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Changes in colonial development, intraspecific heterochrony, morphological integration, and character heritabilities in two populations of the bryozoan species Batostoma jamesi from the Kope Formation (Upper Ordovician, Cincinnatian)

Published online by Cambridge University Press:  14 July 2015

Joseph F. Pachut
Affiliation:
Department of Geology, Indiana University-Purdue University at Indianapolis, Indianapolis 46202-5132,
Margaret M. Fisherkeller
Affiliation:
Indiana State Museum, Indianapolis 46204,

Abstract

Populations of the Upper Ordovician trepostome bryozoan Batostoma jamesi were collected from two different paleoenvironmental settings in the Kope Formation of southeastern Indiana. Within each colony and population, morphologic changes were analyzed during colony growth, or astogeny. Morphological measurements of zooecia, mesozooecia, and acanthostyles display similar patterns of change during colony growth in both populations but magnitudes are generally larger in the high diversity population.

Canonical variates analyses provided multivariate confirmation of univariate character differences found within each population. Statistically significant multivariate morphological differences between growth stages persist even if assignments of colonies to populations are ignored. Results suggest different potentials for altering growth trajectories in different environments with early growth stage flexibility in colonies from lower diversity settings and later-stage flexibility in colonies from higher diversity settings.

Heterochronic changes occur between species populations. Relative to the high-diversity population, the low-diversity population displays the following: 1) progenesis and hypermorphosis for zooecia, reflecting the ability to exist over a broader range of areal densities and surface areas than in populations from high-diversity associations; 2) postdisplacement and progenesis for mesozooecia, producing mature mesozooecial densities earlier in growth and at smaller sizes while the onset of mesozooecial development is delayed; and 3) acceleration, predisplacement, and progenesis for acanthostyles, resulting in a more rapid rate of development, an earlier onset of style development and more styles, and an earlier time of maturation, respectively.

The estimated level of morphological integration is higher in the high diversity population regardless of stage of colony growth. Within populations, integration is stronger during early growth stages in colonies from high diversity settings and during later growth stages in colonies from low diversity settings. Character heritabilities are high in both diversity-level populations, suggesting that these patterns of morphological integration were not the result of non-heritable phenotypic plasticity. Mean heritability is greater in the high diversity population and differs statistically only between the late growth stages of populations. Patterns of morphological integration may result from differing levels of stabilizing selection in different environments. Depending on the timing of selection, these different levels of integration are capable of affecting the outcome of selection on species populations.

