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Quantitative comparisons and models of time-averaging in bivalve and brachiopod shell accumulations

  • Richard A. Krause (a1), Susan L. Barbour (a2), Michał Kowalewski (a3), Darrell S. Kaufman (a4), Christopher S. Romanek (a5), Marcello G. Simões (a6) and John F. Wehmiller (a7)...


The variation in time-averaging between different types of marine skeletal accumulations within a depositional system is not well understood. Here we provide quantitative data on the magnitude of time-averaging and the age structure of the sub-fossil record of two species with divergent physical and ecological characteristics, the brachiopod Bouchardia rosea and the bivalve Semele casali. Material was collected from two sites on a mixed carbonate-siliciclastic shelf off the coast of Brazil where both species are dominant components of the local fauna.

Individual shells (n = 178) were dated using amino acid racemization (aspartic acid) calibrated with 24 AMS radiocarbon dates. Shell ages range from modern to 8118 years b.p. for brachiopods, and modern to 4437 years for bivalves. Significant differences in the shape and central tendency of age-frequency distributions are apparent between each sample. Such differences in time-averaging magnitude confirm the assumption that taphonomic processes are subject to stochastic variation at all spatial and temporal scales. Despite these differences, each sample is temporally incomplete at centennial resolution and three of the four samples have similar right-skewed age-frequency distributions. Simulations of temporal completeness indicate that samples of both species from the shallow site are consistent with a more strongly right-skewed and less-complete age-frequency distribution than those from the deep site.

We conclude that intrinsic characteristics of each species exert less control on the time-averaging signature of these samples than do extrinsic factors such as variation in rates of sedimentation and taphonomic destruction. This suggests that brachiopod-dominated and bivalve-dominated shell accumulations may be more similar in temporal resolution than previously thought, and that the temporal resolution of multi-taxic shell accumulations may depend more on site-to-site differences than on the intrinsic properties of the constituent organisms.



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Allmon, W. D. 1989. Paleontological completeness of the record of lower Tertiary mollusks, U.S. Gulf and Atlantic Coastal Plains: implications for phylogenetic studies. Historical Biology 3:141158.
Anderson, L. C., Gupta, B. K. Sen, McBride, R. A., and Byers, M. R. 1997. Reduced seasonality of Holocene climate and pervasive mixing of Holocene marine section: northeastern Gulf of Mexico shelf. Geology 25:127130.
Angulo, R. J., and Lessa, G. C. 1997. The Brazilian sea-level curves: a critical review with emphasis on the curves from the Paranaguá and Cananéia regions. Marine Geology 140:141166.
Angulo, R. J., Giannini, P. C. F., Suguio, K., and Pessenda, L. C. R. 1999. Relative sea-level changes in the last 550 years in southern Brazil (Laguna-Imbituba Region, Santa Catarina State) based on vermetid 14C ages. Marine Geology 159:323339.
Angulo, R. J., de Souza, M. C., Reimer, P. J., and Sasaoka, S. K. 2005. Reservoir effect of the Southern and Southeastern Brazilian Coast. Radiocarbon 47:6773.
Angulo, R. J., Lessa, G. C., and de Souza, M. C. 2006. A critical review of mid- to late-Holocene sea-level fluctuations on the eastern Brazilian coastline. Quaternary Science Reviews 25:486586.
Ayers, G. P. 2001. Comment on regression analysis of air quality data. Atmospheric Environment 35:24232425.
Wood, S. L. Barbour, Krause, R. A. Jr., Kowalewski, M., Wehmiller, J. F., and Simões, M. G. 2006. Aspartic acid racemization dating of Holocene brachiopods and bivalves from the Southern Brazilian Shelf, South Atlantic. Quaternary Research 66:323331.
Behrensmeyer, A. K., Kidwell, S. M., and Gastaldo, R. A. 2000. Taphonomy and paleobiology. In Erwin, D. H. and Wing, S. L., eds. Deep time: Paleobiology's perspective Paleobiology 26(Suppl. to No. 4):103147.
Behrensmeyer, A. K., Fürsich, F. T., Gastaldo, R. A., Kidwell, S. M., Kosnik, M. A., Kowalewski, M., Plotnick, R. E., Rogers, R. R., and Alroy, J. 2005. Are the most durable shelly taxa also the most common in the marine fossil record? Paleobiology 31:607623.
