Hostname: page-component-8448b6f56d-xtgtn Total loading time: 0 Render date: 2024-04-24T19:32:51.551Z Has data issue: false hasContentIssue false

Implications of intracolonial variation in a Paleozoic bryozoan

Published online by Cambridge University Press:  14 July 2015

Eric J. Holdener
Affiliation:
Department of Biology, Kenyon College, Gambier, OH 43022
Steven J. Hageman
Affiliation:
Department of Geology, Field Museum of Natural History, Chicago, IL 60605

Abstract

Relative differences between environmentally controlled variation and genetically controlled variation are important when investigating morphologic variation in general, especially when establishing species concepts. The colonial nature of bryozoans provides a means for distinguishing between the two sources; variation can be partitioned into within-colony (microenvironmental) and among-colony (environmental + genetic) components. For the Paleozoic order Cryptostomata, biologically and taxonomically significant morphologic characters are well defined and methods for recognizing morphotaxa are well established.

The importance of within-colony variation to the morphometric treatment of fenestrate species was assessed after the discovery of a large specimen of Hemitrypa sp. Variation within the colony was compared to variation among and within two congeneric species. The distribution of study segments across the colony allowed assessment of variation both along the growth axis and laterally between segments of approximately equivalent generational age. Repeatability of methods was assessed using data measured independently from identical segments by three workers.

Variation within the large colony is less than variation among congeneric species, indicating that genetic differences among species exceed variation resulting from combined phenotypic and genotypic sources within species. Neither astogenetic nor ontogenetic morphologic gradients are recognized. Variation between data collected from identical segments by pairs of workers falls within the range of variation for the entire colony. Thus, multiple workers can reproduce data to the finest level of meaningful resolution. Cryptostome morphospecies concepts are validated.

The potential for partitioning genotypic versus environmental variation in reduced, multidimensional morphospace is reinforced. Studies of microevolution and speciation may be designed that account for these factors.

Type
Research Article
Copyright
Copyright © The Paleontological Society 

Access options

Get access to the full version of this content by using one of the access options below. (Log in options will check for institutional or personal access. Content may require purchase if you do not have access.)

