Skip to main content Accessibility help

Multiple paleoecological controls on the composition of marine fossil assemblages from the Frasnian (Late Devonian) of Virginia, with a comparison of ordination methods

  • Andrew M. Bush (a1) and Roderic I. Brame (a2)


Ecological ordination can reveal gradients in the species composition of fossil assemblages that can be correlated with paleoenvironmental gradients. Ordinations of simulated data sets suggest that nonmetric multidimensional scaling (NMDS) generally produces less distorted results than detrended correspondence analysis (DCA). We ordinated 113 brachiopod-dominated samples from the Frasnian (Late Devonian) Brallier, Scherr, and lower Foreknobs Formations of southwest Virginia, which represent a range of siliciclastic marine paleoenvironments. A clear environmental signal in the ordination results was obscured by (apparently) opportunistic species that occurred at high abundance in multiple environments; samples dominated by these species aggregated in ordination space regardless of paleoenvironmental provenance. After the opportunist-dominated samples were removed, NMDS revealed a gradient in species composition that was highly correlated with substrate (grain size); a second, orthogonal gradient likely reflects variation in disturbance intensity or frequency within grain-size regimes. Additional environmental or ecological factors, such as oxygenation, may also be related to the gradients. These two gradients, plus the environmental factors that controlled the occurrence of opportunistic species, explain much of the variation in assemblage composition in the fauna. In general, the composition of fossil assemblages is probably influenced by multiple paleoecological and paleoenvironmental factors, but many of these can be decomposed and analyzed.