Blackwell, B. A. B., Last, W. M., and Rutter, N. W. 2000. Biogeochemical diagenesis in recent mammalian bones from saline lakes in western Victoria, Australia. Pp. 88107 in Goodfriend, et al. 2000.
Boss, K. J. 1972. The genus Semele in the Western Atlantic. Johnsonia 5(49):132.
Braga, E. S., and Muller, T. J. 1998. Observation of regeneration of nitrate, phosphate and silicate during upwelling off Ubatuba, Brazil, 23 degrees south. Continental Shelf Research 18:915922.
Brigham, J. K. 1983. Intrashell variations in amino acid concentrations and isoleucine epimerization ratios in fossil Hiatella arctica . Geology 11:509513.
Brunton, C. H. C. 1996. The functional morphology of the Recent brachiopod Bouchardia rosea . Acta Zoologica 77:233240.
Burnham, R. J. 1993. Time resolution in terrestrial macrofloras; guidelines from modern accumulations. Pp. 5778 in Kidwell, and Behrensmeyer, 1993.
Burone, L. 2002. Foraminíferos bentônicos e parâmetros físicoquímicos da enseada de Ubatuba, São Paulo: estudo ecológico em uma área com poluição orgânica. Ph.D. dissertation. University of São Paulo, Institute of Oceanography, São Paulo.
Burone, L., Muniz, P., Pires-Vanin, A. M. S., and Rodrigues, M. 2003. Spatial distribution of organic matter in the surface sediments of Ubatuba Bay (Southeastern Brazil). Anais da Academia Brasileira de Ciências 75:7790.
Bush, A. M., Powell, M. G., Arnold, W. S., Bert, T. M., and Daley, G. M. 2002. Time-averaging, evolution, and morphologic variation. Paleobiology 28:925.
Callender, W. R., Staff, G. M., Powell, E. N., and MacDonald, I. R. 1990. Gulf of Mexico hydrocarbon seep communities. V. Biofacies and shell orientation of autochthonous shell beds below storm wave base. Palaios 5:214.
Callender, W. R., Powell, E. N., Staff, G. M., Davies, D. J., Beauchamp, B., and von Bitter, P. H. 1992. Distinguishing autochthony, parautochthony and allochthony using taphofacies analysis; can cold seep assemblages be discriminated from assemblages of the nearshore and continental shelf? Palaios 7:409421.
Callender, W. R., Powell, E. N., and Staff, G. M. 1994. Taphonomic rates of molluscan shells placed in autochthonous assemblages on the Louisiana continental slope. Palaios 9:6073.
Callender, W. R., Staff, G.M., Parsons-Hubbard, K. M., Powell, E. N., Rowe, G. T., Walker, S. E., Brett, C. E., Raymond, A., Carlson, D., White, S., and Heiss, E. A. 2002. Taphonomic trends along a forereef slope: Lee Stocking Island, Bahamas. I. Location and water depth. Palaios 17:5065.
Campos, E. J. D., Velhot, D., and da Silveira, I. C. A. 1995. Water mass characteristics and geostrophic circulation in the South Brazil Bight: summer of 1991. Journal of Geophysical Research 100:1853718550.
Campos, E. J. D., Velhot, D., and da Silveira, I. C. A. 2000. Shelf break upwelling driven by Brazil Current cyclonic meanders. Geophysical Research Letters 27:751754.
Carroll, M., Kowalewski, M., Simões, M. G., and Goodfriend, G. A. 2003. Quantitative estimates of time-averaging in brachiopod shell accumulations from a modern tropical shelf. Paleobiology 29:381402.
Castelao, R. M., Campos, E. J. D., and Miller, J. L. 2004. A modeling study of coastal upwelling driven by wind and meanders of the Brazil Current. Journal of Coastal Research 20:662671.
Cherns, L., and Wright, V. P. 2000. Missing mollusks as evidence of large-scale, early skeletal aragonite dissolution in a Silurian sea. Geology 28:791794.
Coan, E. V. 1988. Recent eastern Pacific species of the bivalve genus Semele . Veliger 31:142.
