurran, A. E., and Henderson, P. A. 2003. Explaining the excess of rare species in natural species abundance distributions. Nature 422:714–716.Google Scholar
Mancini, E. A., and Tew, B. H. 1992. Paleogene unconformity-bounded depositional sequences of southwest Alabama: lithofacies, systems tracts, and sequence boundaries. Alabama Geological Society Guidebook 29:1–72.Google Scholar
Marquet, P. A., Keymer, J. A., and Cofre, H. 2003. Breaking the stick in space: of niche models, metacommunities and patterns in the relative abundance of species. Pp. 64–86 in Blackburn, T. M. and Gaston, K. J., eds. Macroecology: concepts and consequences. Blackwell Science, Oxford.Google Scholar
McCullagh, P., and Nelder, J. A. 1989. Generalized linear models. Chapman and Hall, London.Google Scholar
McGill, B. J. 2003. Does Mother Nature really prefer rare species or are log-left-skewed SADs a sampling artefact? Ecology Letters 6:766–773.Google Scholar
McGill, B. J., Etienne, R. S., Gray, J. S., Alonso, D., Anderson, M. J., Benecha, H. K., Dornelas, M., Enquist, B. J., Green, J. L., He, F., Hurlbert, A. H., Magurran, A. E., Marquet, P. A., Maurer, B. A., Ostling, A., Soykan, C. U., Ugland, K. I., and White, E. P. 2007. Species abundance distributions: moving beyond single prediction theories to integration within an ecological framework. Ecology Letters 10:995–1015.Google Scholar
Meldahl, K. H. 1990. Sampling, species abundance, and the stratigraphic signature of mass extinction: a test using Holocene tidal flat mollusks. Geology 18:890–893.Google Scholar
Mouillot, D., and Lepretre, A. 1999. A comparison of species diversity estimators. Researches on Population Ecology 41:203–215.Google Scholar
Mouillot, D., and Lepretre, A. 2000. Introduction of relative abundance distribution (RAD) indices, estimated from the rank-frequency diagrams (RD), to assess changes in community diversity. Environmental Monitoring and Assessment 63:279–295.Google Scholar
Novack-Gottshall, P. M. 2007. Using a theoretical ecospace to quantify the ecological diversity of Paleozoic and modern marine biotas. Paleobiology 33:273–294.Google Scholar
Nunes, F., Fukami, H., Vollmer, S. V., Norris, R. D., and Knowlton, N. 2008. Re-evaluation of the systematics of the endemic corals of Brazil by molecular data. Coral Reefs 27:423–432.Google Scholar
Oksanen, J., Kindt, R., Legendre, P., and O'Hara, R. B. 2007. Vegan: R functions for vegetation ecologists, Version 1. 8.5. URL: http://cc.oulu.fi/~jarioksa/.Google Scholar
Olszewski, T. D. 2004. A unified mathematical framework for the measurement of richness and evenness within and among multiple communities. Oikos 104:377–387.Google Scholar
Olszewski, T. D., and Erwin, D. H. 2004. Dynamic response of Permian brachiopod communities to long-term environmental change. Nature 428:738–741.Google Scholar
Palmer, K. V. W., and Brann, D. C. 1965. Catalogue of the Paleocene and Eocene Mollusca of the southern and eastern United States, Part I. Pelecypoda, Amphineura, Pteropoda, Scaphopoda, and Cepholopoda. Bulletins of American Paleontology 48(218):1–466.Google Scholar
Palmer, K. V. W., and Brann, D. C. 1966. Catalogue of the Paleocene and Eocene Mollusca of the southern and eastern United States, Part II. Gastropoda. Bulletins of American Paleontology 48:467–1057.Google Scholar
Pandolfi, J. M. 1996. Limited membership in Pleistocene reef coral assemblages from the Huon Peninsula, Papua New Guinea: constancy during global change. Paleobiology 22:152–176.Google Scholar
Pasley, M. A., and Hazel, J. E. 1995. Revised sequence stratigraphic interpretation of the Eocene-Oligocene boundary interval, Mississippi and Alabama, Gulf-Coast Basin, USA. Journal of Sedimentary Research 65:160–169.Google Scholar
Peters, S. E. 2004. Evenness of Cambrian–Ordovician benthic marine communities in North America. Paleobiology 30:325–346.2.0.CO;2>CrossRefGoogle Scholar
Peters, S. E. 2006. Genus richness in Cambrian-Ordovician benthic marine communities in North America. Palaios 21:580–587.Google Scholar
Petersen, F. T., Meier, R., and Nykjaer, M. 2003. Testing species richness estimation methods using museum label data on the Danish Asilidae. Biodiversity and Conservation 12:687–701.Google Scholar