Hide All
Alexander, R. 1977. Growth, morphology and ecology of Paleozoic and Mesozoic opportunistic species of brachiopods from Idaho-Utah. Journal of Paleontology 51:11331149.
Aseltine-Neilson, D. A., Bernstein, B. B., Palmer-Zwahlen, M. L., Riege, L. E., and Smith, R. W. 1999. Comparisons of turf communities from Pendleton Artificial Reef, Torrey Pines Artificial Reef, and a natural reef using multivariate techniques. Bulletin of Marine Science 65:3757.
Austin, M. P., and Noy-Meir, I. 1971. The problem of nonlinearity in ordination: experiments with two-gradient models. Journal of Ecology 59:763773.
Bambach, R. K., and Bennington, J B. 1996. Do communities evolve? A major question in evolutionary paleoecology. Pp. 123160 in Jablonski, D., Erwin, D. H., and Lipps, J., eds. Evolutionary paleobiology: essays in honor of James W. Valentine. University of Chicago Press, Chicago.
Bassett, M. G. 1984. Life strategies of Silurian brachiopods. Special Papers in Palaeontology 32:237263.
Bennington, J B., and Bambach, R. K. 1996. Statistical testing for paleocommunity recurrence: are similar fossil assemblages ever the same? Palaeogeography, Palaeoclimatology, Palaeoecology 127:107133.
Bergen, M., Weisberg, S. B., Smith, R. W., Cadien, D. B., Dalkey, A., Montagne, D. E., Stull, J. K., Velarde, R. G., and Ranasinghe, J. A. 2001. Relationship between depth, sediment, latitude, and the structure of benthic infaunal assemblages on the mainland shelf of southern California. Marine Biology 138:637647.
Bizzarro, M. 1995. The Middle Devonian chonetoidean brachiopods from the Hamilton Group of New York. Documents des Laboratoires de Géologie de Lyon 136:149189.
Bonelli, J. R. Jr., Brett, C. E., Miller, A. I., and Bennington, J. B. 2006. Testing for faunal stability across a regional biotic transition: quantifying stasis and variation among recurring coral-rich biofacies in the Middle Devonian Appalachian Basin. Paleobiology 32:2037.
Bowen, Z. P., Sutton, R. G., McAlester, A. L., and Rhoads, D. C. 1970. Upper Devonian deltaic environments. In Heaslip, W. G., ed. Annual Meeting of the New York State Geological Association, Guidebook 42:B1B11.
Bowen, Z. P., Rhoads, D. C., and McAlester, A. L. 1974. Marine benthic communities in the Upper Devonian of New York. Lethaia 7:93120.
Boyce, R. L., and Ellison, P. C. 2001. Choosing the best similarity index when performing fuzzy set ordination on binary data. Journal of Vegetation Science 12:711720.
Boyer, D. L., and Droser, M. L. 2009. Palaeoecological patterns within the dysaerobic biofacies: Examples from Devonian black shales of New York state. Palaeogeography, Palaeoclimatology, Palaeoecology 276:206216.
Bradfield, G. E., and Kenkel, N. C. 1987. Nonlinear ordination using flexible shortest path adjustment of ecological distances. Ecology 68:750753.
Brame, R. I. 2001. Revision of the Upper Devonian in the central-southern Appalachian Basin: biostratigraphy and lithostratigraphy. , Virginia Tech, Blacksburg, Va.
Brett, C. E., Hendy, A. J. W., Bartholomew, A. J., Bonelli, J. R. Jr., and McLaughlin, P. I. 2007a. Response of shallow marine biotas to sea-level fluctuations: a review of faunal replacement and the process of habitat tracking. Palaios 22:228244.
Brett, C. E., Bartholomew, A. J., and Baird, G. C. 2007b. Biofacies recurrence in the Middle Devonian of New York State: an example with implications for evolutionary paleoecology. Palaios 22:306324.
Brunton, H. 1972. The shell structure of chonetacean brachiopods and their ancestors. Bulletin of the British Museum (Natural History) Geology 21:126.
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 253:490508.
Bush, A. M., and Daley, G. M. 2008. Comparative paleoecology of fossils and fossil assemblages. In Kelley, P. H. and Bambach, R. K., eds. From evolution to geobiology: research questions driving paleontology at the start of a new century. Paleontological Society Paper 14:289317.
Clarke, K. R. 1993. Non-parametric multivariate analyses of changes in community structure. Australian Journal of Ecology 18:117143.
Clarke, K. R., and Warwick, R. M. 2001. Change in marine communities: an approach to statistical analysis and interpretation, 2d ed. Primer-E, Plymouth, U.K.
Copper, P. 1966. Ecological distribution of Devonian atrypid brachiopods. Palaeogeography, Palaeoclimatology, Palaeoecology 2:245266.
Copper, P. 1967. Adaptations and life habits of Devonian atrypid brachiopods. Palaeogeography, Palaeoclimatology, Palaeoecology 3:363379.