Cummins, H., Powell, E. N., Stanton, R. J. Jr., and Staff, G. 1986. The rate of taphonomic loss in modern benthic habitats: how much of the potentially preservable community is preserved? Palaeogeography Palaeoclimatology Palaeoecology 89:193204.
da Rocha, J., Milliman, J. D., Santana, C. I., and Vicalvi, M. A. 1975. Southern Brazil. In Milliman, J. D., and Summerhayes, C. P., eds. Upper continental margin sedimentation off Brazil. Contributions to Sedimentology No. 4:117150. Schweizerbart, Stuttgart.
Depetris, P. J., Kemp, S., Latif, M., and Mook, W. G. 1996. ENSO-controlled flooding in the Paraná River (1904–1991). Naturwissenschaften 83:127129.
Diaconis, P., and Efron, B. 1983. Computer intensive methods in statistics. Scientific American 248:116130.
DiCiccio, T. J., and Romano, J. P. 1988. A review of bootstrap confidence intervals. Journal of the Royal Statistical Society B 50:338354.
Domaneschi, O. 1995. A comparative study of the functional morphology of Semele purpurascens (Gmelin, 1791) and Semele proficua . Veliger 38:323342.
Efron, B. 1981. Nonparametric standard errors and confidence intervals. Canadian Journal of Statistics 9:139172.
Flessa, K. W. 1998. Well-traveled cockles: shell transport during the Holocene transgression of the southern North Sea. Geology 26:187190.
Flessa, K. W., and Kowalewski, M. 1994. Shell survival and time averaging in nearshore and shelf environments: estimates from the radiocarbon literature. Lethaia 27:153165.
Flessa, K. W., Cutler, A. H., and Meldahl, K. H. 1993. Time and taphonomy: quantitative estimates of time-averaging and stratigraphic disorder in a shallow marine habitat. Paleobiology 19:266286.
Folk, R. L. 1959. Practical petrographic classification of limestones. American Association of Petroleum Geologists Bulletin 43:138.
Fürsich, F. T., and Aberhan, M. 1990. Significance of time averaging for paleocommunity analysis. Lethaia 23:143152.
Goodfriend, G. A. 1987. Chronostratigraphic studies of sediments in the Negev Desert, using amino acid epimerization analysis of land snail shells. Quaternary Research 28:374392.
Goodfriend, G. A. 1989. Complementary use of amino acid epimerization and radiocarbon analysis for dating mixed-age fossil assemblages. Radiocarbon 31:10411047.
Goodfriend, G. A. 1992. Rapid racemization of aspartic acid in mollusc shells and potential for dating over Recent centuries. Nature 357:399401.
Goodfriend, G. A., and Stanley, D. J. 1996. Reworking and discontinuities in Holocene sedimentation in the Nile Delta: documentation from amino acid racemization and stable isotopes in mollusk shells. Marine Geology 129:271283.
Goodfriend, G. A., Brigham-Grette, J., and Miller, G. H. 1996. Enhanced age resolution of the marine Quaternary record in the Arctic using aspartic acid racemization dating of bivalve shells. Quaternary Research 45:176187.
Goodfriend, G. A., Collins, M. J., Fogel, M. L., Macko, S. A., and Wehmiller, J. F., eds. 2000. Perspectives in amino acid and protein geochemistry. Oxford University Press, Oxford.
Greenwood, D. R. 1991. The taphonomy of plant macrofossils. Pp. 141169 in Donovan, S. K., ed. The processes of fossilization. Columbia University Press, New York.
Griffiths, D. 1998. Sampling effort, regression method, and the shape and slope of size-abundance relations. Journal of Animal Ecology 67:795804.
Hall, P. 1992. Efficient bootstrap simulations. Pp. 127143 in Lepage, R. and Billard, L. L., eds. Exploring the limits of bootstrap. Wiley, New York.
Hearty, P. J., 1987. New data on the Pleistocene of Mallorca. Quaternary Science Reviews 6:245257.
Hendry, J. P., Ditchfield, P. W., and Marshall, J. D. 1995. Two stage metamorphism of Jurassic aragonitic bivalves: implications for early diagenesis. Journal of Sedimentary Research A65:214224.
Hendry, J. P., Trewin, N. H., and Fallick, A. E. 1996. Low Mgcalcite marine cement in Cretaceous turbidites: origin, spatial distribution, and relationship to sea-water chemistry. Sedimentology 43:877900.