Type
Research Article
Copyright
Copyright © The Paleontological Society

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References

Alberch, P. 1980. Ontogenesis and morphological diversification. American Zoologist, 20:653667.CrossRefGoogle Scholar
Alberch, P. 1982. Developmental constraints in evolutionary processes, p. 313332. In Bonner, J. T. (ed.), Evolution and Development. Springer-Verlag, New York.CrossRefGoogle Scholar
Alberch, P., and Alberch, J. 1981. Heterochronic mechanisms of morphological diversification and evolutionary change in the neotropical salamander Bolitoglossa occidentalis (Amphibia: Plethodontidae). Journal of Morphology, 167:249264.CrossRefGoogle Scholar
Alberch, P., Gould, S. J., Oster, G. F., and Wake, D. B. 1979. Size and shape in ontogeny and phylogeny. Paleobiology, 5:296317.CrossRefGoogle Scholar
Anstey, R. L. 1987. Astogeny and phylogeny: evolutionary heterochrony in Paleozoic bryozoans. Paleobiology, 13:2043.CrossRefGoogle Scholar
Anstey, R. L., and Bartley, J. W. 1984. Quantitative stereology: an improved thin section biometry for bryozoans and other colonial organisms. Journal of Paleontology, 58:612625.Google Scholar
Anstey, R. L., and Perry, T. G. 1973. Eden Shale bryozoans: a numerical study (Ordovician, Ohio Valley). Michigan State University, Publications of the Museum, Paleontological Series, 1:180.Google Scholar
Anstey, R. L., Pachut, J. F., and Prezbindowski, D. R. 1976. Morphogenetic gradients in Paleozoic bryozoan colonies. Paleobiology, 2:131146.CrossRefGoogle Scholar
Boardman, R. S. 1954. Morphologic variation and mode of growth of Devonian trepostomatous Bryozoa. Science, 120:322.CrossRefGoogle ScholarPubMed
Boardman, R. S. 1960. Trepostomatous Bryozoa of the Hamilton Group of New York State. U.S. Geological Survey Professional Paper 340, 87 p.CrossRefGoogle Scholar
Boardman, R. S., Ceetham, A. H., and Cook, P. L. 1970. Intracolony variation and the genus concept in Bryozoa. North American Paleontological Convention, Chicago, 1969, Proceedings C, p. 294320.Google Scholar
Boardman, R. S., Cheetham, A. H., Blake, D. B., Utgaard, J., Karklins, O. L., Cook, P. L., Sandberg, P. A., Lutaud, G., and Wood, T. S. 1983. Part G, Bryozoa, Revised. In Robison, R. A. (ed.), Treatise on Invertebrate Paleontology. Geological Society of America and University of Kansas Press, Lawrence, 625 p.Google Scholar
Cheetham, A. H., Jackson, J. B. C., and Hayek, L.-A. 1993. Quantitative genetics of bryozoan phenotypic evolution. I. Rate tests for random change versus selection in differentiation of living species. Evolution, 47:15261538.CrossRefGoogle ScholarPubMed
Cheetham, A. H., Jackson, J. B. C., and Hayek, L.-A. 1995. Quantitative genetics of bryozoan phenotypic evolution. III. Phenotypic plasticity and the maintenance of genetic variation. Evolution, 49:290296.CrossRefGoogle ScholarPubMed
Cheverud, J. M. 1982. Phenotypic, genetic, and environmental morphological integration in the cranium. Evolution, 36:499516.CrossRefGoogle ScholarPubMed
Cheverud, J. M. 1984. Quantitative genetics and developmental constraints on evolution by selection. Journal of Theoretical Biology, 110:155171.CrossRefGoogle ScholarPubMed
Cheverud, J. M. 1988. A comparison of genetic and phenotypic correlations. Evolution, 42:958968.CrossRefGoogle ScholarPubMed
Cheverud, J. M., Rutledge, J. J., and Atchley, W. R. 1983. Quantitative genetics of development: genetic correlations among age-specific values and the evolution of ontogeny. Evolution, 37:895905.Google ScholarPubMed
Cheverud, J. M., Wagner, G. P., and Dow, M. M. 1989. Methods for the comparative analysis of variation patterns. Systematic Zoology, 38:201213.CrossRefGoogle Scholar
Crow, J. F. 1986. Basic Concepts in Population, Quantitative, and Evolutionary Genetics. W. H. Freeman and Company, New York, 273 p.Google Scholar