Copper, P. 1998. Evaluating the Frasnian-Famennian mass extinction: comparing brachiopod faunas. Acta Palaeontologica Polonica 43:137154.
Day, J., and Over, D. J. 2002. Post-extinction survivor fauna from the lowermost Famennian of eastern North America. Acta Palaeontologica Polonica 47:189202.
De'ath, G. 1999. Extended dissimilarity: a method of robust estimation of ecological distances from high beta diversity data. Plant Ecology 144:191199.
Dean, S. L., Kulander, B. R., and Skinner, J. M. 1988. Structural chronology of the Alleghanian Orogeny in southeastern West Virginia. Geological Society of America Bulletin 100:299310.
Dennison, J. M. 1970. Stratigraphic divisions of the Upper Devonian Greenland Gap Group (“Chemung Formation”) along Allegheny Front in West Virginia, Maryland, and Highland County, Virginia. Southeastern Geology 12:5382.
Dennison, J. M., Filer, J. K., and Rossbach, T. J. 1996. Devonian strata of southeastern West Virginia and adjacent Virginia. Pp. 354 in Dennison, J. M., ed. Geologic field guide to Devonian hydrocarbon stratigraphy of southeastern West Virginia and adjacent Virginia. Appalachian Geological Society, Charleston, W. Va.
Diener, D. R., Fuller, S. C., Lissner, A., Haydock, C. I., Maurer, D., Robertson, G., and Gerlinger, T. 1995. Spatial and temporal patterns of the infaunal community near a major ocean outfall in Southern California. Marine Pollution Bulletin 30:861878.
Dominici, S., and Kowalke, T. 2007. Depositional dynamics and the record of ecosystem stability: Early Eocene faunal gradients in the Pyrenean foreland, Spain. Palaios 22:268284.
Faith, D. P., Minchin, P. R., and Belbin, L. 1987. Compositional dissimilarity as a robust measure of ecological distance. Vegetatio 69:5768.
Fasham, M. J. R. 1977. A comparison of nonmetric multidimensional scaling, principal components and reciprocal averaging for the ordination of simulated coenoclines, and coenoplanes. Ecology 58:551561.
Filer, J. K. 2002. Late Frasnian sedimentation cycles in the Appalachian Basin: possible evidence for high frequency eustatic sea-level changes. Sedimentary Geology 154:3152.
Finnegan, S., and Droser, M. L. 2008. Reworking diversity: effects of storm deposition on evenness and sampled richness, Ordovician of the Basin and Range, Utah and Nevada, USA. Palaios 23:8796.
Fürsich, F. T., and Hurst, J. M. 1974. Environmental factors determining the distribution of brachiopods. Palaeontology 17:879900.
Goldman, D., and Mitchell, C. E. 1990. Morphology, systematics, and evolution of Middle Devonian Ambocoeliidae (Brachiopoda), western New York. Journal of Paleontology 64:7999.
Greiner, H. 1957. “Spirifer disjunctus”: its evolution and paleoecology in the Catskill Delta. Peabody Museum of Natural History, Yale University, Bulletin 11.
Harrington, J. W. 1970. Benthic communities of the Genesee Group (Upper Devonian). In Heaslip, W. G., ed. Annual Meeting of the New York State Geological Association, Guidebook 42:A1A18.
Hendy, A. J. W., and Kamp, P. J. J. 2007. Paleoecology of Late Miocene-Early Pliocene sixth-order glacioeustatic sequences in the Manutahi-1 core, Wanganui-Taranaki Basin, New Zealand. Palaios 22:325337.
Hill, M. O., and Gauch, H. G. Jr. 1980. Detrended correspondence analysis: an improved ordination technique. Vegetatio 42:4758.
Holland, S. M. 1995. The stratigraphic distribution of fossils. Paleobiology 21:92109.
Holland, S. M. 2003. Confidence limits on fossil ranges that account for facies changes. Paleobiology, 29:468479.
Holland, S. M. 2005. The signatures of patches and gradients in ecological ordinations. Palaios 20:573580.
Holland, S. M., and Patzkowsky, M. E. 2007. Gradient ecology of a biotic invasion: biofacies of the type Cincinnatian Series (Upper Ordovician), Cincinnati, Ohio region, USA. Palaios 22:392407.
Holland, S. M., Miller, A. I., Meyer, D. L., and Dattilo, B. F. 2001. The detection and importance of subtle biofacies within a single lithofacies: the Upper Ordovician Kope Formation of the Cincinnati, Ohio region. Palaios 16:205217.
Kendall, D. G. 1971. Seriation from abundance matrices. Pp. 215252 in Hodson, F. R., Kendall, D. G., and Tautu, P., eds. Mathematics in the archeological and historical sciences. Edinburgh University Press, Edinburgh.
Kenkel, N. C., and Orlóci, L. 1986. Applying metric and nonmetric multidimensional scaling to ecological studies: some new results. Ecology 67:919928.