Hughen, K. A., Baillie, M. G. L., Bard, E., Bayliss, A., Bertrand, C. J. H., Blackwell, P. G., Buck, C. E., Burr, G. S., Cutler, K. B., Damon, P. E., Edwards, R. L., Fairbanks, R. G., Friedrich, M., Guilderson, T. P., Kromer, B., McCormac, F. G., Manning, S. W., Bronk Ramsey, C., Reimer, P. J., Reimer, R. W., Remmele, S., Southon, J. R., Stuiver, M., Talamo, S., Taylor, F. W., van der Plicht, J., and Weyhenmeyer, C. E. 2004. Marine04 marine radiocarbon age calibration, 26–0 ka BP. Radiocarbon 46:10591086.
Ikemoto, T., and Takai, K. 2000. A new linearized formula for the law of total effective temperature and the evaluation of line-fitting methods with both variables subject to error. Environmental Entomology 29:671682.
Kaufman, D. S. 2000. Amino acid racemization in ostracodes. Pp. 145160 in Goodfriend, et al. 2000.
Kaufman, D. S., and Manley, W. F. 1998. A new procedure for determining DL amino acid ratios in fossils using reverse phase liquid chromatography. Quaternary Science Reviews 17:9871000.
Keil, R. G., Tsamakis, E., and Hedges, J. I. 2000. Early diagenesis of particulate amino acids in marine systems. Pp. 6982 in Goodfriend, et al. 2000.
Kidwell, S. M. 1993. Patterns of time averaging in the shallow marine fossil record. Pp. 275300 in Kidwell, and Behrensmeyer, 1993.
Kidwell, S. M. 1998. Time averaging in the marine fossil record: overview of strategies and uncertainties. Geobios 30:977995.
Kidwell, S. M., and Behrensmeyer, A. K., eds. 1993. Taphonomic approaches to time resolution in fossil assemblages. Short Courses in Paleontology 6. Paleontological Society, Knoxville, Tenn.
Kidwell, S. M., and Bosence, D. W. J. 1991. Taphonomy and time-averaging of marine shelly faunas. Pp. 116188 in Allison, P. A. and Briggs, D. E. G., eds. Taphonomy: releasing the data locked in the fossil record. Plenum Press, New York.
Kidwell, S. M., and Brenchley, P. J. 1994. Patterns in bioclastic accumulation through the Phanerozoic: Changes in input or in destruction? Geology 22:11391143.
Kidwell, S. M., and Brenchley, P. J. 1996. Evolution of the fossil record: thickness trends in marine skeletal accumulations and their implications. Pp. 290336 in Jablonski, D., Erwin, D. H., and Lipps, J. H., eds. Evolutionary paleobiology. University of Chicago Press, Chicago.
Kidwell, S. M., Best, M. M. R., and Kaufman, D. S. 2005. Taphonomic trade-offs in tropical marine death assemblages: differential time averaging, shell loss, and probable bias in siliciclastic vs. carbonate facies. Geology 33:729732.
Kimber, R. W. L., and Griffin, C. V. 2000. Interpretation of D/L amino acid data from agricultural soils. Pp. 195201 in Goodfriend, et al. 2000.
Kosnik, M. A., and Kaufman, D. S. 2008. Identifying outliers and assessing the accuracy of amino acid racemization measurements for use in geochronology. II. Data screening. Quaternary Geochronology 3:328341.
Kosnik, M. A., Hua, Q., Jacobsen, G. E., Kaufman, D. S., and Wüst, R. A. 2007. Sediment mixing and stratigraphic disorder revealed by the age-structure of Tellina shells in Great Barrier Reef sediment. Geology 35:811814.
Kosnik, M. A., Kaufman, D. S., and Hua, Q. 2008. Identifying outliers and assessing the accuracy of amino acid racemization measurements for use in geochronology. I. Age calibration curves. Quaternary Geochronology 3:308327.
Kosnik, M. A., Hua, Q., Kaufman, D. S., and Wüst, R. A. 2009. Taphonomic bias and time-averaging in tropical molluscan death assemblages: differential shell half-lives in Great Barrier Reef sediment. Paleobiology 35:565586.
Kowalewski, M. 1996a. Time-averaging, overcompleteness, and the geological record. Journal of Geology 104:317326.