Cumings, E. R. 1904. Development of some Paleozoic Bryozoa. American Journal of Sciences, 17:4978.Google Scholar
Cumings, E. R. 1912. Development and systematic position of the monticuliporoids. Geological Society of America Bulletin, 23:357370.CrossRefGoogle Scholar
Delmet, D. A., and Anstey, R. L. 1974. Fourier analysis of morphologic plasticity within an Ordovician bryozoan colony. Journal of Paleontology, 48:217226.Google Scholar
Dodd, J. R., and Stanton, R. J. Jr. 1990. Paleoecology, Concepts and Applications. John Wiley and Sons, New York, 502 p.Google Scholar
Goodman, D. 1975. The theory of diversity-stability relationships in ecology. Quarterly Review of Biology, 50:237266.CrossRefGoogle Scholar
Gould, S. J. 1977. Ontogeny and Phylogeny. Belknap Press, Cambridge, Massachusetts, 501 p.Google Scholar
Gould, S. J. 1982. Change in developmental timing as a mechanism of macroevolution, p. 333346. In Bonner, J. T. (ed.), Evolution and Development. Springer-Verlag, New York.CrossRefGoogle Scholar
Gould, S. J. 1989. A developmental constraint in Cerion, with comments on the definition and interpretation of constraint in evolution. Evolution, 43:516539.Google Scholar
Hageman, S. J., Bayer, M. M., and Todd, C. D. 1999. Partitioning phenotypic variation: genotypic, environmental and residual components from bryozoan skeletal morphology. Journal of Natural History, 33:17131735.CrossRefGoogle Scholar
Hickey, D. R. 1988. Bryozoan astogeny and evolutionary novelties: their role in the origin and systematics of the Ordovician monticuliporid trepostome genus Peronopora . Journal of Paleontology, 62:180203.CrossRefGoogle Scholar
Horowitz, A. S., and Pachut, J. F. 1989. In search of the elusive earliest stages of colony development in the Paleozoic bryozoan order Cystoporida—an aid in phylogenetic reconstruction? Geological Society of America, Annual Meeting, Abstracts with Programs, 22:113.Google Scholar
Huston, M. 1979. A general hypothesis of species diversity. American Naturalist, 113:81101.CrossRefGoogle Scholar
Jackson, J. B. C., and Cheetham, A. H. 1990. Evolutionary significance of morphospecies: a test with cheilostome Bryozoa. Science, 248:579583.CrossRefGoogle ScholarPubMed
Kachigan, S. K. 1991. Multivariate Statistical Analysis. Radius Press, New York, 303 p.Google Scholar
Key, M. M. Jr. 1990. A new family of trepostome bryozoans from the Ordovician Simpson Group of Oklahoma. Journal of Paleontology, 64:700724.CrossRefGoogle Scholar
Lande, R. 1980. The genetic covariance between characters maintained by pleiotropic mutation. Genetics, 94:203215.CrossRefGoogle Scholar
Lande, R. 1982. A quantitative genetic theory of life history evolution. Ecology, 63:607615.CrossRefGoogle Scholar
Lande, R. 1984. The genetic correlation between characters maintained by selection, linkage and inbreeding. Genetical Research, 44:309320.CrossRefGoogle ScholarPubMed
Lande, R. 1986. The dynamics of peak shifts and the pattern of morphological evolution. Paleobiology, 12:343354.CrossRefGoogle Scholar
Lande, R., and Arnold, S. J. 1983. The measurement of selection on correlated characters. Evolution, 37:12101226.CrossRefGoogle ScholarPubMed
Maderson, P. F. A., Alberch, P., Goodwin, B. C., Gould, S. J., Hoffman, A., Murray, J. D., Raup, D. M., de Ricqles, A., Seilacher, A., Wagner, G. P., and Wake, D. B. 1982. The role of development in macroevolutionary change, p. 279312. In Bonner, J. T. (ed.), Evolution and Development. Springer-Verlag, New York.Google Scholar
McKinney, F. K. 1977. Autozooecial budding patterns in dendroid Paleozoic bryozoans. Journal of Paleontology, 51:303329.Google Scholar
McKinney, F. K. 1978. Astogeny of the lyre-shaped Carboniferous fenestrate bryozoan Lyroporella . Journal of Paleontology, 52:8390.Google Scholar