Kirchgessner, D. A. 1973. Sedimentology and petrology of upper Devonian Greenland Gap Group along the Allegheny Front, Virginia, West Virginia and Maryland. Ph.D. dissertation. University of North Carolina, Chapel Hill.
Kruskal, J. B. 1964a. Multidimensional scaling by optimizing goodness of fit to a nonmetric hypothesis. Psychometrika 29:127.
Kruskal, J. B. 1964b. Nonmetric multidimensional scaling: a numerical method. Psychometrika 29:115129.
Leighton, L. R. 2000. Environmental distribution of spinose brachiopods from the Devonian of New York: test of the soft-substrate hypothesis. Palaios 15:184193.
Levinton, J. S. 1970. The paleoecologic significance of opportunistic species. Lethaia 3:6978.
Linsley, D. M. 1994. Devonian paleontology of New York. Paleontology Research Institution Special Publication 21.
Lundegard, P. D., Samuels, N. D., and Pryor, W. A. 1985. Upper Devonian turbidite sequence, central and southern Appalachian basin: contrasts with submarine fan deposits. Pp. 107121 in Woodrow, and Sevon, 1985.
McAlester, A. L. 1962. Upper Devonian pelecypods of the New York Chemung Stage. Peabody Museum of Natural History, Yale University, Bulletin 16.
McCune, B., and Grace, J. B. 2002. Analysis of ecological communities. MjM Software Design. Gleneden Beach, Ore.
McGhee, G. R. Jr. 1976. Late Devonian benthic marine communities of the central Appalachian Allegheny Front. Lethaia 9:111136.
McGhee, G. R. Jr. 1977. The Frasnian-Famennian (Late Devonian) boundary within the Foreknobs Formation, Maryland and West Virginia. Geological Society of America Bulletin 88:806808.
McGhee, G. R. Jr., and Dennison, J. M. 1976. The Red Lick Member, a new subdivision of the Foreknobs Formation (Upper Devonian) in Virginia, West Virginia, and Maryland. Southeastern Geology 18:4957.
McGhee, G. R. Jr., and Dennison, J. M. 1980. Late Devonian chronostratigraphic correlations between the central Allegheny Front and central and western New York. Southeastern Geology 21:279286.
McGhee, G. R. Jr., and Sutton, R. G. 1981. Late Devonian marine ecology and zoogeography of the central Appalachians and New York. Lethaia 14:2743.
McGhee, G. R. Jr., and Sutton, R. G. 1983. Evolution of late Frasnian (Late Devonian) marine environments in New York and the central Appalachians. Alcheringa 7:921.
McGhee, G. R. Jr., and Sutton, R. G. 1985. Late Devonian marine ecosystems of the lower West Falls Group in New York. Pp. 199209 in Woodrow, and Sevon, 1985.
McLaughlin, P. I., and Brett, C. E. 2007. Signatures of sea-level rise on the carbonate margin of a Late Ordovician foreland basin: a case study from the Cincinnati Arch, USA. Palaios 22:245267.
Miller, A. I. 1988. Spatial resolution in subfossil remains: implications for paleobiological analysis. Paleobiology 14:91103.
Miller, A. I., Holland, S. M., Meyer, D. L., and Datillo, B. F. 2001. The use of faunal gradient analysis for intraregional correlation and assessment of changes in sea-floor topography in the type Cincinnatian. Journal of Geology 109:603613.
Minchin, P. R. 1987a. An evaluation of the relative robustness of techniques for ecological ordination. Vegetatio 69:89107.
Minchin, P. R. 1987b. Simulation of multidimensional community patterns: towards a comprehensive model. Vegetatio 71:145156.
Minchin, P. R. 1989. Montane vegetation of the Mt. Field massif, Tasmania: a test of some hypotheses about properties of community patterns. Vegetatio 83:97110.
Muir-Wood, H. M. 1962. On the morphology and classification of the brachiopod suborder Chonetoidea. British Museum (Natural History), London.
Noy-Meir, I., and Austin, M. P. 1970. Principal component ordination and simulated vegetational data. Ecology 51:551552.
Oksanen, J., and Minchin, P. R. 2002. Continuum theory revisited: what shape are species responses along ecological gradients? Ecological Modelling 157:119129.
R Development Core Team. 2007. R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna.
Racheboeuf, P. 1990. Les brachiopodes chonetacés dans les assemblages benthiques siluriens et devoniens. Palaeogeography, Palaeoclimatology, Palaeoecology 81:141171.
Redman, C. M., Leighton, L. R., Schellenberg, S. A., Gale, C. N., Nielsen, J. L., Dressler, D. L., and Klinger, M. K. 2007. Influence of spatiotemporal scale on the interpretation of paleocommunity structure: lateral variation in the Imperial Formation of California. Palaios 22:630641.
Roberts, J., and Ludwig, J. A. 1991. Riparian vegetation along current-exposure gradients in floodplain wetlands of the River Murray, Australia. Journal of Ecology 79:117127.