Kowalewski, M. 1996b. Taphonomy of a living fossil: the lingulide brachiopod Glottidia palmeri Dall from Baja California, Mexico. Palaios 11:244265.
Kowalewski, M. 1997. The reciprocal taphonomic model. Lethaia 30:8688.
Kowalewski, M., and Bambach, R. K. 2003. The limits of paleontological resolution. Pp. 148 in Harries, P. J. ed. Approaches in high-resolution stratigraphic paleontology. Kluwer Academic/Plenum Publishers, New York.
Kowalewski, M., and Rimstidt, J. D. 2003. Average lifetime and age spectra of detrital grains: toward a unifying theory of sedimentary particles. Journal of Geology 111:427439.
Kowalewski, M., Goodfriend, G. A., and Flessa, K. W. 1998. High-resolution estimates of temporal mixing within shell beds: the evils and virtues of time-averaging. Paleobiology 24:287304.
Kowalewski, M., Serrano, G. E. A., Flessa, K. W., and Goodfriend, G. A. 2000. Dead delta's former productivity: two trillion shells at the mouth of the Colorado River. Geology 28:10591062.
Kowalewski, M., Simões, M. G., Carroll, M., and Rodland, D. L. 2002. Abundant brachiopods on a tropical upwelling-influenced shelf (Southeast Brazilian Bight, South Atlantic). Palaios 17:277286.
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 1:4365.
Lajoie, K. R., Wehmiller, J. F., and Kennedy, G. L. 1980. Inter- and intrageneric trends in apparent racemization kinetics of amino acids in Quaternary mollusks. Pp. 305340 in Hare, P. E., Hoering, T. C., and King, K. Jr., eds. Biogeochemistry of amino acids. Wiley, New York.
Lee, D. E. 1991. Aspects of the ecology and distribution of the living Brachiopoda of New Zealand. Pp. 273279 in MacKinnon, D. L., Lee, D. L., and Campbell, J. D., eds. Brachiopods through time. Balkema, Rotterdam.
Lever, J., Kessler, A., Van Overbeeke, A. P., and Thijssen, R. 1961. Quantitative beach research. II. The “hole effect.” Netherlands Journal of Sea Research 1:339358.
Mahiques, M. M. d. 1995. Sedimentary dynamics of the bays off Ubatuba, state of Sao Paolo. Boletim do Institute Oceanografico, São Paulo 43:111122.
Mahiques, M. M. d., Tessler, M. G., and Furtado, V. V. 1998. Characterization of energy gradient in enclosed bays of Ubatuba region, Southeastern Brazil. Estuarine, Coastal and Shelf Science 47:431446.
Mahiques, M. M., Tessler, M. G., Ciotti, A. M., da Silveira, I. C. A., Sousa, S. H. M., Figueria, R. C. L., Tassinari, C. C. G., Furtado, V. V., and Passos, R. F. 2004. Hydrodynamically-driven patterns of recent sedimentation in the shelf and upper slope off southeast Brazil. Continental Shelf Research 24:16851697.
Manceñido, M., and Griffin, M. 1988. Distribution and palaeoenvironmental significance of the genus Bouchardia (Brachiopoda, Terebratellidina): its bearing on the Cenozoic evolution of the South Atlantic. Revista Brasileira de Geosciências 18:201211.
Manley, W. F., Miller, G. H., and Czywezynski, J. 2000. Kinetics of aspartic acid racemization in Mya and Hiatella: modeling age and paleotemperature of high-latitude Quaternary mollusks. Pp. 202218 in Goodfriend, et al. 2000.
Manly, B. F. J. 1997. Randomization and Monte Carlo methods in biology, 2d ed. Chapman and Hall, London.
Mantelatto, F. L. M., and Fransozo, A. 1999. Characterization of the physical and chemical parameters of Ubatuba Bay, northern coast of São Paulo State, Brazil. Revista Brasileira de Biologia 59:2331.
Martin, L., Bittencourt, A. C. S. P., Dominguez, J. M. L., Flexor, J., and Suguio, K. 1998. Oscillations or not oscillations, that is the question: Comment on Angulo, R.J., and Lessa, G.C. “The Brazilian sea-level curves: a critical review with emphasis on the curves from the Paranaguá and Cananéia regions” [Mar. Geol. 140, 141–166]. Marine Geology 150:179187.