References

Anstey, R. L., and Perry, T. G. 1970. Biometric procedures in taxonomic studies of Paleozoic bryozoans. Journal of Paleontology, 44:383398.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., Cheetham, A. H., and Cook, P. L. 1983. Introduction to the Bryozoa, p. 348. In Robinson, R. A., (ed.), Treatise on Invertebrate Paleontology, Part G, Bryozoa Revised. The Geological Society of America and the University of Kansas, Boulder, Colorado and Lawrence, Kansas.Google Scholar
Brande, S., and Bretsky, S. S. 1982. Avoid improper statistical analysis in bryozoans: analysis of variance is suitable for study of hierarchical variation. Journal of Paleontology, 56:12071212.Google Scholar
Cheetham, A. H., Jackson, J. B. C., and Hayek, L. C. 1993. Quantitative genetics of bryozoan phenotypic evolution. I. Rate tests for random change versus selection in differentiation of living species. Evolution, 47:15261538.Google ScholarPubMed
Cheetham, A. H., Jackson, J. B. C., and Hayek, L. C. 1994. Quantitative genetics of bryozoan phenotypic evolution. II. Analysis of selection and random change in fossil species using reconstructed genetic parameters. Evolution, 48:360375.Google ScholarPubMed
Cheetham, A. H., Jackson, J. B. C., and Hayek, L. C. 1995. Quantitative genetics of bryozoan phenotypic evolution. III. Phenotypic plasticity and the maintenance of genetic variation. Evolution, 49:290296.Google ScholarPubMed
Elias, M. K. 1964. Stratigraphy and paleoecology of some Carboniferous bryozoans. Cinquieme Congres International de Stratigraphie et de Geologie du Carbonifere, Compte Rendu, 1:375382.Google Scholar
Hageman, S. J. 1991. Approaches to systematic and evolutionary studies of perplexing groups: an example using fenestrate Bryozoa. Journal of Paleontology, 65:630647.CrossRefGoogle Scholar
Hageman, S. J. 1993. Effects of nonnormality on studies of morphologic variation of a rhabdomesine bryozoan, Streblotrypa (Streblascopora) prisca (Gabb and Horn). The University of Kansas Paleontological Contributions, New Series, Number 4, 13 p.Google Scholar
Hageman, S. J. 1994. Microevolutionary implications of clinal variation in the Paleozoic bryozoan Streblotrypa. Lethaia, 27:209222.CrossRefGoogle Scholar
Hageman, S. J. 1995. Observed phenotypic variation in a Paleozoic bryozoan. Paleobiology, 21:314328.CrossRefGoogle Scholar
Hageman, S. J., and Blake, D. B. 1992. Microenvironmental effects on intracolonial morphology in Bryozoa. Geological Society of America Abstracts with Programs, 24(7):A99.Google Scholar
Holdener, E. H. 1994. Numerical taxonomy of fenestrate bryozoans: evaluation of methodologies and recognition of intraspecific variation. Journal of Paleontology, 68:12011214.CrossRefGoogle Scholar
Key, M. M. Jr. 1987. Partitioning of morphological variation across stability gradients in Upper Ordovician trepostomes, p. 145152. In Ross, J. R. P. (ed.), Bryozoa: Present and Past. Western Washington University, Bellingham.Google Scholar
McKinney, F. K. 1980. The Devonian fenestrate bryozoan Utropora Pocta. Journal of Paleontology, 54:241252.Google Scholar
McKinney, F. K., and Boardman, R. S. 1985. Zooidal biometry of Stenolaemata, p. 193203. In Nielsen, C. and Larwood, G. P. (eds.), Bryozoa: Ordovician to Recent. Olsen and Olsen, Fredensborg, Denmark.Google Scholar
McKinney, F. K., and Kriz, J. 1986. Lower Devonian Fenestrata (Bryozoa) of the Prague Basin, Barrandian area, Bohemia, Czechoslovakia. Fieldiana, Geology, New Series, Number 15, 90 p.Google Scholar
McKinney, F. K., and Stedman, T. G. 1981. Constancy of characters within helical portions of Archimedes, p. 151157. In Larwood, G. P. and Nielsen, C. (eds.), Recent and Fossil Bryozoa. Olsen and Olsen, Fredensborg, Denmark.Google Scholar
McKinney, F. K., Taylor, P. D., and Zullo, V. A. 1993. Lyre-shaped hornerid bryozoan colonies: homeomorphy in colony form between Paleozoic Fenestrata and Cenozoic Cyclostomata. Journal of Paleontology, 67:343354.CrossRefGoogle Scholar
Miller, T. G. 1962. On Hemitrypa hibernica M'Coy. Geological Magazine, 99:313321.CrossRefGoogle Scholar
Neff, N. A., and Marcus, L. F. 1980. A Survey of Multivariate Methods for Systematics. Privately published, New York, 243 p.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., 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
Schopf, T. J. M. 1976. Environmental versus genetic causes of morphologic variability in bryozoan colonies from the deep sea. Paleobiology, 2:156165.CrossRefGoogle Scholar
Snyder, E. M. 1984. Taxonomy, functional morphology, and paleoecology of the Fenestellidae and Polyporidae (Fenestelloidea, Bryozoa) of the Warsaw Formation (Valmeyeran, Mississippian) of the Mississippi Valley. Unpublished , , 802 p.Google Scholar
Snyder, E. M. 1991. Revised taxonomic procedures and paleoecological applications for some North American Mississippian Fenestellidae and Polyporidae (Bryozoa). Palaeontographica Americana, 57, 275 p.Google Scholar
Stedman, T. G. 1982. Astogeny of fenestrate bryozoans and their potential use in biostratigraphy. Unpublished , , 129p.Google Scholar
Stratton, J. F., and Horowitz, A. S. 1977. Astogenetic variability in a frond of Polypora laevinodata (Hall). Proceedings of the Indiana Academy of Sciences, 86:290292.Google Scholar
Taylor, P. D., and Furness, R. W. 1978. Astogenetic and environmental variation of zooid size within colonies of Jurassic Stomatopora (Bryozoa, Cyclostomata). Journal of Paleontology, 52:10931102.Google Scholar
Winston, J. E. 1977. Feeding in marine bryozoans, p. 233271. In Woollacott, R. M. and Zimmer, R. L. (eds.), Biology of Bryozoans. Academic Press, New York.CrossRefGoogle Scholar