Rode, A. L., and Lieberman, B. S. 2004. Using GIS to unlock the interactions between biogeography, environment, and evolution in Middle and Late Devonian brachiopods and bivalves. Palaeogeography, Palaeoclimatology, Palaeogeography 211:345359.
Rudwick, M. J. S. 1970. Living and fossil brachiopods. Hutchinson University Library, London.
SAS Institute. 2009. SAS software version 9.2. SAS Institute Inc., Cary, N.C.
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 32:989992.
Smith, R. W., Bernstein, B. B., and Cimberg, R. L. 1988. Community-environmental relationships in the benthos: applications of multivariate analytical techniques. Pp. 247326 in Soule, D. F. and Kleppel, G. S., eds. Marine organisms as indicators. Springer, New York.
Smith, R. W., Bergen, M., Weisberg, S. B., Cadien, D., Dalkey, A., Montagne, D., Stull, J. K., and Velarde, R. G. 2001. Benthic response index for assessing infaunal communities on the southern California mainland shelf. Ecological Applications 11:10731087.
Springer, D. A., and Bambach, R. K. 1985. Gradient versus cluster analysis of fossil assemblages: a comparison from the Ordovician of southwestern Virginia. Lethaia 18:181198.
Rode, A. L. Stigall 2005. Systematic revision of the Middle and Late Devonian brachiopods Schizophoria (Schizophoria) and ‘Schuchertella’ from North America. Journal of Systematic Palaeontology 3:133167.
Rode, A. L. Stigall, and Lieberman, B. S. 2005. Using environmental niche modeling to study the Late Devonian biodiversity crisis. Pp. 93180 in Over, D. J., Morrow, J. R., and Wignall, P. B., eds. Understanding Late Devonian and Permian-Triassic biotic and climatic events: towards an integrated approach. Elsevier, Amsterdam.
Sutton, R. G., and McGhee, G. R. Jr. 1985. The evolution of Frasnian marine “community-types” of south central New York. Pp. 211224 in Woodrow, and Sevon, 1985.
Sutton, R. G., Bowen, Z. P., and McAlester, A. L. 1970. Marine shelf environments of the Upper Devonian Sonyea Group of New York. Bulletin of the Geological Society of America 81:29752992.
Swan, J. M. A. 1970. An examination of some ordination problems by use of simulated vegetational data. Ecology 51:89102.
Thayer, C. W. 1974. Marine paleoecology in the Upper Devonian of New York. Lethaia 7:121155.
Tomašových, A. 2006. Brachiopod and bivalve ecology in the Late Triassic (Alps, Austria): onshore-offshore replacements caused by variations in sediment and nutrient supply. Palaios 21:344368.
Virginia Division of Mineral Resources. 1993. Geologic map of Virginia, scale 1:500,000. Virginia Division of Mineral Resources.
Virtanen, R., Oksanen, J., Oksanen, L. and Razzhivin, V. Y. 2006. Broad-scale vegetation-environment relationships in Eurasian high-latitude areas. Journal of Vegetation Science 17:519528.
Webber, A. J. 2002. High-resolution faunal gradient analysis and an assessment of the causes of meter-scale cyclicity in the type Cincinnatian Series (Upper Ordovician). Palaios 17:545555.
Webber, A. J. 2005. The effects of spatial patchiness on the stratigraphic signal of biotic composition (Type Cincinnatian Series; Upper Ordovician). Palaios 20:3750.
Wildi, O. 1992. On the use of Mantel's statistic and flexible shortest path adjustment in the analysis of ecological gradients. Coenoses 7:91101.
Williamson, M. H. 1978. The ordination of incidence data. Journal of Ecology 66:911920.
Williamson, M. H. 1983. The land-bird community of Skokholm: ordination and turnover. Oikos 41:378384.
Woodrow, J. W., and Sevon, W. D., eds. 1985. The Catskill Delta. Geological Society of America Special Paper 201.
Zuschin, M., Harzhauser, M., and Mandic, O. 2007. The stratigraphic and sedimentologic framework of fine-scale faunal replacements in the Middle Miocene of the Vienna Basin (Austria). Palaios 22:285295.
Type Description Title
Supplementary materials

Bush and Brame supplementary material
Supplementary Material

 Unknown (24 KB)
24 KB

Multiple paleoecological controls on the composition of marine fossil assemblages from the Frasnian (Late Devonian) of Virginia, with a comparison of ordination methods

  • Andrew M. Bush (a1) and Roderic I. Brame (a2)


Full text views

Total number of HTML views: 0
Total number of PDF views: 0 *
Loading metrics...

Abstract views

Total abstract views: 0 *
Loading metrics...

* Views captured on Cambridge Core between <date>. This data will be updated every 24 hours.

Usage data cannot currently be displayed