Martin, L., Dominguez, J. M. L., and Bittencourt, A. C. S. P. 2003. Fluctuating Holocene sea levels in eastern and southeastern Brazil: evidence from multiple fossil and geometric indicators. Journal of Coastal Research 19:101124.
Martin, R. E. 1999. Taphonomy: a process approach. Cambridge University Press, Cambridge.
Martin, R. E., Wehmiller, J. F., Harris, M. S., and Liddel, W. D. 1996. Comparative taphonomy of bivalves and foraminifera from Holocene tidal flat sediments, Bahía la Choya, Sonora, Mexico (northern Gulf of California): taphonomic grades and temporal resolution. Paleobiology 22:8090.
McCormac, F. G., Hogg, A. G., Blackwell, P. G., Buck, C. E., Higham, T. F. G., and Reimer, P. J. 2004. SHCa104 Southern hemisphere calibration 0–11.0 cal kyr BP. Radiocarbon 46:10871092.
McKittrick, M. A. 1987. Experiments on the settling of gastropod and bivalve shells: biostratinomic implications. In Flessa, K. W., ed. Paleoecology and taphonomy of Recent to Pleistocene intertidal deposits Gulf of California. Paleontological Society Special Publication 2:150163. Washington, D.C.
Meldahl, K. H., Flessa, K. W., and Cutler, A. H. 1997. Time-averaging and postmortem skeletal survival in benthic fossil assemblages: quantitative comparisons among Holocene environments. Paleobiology 23:207229.
Miller, B. B., and Hare, P. E. 1980. Amino acid geochronology: integrity of the carbonate matrix and potential of molluscan fossils. Pp. 415443 in Hare, P. E., Hoering, T. C., and King, K. Jr., eds. Biogeochemistry of amino acids. Wiley, New York.
Mitterer, R. M., and Kriausakul, N. 1989. Calculation of amino acid racemization ages based on apparent parabolic kinetics. Quaternary Science Reviews 8:353357.
Miller, B. B., McCoy, W. D., and Bleuer, N. K. 1987. Stratigraphic potential of amino acid ratios in Pleistocene terrestrial gastropods: an example from West-Central Indiana. Boreas 16:133138.
Morse, J. W., Zullig, J. J., Bernstein, L. D., Millero, F. J., Milne, P., Mucci, A., and Choppin, G. R. 1985. Chemistry of calcium-rich shallow water sediments in the Bahamas. American Journal of Science 285:147185.
Narchi, W., and Domaneschi, O. 1977. Semele casali – Jurado, 1949 (Mollusca, Bivalvia) in the Brazilian littoral. Studies on Neotropical Fauna and Environment 12:263272.
Olszewski, T. D. 1999. Taking advantage of time-averaging. Paleobiology 25:226238.
Olszewski, T. D. 2004. Modeling the influence of taphonomic destruction, reworking, and burial, on time-averaging in fossil accumulations. Palaios 19:3950.
Parsons-Hubbard, K.M., Callender, W. R., Powell, E. N., Brett, C. E., Walker, S. E., Raymond, A. L., and Staff, G. M. 1999. Rates of burial and disturbance of experimentally-deployed molluscs: implications for preservation potential. Palaios 14:337351.
Parsons-Hubbard, K. M., Powell, E. N., Staff, G. M., Callender, W. R., Brett, C. E., and Walker, S. E. 2001. The effect of burial on shell preservation and epibiont cover in Gulf of Mexico and Bahamas shelf and slope environments after two years: an experimental approach. In Aller, J. Y., Woodin, S. A., and Aller, R. C., eds. Organism-sediment interactions. Belle W. Baruch Library in Marine Science No. 21:297314. University of South Carolina Press, Columbia.
Pohlo, R. 1982. Evolution of the Tellinacea (Bivalvia). Journal of Molluscan Studies 48:245256.
Powell, E. N., Parsons-Hubbard, K. M., Callender, W. R, Staff, G. M., Rowe, G. T., Brett, C. E., Walker, S. E., Raymond, A., Carlson, D. D., White, S., and Heise, E. A. 2002. Taphonomy on the continental shelf and slope: two-year trends—Gulf of Mexico and Bahamas. Palaeogeography, Palaeoclimatology, Palaeoecology 184:135.