McKinney, M. L. 1984. Allometry and heterochrony in an Eocene echinoid lineage: morphological change as a by-product of size selection. Paleobiology, 10:407419.CrossRefGoogle Scholar
McKinney, M. L. 1986. Ecological causation of heterochrony: a test and implications for evolutionary theory. Paleobiology, 12:282289.CrossRefGoogle Scholar
McNamara, K. J. 1986. A guide to the nomenclature of heterochrony. Journal of Paleontology, 60:413.CrossRefGoogle Scholar
Meyer, A. 1987. Phenotypic plasticity and heterochrony in Chiclasoma managuense (Pisces, Cichlidae) and their implications for speciation in cichlid fishes. Evolution, 41:13571369.Google ScholarPubMed
Nickles, J. M. 1902. The geology of Cincinnati (Ohio). Cincinnati Society of Natural History Journal, 20:49110.Google Scholar
Olson, E. C., and Miller, R. L. 1958. Morphological Integration. University of Chicago Press, Chicago.Google Scholar
Pachut, J. F. 1982. Morphologic variation within and among genotypes in two Devonian bryozoan species: an independent indicator of paleostability? Journal of Paleontology, 56:703716.Google Scholar
Pachut, J. F. 1987. Population genetics of four species of Ordovician bryozoans: Stereology and jackknifed analysis of variance. Journal of Paleontology, 61:927941.CrossRefGoogle Scholar
Pachut, J. F. 1989. Heritability and intraspecific heterochrony in Ordovician bryozoans from environments differing in diversity. Journal of Paleontology, 63:182194.CrossRefGoogle Scholar
Pachut, J. F. 1992. Morphological integration and covariance during astogeny of an Ordovician trepostome bryozoan from communities of different diversities. Journal of Paleontology, 66:750757.CrossRefGoogle Scholar
Pachut, J. F., and Anstey, R. L. 1979. A developmental explanation of stability-diversity-variation hypotheses: morphogenetic regulation in Ordovician bryozoan colonies. Paleobiology, 5:168187.CrossRefGoogle Scholar
Pachut, J. F., and Cuffey, R. J. 1991. Clinal variation, intraspecific heterochrony, and microevolution in the Permian bryozoan Tabulipora carbonaria . Lethaia, 24:165185.CrossRefGoogle Scholar
Pachut, J. F., Cuffey, R. J., and Anstey, R. L. 1991. The concepts of astogeny and ontogeny in stenolaemate bryozoans, and their illustration in colonies of Tabulipora carbonaria from the Lower Permian of Kansas. Journal of Paleontology, 65:213233.CrossRefGoogle Scholar
Perry, T. G., and Hattin, D. E. 1958. Astogenetic study of fistuliporoid bryozoans. Journal of Paleontology, 32:10391050.Google Scholar
Pielou, E. C. 1974. Population and Community Ecology. Gordon & Breach Science Publishers, New York, 424 p.Google Scholar
Podell, M. E., and Anstey, R. L. 1979. The interrelationship of early colony development, monticules and branches in Paleozoic bryozoans. Palaeontology, 22:965982.Google Scholar
Russ, J. C. 1986. Practical Stereology. Plenum Press, New York, 185 p.CrossRefGoogle Scholar
Smith, J. M., Burian, R., Kauffman, S., Alberch, P., Campbell, J., Goodwin, B., Lande, R., Raup, D., and Wolpert, L. 1985. Developmental constraints and evolution. Quarterly Review of Biology, 60:265287.CrossRefGoogle Scholar
Tissot, B. N. 1988. Geographic variation and heterochrony in two species of cowries (Genus Cypraea). Evolution, 42:103117.Google Scholar
Zaret, T. M. 1982. The stability/diversity controversy: a test of hypotheses. Ecology, 63:721731.CrossRefGoogle Scholar
Zelditch, M. L. 1988. Ontogenetic variation in patterns of phenotypic integration in the laboratory rat. Evolution, 42:2841.CrossRefGoogle ScholarPubMed
Zelditch, M. L., and Carmichael, A. C. 1989. Ontogenetic variation in patterns of developmental and functional integration in skulls of Sigmodon fulviventer . Evolution, 43:814824.CrossRefGoogle ScholarPubMed
Zelditch, M. L., Straney, D. O., Swiderski, D. L., and Carmichael, A. C. 1990. Variation in developmental constraints in Sigmodon . Evolution, 44:17381747.CrossRefGoogle ScholarPubMed