Powell, M. G., and Kowalewski, M. 2002. Increase in evenness and sampled alpha diversity through the Phanerozoic: comparison of early Paleozoic and Cenozoic marine fossil assemblages. Geology 30:331–334.Google Scholar
Preston, F. W. 1948. The commonness, and rarity, of species. Ecology 29:254–283.Google Scholar
R Development Core Team. 2006. R: a language and environment for statistical computing, Version 2. 3.1. R Foundation for Statistical Computing, Vienna.Google Scholar
Raup, D. M. 1975. Taxonomic diversity estimation using rarefaction. Paleobiology 1:333–342.Google Scholar
Sanders, H. L. 1968. Marine benthic diversity—a comparative study. American Naturalist 102:243–282.Google Scholar
Scarponi, D., and Kowalewski, M. 2004. Stratigraphic paleoecology: bathymetric signatures and sequence overprint of mollusk associations from upper Quaternary sequences of the Po Plain, Italy. Geology 32:989–992.Google Scholar
Scarponi, D., and Kowalewski, M. 2007. Sequence stratigraphic anatomy of diversity patterns: Late Quaternary benthic mollusks of the Po Plain, Italy. Palaios 22:296–305.Google Scholar
Schlacher, T. A., Newell, P., Clavier, J., Schlacher-Hoenlinger, M. A., Chevillon, C., and Britton, J. 1998. Soft-sediment benthic community structure in a coral reef lagoon—the prominence of spatial heterogeneity and ‘spot endemism.’ Marine Ecology Progress Series 174:159–174.Google Scholar
Shin, P. K. S., and Ellingsen, K. E. 2004. Spatial patterns of soft-sediment benthic diversity in subtropical Hong Kong waters. Marine Ecology Progress Series 276:25–35.Google Scholar
Signor, P. W. III., and Lipps, J. H. 1982. Sampling bias, gradual extinction patterns, and catastrophes in the fossil record. In Silver, L. T. and Schultz, P. H., eds. Geological implications of impacts of large asteroid and comets on the earth. Geological Society of America Special Paper 190:291–296.Google Scholar
Swindel, D. B. 1986. A paleoecological study of the Gosport Sand (Claibornian: Middle Eocene) in southwest Alabama. . University of Alabama, Tuscaloosa.Google Scholar
Tew, B. H., and Mancini, E. A. 1995. An integrated stratigraphic method for paleogeographic reconstruction: examples from the Jackson and Vicksburg Groups of the eastern Gulf Coastal Plain. Palaios 10:133–153.CrossRefGoogle Scholar
Thompson, W. L., ed. 2004. Sampling rare or elusive species. Island Press, Washington, D.C. Google ScholarPubMed
Tipper, J. C. 1979. Rarefaction and rarefiction: use and abuse of a method in paleoecology. Paleobiology 5:423–434.Google Scholar
Ulrich, W., and Ollik, M. 2004. Frequent and occasional species and the shape of relative-abundance distributions. Diversity and Distributions 10:263–269.Google Scholar
Wagner, P. J., Kosnik, M. A., and Lidgard, S. 2006. Abundance distributions imply elevated complexity of post-Paleozoic marine ecosystems. Science 314:1289–1292.Google Scholar
Wagner, P. J., Aberhan, M., Hendy, A., and Kiessling, W. 2007. The effects of taxonomic standardization on sampling-standardized estimates of historical diversity. Proceedings of the Royal Society of London B 274:439–444.Google Scholar
Webster, M., Sadler, P. M., Kooser, M. A., and Fowler, E. 2003. Combining stratigraphic sections and museum collections to increase biostratigraphic resolution. Pp. 95–128 in Harries, P. J., ed. Approaches in high-resolution stratigraphic paleontology. Kluwer Academic, Amsterdam.Google Scholar
Zuschin, M., and Oliver, P. G. 2005. Diversity patterns of bivalves in a coral dominated shallow-water bay in the northern Red Sea—high species richness on a local scale. Marine Biology Research 1:396–410.Google Scholar
Zuschin, M., and Stanton, R. J. 2002. Paleocommunity reconstruction from shell beds: a case study from the Main Glauconite Bed, Eocene, Texas. Palaios 17:602–614.Google Scholar
Zuschin, M., Harzhauser, M., and Sauermoser, K. 2006. Patchiness of local species richness and its implication for large-scale diversity patterns: an example from the middle Miocene of the Paratethys. Lethaia 39:65–88.Google Scholar
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