Powell, E. N., Callender, W. R, Staff, G. M., Parsons-Hubbard, K. M., Brett, C. E., Walker, S. E., Raymond, A., and Ashton-Alcox, K. A. 2008. Molluscan shell condition after eight years on the sea floor — Taphonomy in the Gulf of Mexico and Bahamas. Journal of Shellfish Research 27:191225.
Richardson, J. R. 1981. Brachiopods and pedicles. Paleobiology 7:8795.
Rodland, D. L., Kowalewski, M., Carroll, M., and Simões, M. G. 2004. Colonization of a “lost world” encrustation patterns in modern subtropical brachiopod assemblages. Palaios 19:381395.
Rodland, D. L., Kowalewski, M., Carroll, M., and Simões, M. G. 2006. The temporal resolution of epibiont assemblages; are they ecological snapshots or overexposures? Journal of Geology 114:313324.
Rodrigues, S. C. 2006. Tafonomia de moluscos bivalves e braquiópodes das enseadas de Ubatuba e Picinguaba, norte do Estado de São Paulo: implicações do uso de assinaturas tafonômicas no reconhecimento de gradientes ambientais. Ph.D. dissertation. University of São Paulo, Institute of Geosciences, São Paulo.
Rodrigues, S. C., Simóes, M. G., Kowalewski, M., Petti, M. A. V., Nonato, E. F., Martinez, S., and Del Rio, C. J. Biotic interaction between spionid polychaetes and bouchardiid brachiopods: Paleoecological, taphonomic, and evolutionary implications. Acta Palaeontologica Polonica 53:657668.
Sadler, P. M. 1981. Sediment accumulation and the completeness of stratigraphic sections. Journal of Geology 89:569584.
Sheskin, D. J. 2004. Handbook of parametric and nonparametric statistical procedures, 3d ed. Chapman and Hall, London.
Simões, M. G., Kowalewski, M., Mello, L. H. C., Rodland, D. L., and Carroll, M. 2004. Recent brachiopods from the Southern Brazilian Shelf: palaeontological and biogeographical implications. Palaeontology 47:515533.
Simóes, M. G., Rodrigues, S. C., Leme, J. M., and Bissaro, M. C. Junior. 2005. The settling pattern of brachiopod shells: stratigraphic and taphonomic implications to shell bed formation and paleoecology. Revista Brasileira de Geosciências 35:383391.
Simões, M. G., Rodrigues, S.C., and Kowalewski, M. 2009. Bouchardia rosea, a vanishing brachiopod species of the Brazilian platform: taphonomy, historical ecology, and conservation paleobiology. Historical Biology 21:123137.
Simões, M. G., Rodrigues, S. C., Neves, J. P., and Kowalewski, M. 2007. Experimental and field observations on the preservational bias of brachiopod valves in present-day assemblages of Bouchardia rosea from the Brazilian shelf: paleoecological implications. Geological Society of America Abstracts with Programs 39(2):96.
Soares-Gomes, A., and Pires-Vanin, A. M. S. 2003. Padrões de abundância, riqueza e diversidade de moluscos bivalves na plataforma continental ao largo de Ubatuba, São Paulo, Brasil: uma comparação metológica. Revista Brasileira de Zoologia, 20:717725.
Stead, R. A., Urrutia, G., Garrido, O., Clasing, E., Lardies, M. A., and Arratia, L. P. 2002. The significance of contrasting feeding strategies on the reproductive cycle in two coexisting tellinacean bivalves. Journal of the Marine Biological Association of the United Kingdom 82:443–53.
Stempien, J. A. 2005. Brachyuran taphonomy in a modern tidal-flat environment: preservation potential and anatomical bias. Palaios 20:400410.
Stuiver, M., Reimer, P. J., and Reimer, R. W. 2005. CALIB 5.0.2, radiocarbon calibration program,
Teece, M. A., Tuross, N., Kress, W. M., Peterson, P. M., Russell, G., and Fogel, M. L. 2000. Preservation of amino acids in museum herbarium samples. Pp. 8387 in Goodfriend, et al. 2000.
Teixeira, C., and Tundisi, J. G. 1981. The effects of nitrogen and phosphorous enrichment on phytoplankton in the region of Ubatuba. Solatum do Instituto Oceanografico São Paulo 30:7786.
Tomašových, A., and Rothfus, T. A. 2005. Differential taphonomy of modern brachiopods (San Juan Islands, Washington State): effect of intrinsic factors on damage and community-level abundance. Lethaia 38:271292.
Tomašových, A., Fürsich, F. T., and Olszewski, T. D. 2006a. Modeling shelliness and alteration in shell beds: variation in hardpart input and burial rates leads to opposing predictions. Paleobiology 32:278298.
Tomašových, A., Fürsich, F. T., and Wilmsen, M. 2006b. Preservation of autochthonous shell beds by positive feedback between increased hardpart-input rates and increased sedimentation rates. Journal of Geology 114:287312.
Tommasi, L. R. 1970. Sobre o braquiópode Bouchardia rosea (Mawe, 1823). Boletim do Instituto Oceanográfico 19:3-3.
Torello, F. F. 2004. Tafonomia experimental do fossil-vivo Bouchardia rosea (Brachiopoda, Terebratellidae) e suas aplicações em Paleontologia. Ph.D. dissertation. University of São Paulo, Institute of Geosciences, São Paulo.
Torello, F. F., and Simões, M. G. 2003. Comparative bivalve and brachiopod shell resistance to abrasion tested in a taphonomic tumbling barrel. Third Latin American Congress of Sedimentology, Belém 2003. Abstracts, pp. 223224.
Waite, E. R., and Collins, M. J. 2000. The interpretation of aspartic acid racemization of dentine proteins. Pp. 182194 in Goodfriend, et al. 2000.
Walker, K. R., and Bambach, R. K. 1971. The significance of fossil assemblages from fine-grained sediments: time-averaged communities. Geological Society of America Abstracts with Programs 3(7):783784.
Walker, S. E., and Voight, J. R. 1994. Paleoecologic and taphonomic potential of deep sea gastropods. Palaios 9:4859.
Wehmiller, J. F. 1982. A review of amino acid racemization studies in Quaternary mollusks: stratigraphic and chronologic applications in coastal and interglacial sites, Pacific and Atlantic coasts, United States, United Kingdom, Baffin Island and tropical islands. Quaternary Science Reviews 1:83120.
Wehmiller, J. F., and Miller, G. H. 2000. Aminostratigraphic dating methods in Quaternary geology. In Noller, J. S., Sowers, J. M., and Lettis, W. R., eds. Quaternary geochronology, methods and applications. American Geophysical Union Reference Shelf 4:187222.
Wehmiller, J. F., York, L. L., and Bart, M. L. 1995. Amino-acid racemization geochronology of reworked Quaternary mollusks on U.S. Atlantic coast beaches: implications for chronostratigraphy, taphonomy, and coastal sediment transport. Marine Geology 124:303337.
Williams, A., and Rowell, A. J. 1965. Morphology. Pp. H57H155 in Williams, A. et al. Brachiopoda 1. Part H of Moore, R. C. ed. Treatise on invertebrate paleontology. Geological Society of America, Boulder, Colo., and University of Kansas, Lawrence.
Williams, A., James, M. A., Emig, C. C., MacKay, S., and Rhodes, M. C. 1997. Anatomy. Pp. 7188 in Williams, A. et al. Brachiopoda 1. Part H (revised) of Kaesler, R. L., ed. Treatise on invertebrate paleontology. Geological Society of America, Boulder, Colo., and University of Kansas, Lawrence.
Wilson, M. V. H. 1988. Taphonomic processes: information loss and information gain. Geoscience Canada 15 (Paleoscene No. 9):131148.
Wing, S. L., and DiMichele, W. A. 1995. Conflict between local and global changes in plant diversity through geological time. Palaios 10:551564.
Wright, P., Cherns, L., and Hodges, P. 2003. Missing molluscs: field testing taphonomic loss in the Mesozoic through early large-scale aragonite dissolution. Geology 31:211214.
Yanes, Y., Kowalewski, M., Ortiz, J. E., Castillo, C., Torres, T., and Nuez, J. 2007. Scale and structure of time-averaging (age mixing) in terrestrial gastropod assemblages from Quaternary eolian deposits of the eastern Canary Islands. Palaeogeography, Palaeoclimatology, Palaeoecology 251:283299.
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