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Cladistic and phenetic recognition of species in the Ordovician bryozoan genus Peronopora

Published online by Cambridge University Press:  20 May 2016

Robert L. Anstey  and
Joseph F. Pachut
Robert L. Anstey
Affiliation:
Department of Geological Sciences, Michigan State University, East Lansing 48824–1115,
Joseph F. Pachut
Affiliation:
Department of Geology, Indiana University-Purdue University at Indianapolis 46202–5132,
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Abstract

We compare cladistic, phenetic, stratophenetic, and typological methods of recognizing species within a single well-sampled, long-lived Ordovician bryozoan genus, Peronopora Nicholson, 1881. A consensus of 11 methods recognizes 14 species within Peronopora, each of which has both cladistic and phenetic support, and support by at least four different methods. Comparison of methods was based on: 1) the number of species recognized; 2) the degree of splitting of consensus groups; 3) consistency in determining first and last appearance datums, and stratigraphic ranges; 4) consistency in the number of specimens assigned to species; 5) stability in recognizing geographic distributions; and 6) correlations with character heritability. The five methods most closely approaching consensus were: 1) Ward clustering on Euclidean distance; 2) K-means splitting; 3) Ward clustering on correlation coefficients; 4) cladistic parsimony; and 5) cladistic parsimony plus iterative discriminant analysis. The three methods farthest from consensus were: 1) stratophenetics; 2) average linkage on Euclidean distance; and 3) conventional typology. Cladistic parsimony is the only method that can recognize all 14 consensus clusters. We argue that the overall advantages of parsimony analysis outweigh the merits of the various phenetic approaches in recognizing paleontological species. We also argue that given sufficient allochrony in sampling, cladistic structure is detectable both within and among species. Recognizing species based on fixed character states would require at least 72 species in our material.

The crown groups of Pachut and Anstey (2002) are herein recognized as monophyletic species in the sense of Mishler and Theriot (2000), one of which we designate as a new species, P. browni. Six of our eight nonmonophyletic stem groups are recognized in this paper as metaspecies in the sense of Donoghue (1985) and Gauthier et al. (1988). Two of our stem groups are phenetically indistinguishable from phyletically contiguous crown groups, and we attribute the failure of parsimony analysis to recognize the monophyly of these groups to homoplasy.

Type
Research Article
Information
Journal of Paleontology , Volume 78 , Issue 4 , July 2004 , pp. 651 - 674
Copyright
Copyright © The Paleontological Society 

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References

Adrain, J. M., and Westrop, S. R. 2001. Stratigraphy, phylogeny, and species sampling in time and space, p. 291–322. In Adrain, J. M., Edgecombe, G. D., and Lieberman, B. S. (eds.), Fossils, Phylogeny, and Form: An Analytical Approach. Kluwer Academic, New York.Google Scholar
Anstey, R. L. 1981. Zooid orientation structures and water flow patterns in Paleozoic bryozoan colonies. Lethaia, 14:287–302.Google Scholar
Anstey, R. L. 1987a. Astogeny and phylogeny: evolutionary heterochrony in Paleozoic bryozoans. Paleobiology, 13:20–43.Google Scholar
Anstey, R. L. 1987b. Colony patterning and functional morphology of water flow in Paleozoic stenolaemate bryozoans, p. 1–8. In Ross, J. R. P. (ed.), Bryozoa: Present and Past. Western Washington University, Bellingham.Google 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, 58:612–625.Google Scholar
Anstey, R. L., and Perry, T. G. 1969. Redescription of cotypes of Peronopora vera, a Cincinnatian (Late Ordovician) ectoproct species. Journal of Paleontology, 43:245–251.Google Scholar
Anstey, R. L., and Perry, T. G. 1970. Biometric procedures in taxonomic studies of Paleozoic bryozoans. Journal of Paleontology, 44:383–398.Google Scholar
Anstey, R. L., and Perry, T. G. 1973. Eden Shale bryozoans: a numerical study (Ordovician, Ohio Valley). Publications of the Museum, Michigan State University (Paleontological Series), 1(1): 1–80.Google Scholar
Bassler, R. S. 1953. Bryozoa, p. G1–G253. In Moore, R. C. (ed.), Treatise on Invertebrate Paleontology, Pt. G, Bryozoa. Geological Society of America and University of Kansas Press, Lawrence.Google Scholar
Bell, M. A. 2002. Speciation and morphological change, p. 1160–1165. In Singer, R. (ed.), Encyclopedia of Paleontology. Fitzroy Dearborn Publishers, Chicago.Google Scholar
Boardman, R. S. 1983. General features of the Class Stenolaemata, p. 49–137. In Boardman, R. S. et al. (eds.), Treatise on Invertebrate Paleontology, Pt. G, Bryozoa (revised). Geological Society of America and University of Kansas Press, Lawrence.Google Scholar
Boardman, R. S., and Utgaard, J. 1966. A revision of the Ordovician bryozoan genera Monticulipora, Peronopora, Heterotrypa, and Dekayia. Journal of Paleontology, 40:1082–1108.Google Scholar
Brochu, C. A., and Norell, M. A. 2001. Time and trees: quantitative assessment of temporal congruence in the bird origins debate, p. 511–535. In Gauthier, J. A. and Gall, L. F. (eds.), New Perspectives on the Origin and Evolution of Birds: Proceedings of the International Symposium in Honor of John H. Ostrom. Peabody Museum of Natural History, Yale University, New Haven.Google Scholar
Brown, G. D., and Daly, E. J. 1985. Trepostome Bryozoa from the Dillsboro Formation (Cincinnatian) in southeastern Indiana. Indiana Department of Natural Resources Geological Survey Special Report, 33:1–95.Google Scholar
Cheetham, A. H. 1986. Tempo of evolution in a Neogene bryozoan: rates of morphologic change within and across species boundaries. Paleobiology, 12:190–202.CrossRefGoogle 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, 47:1526–1538.Google Scholar
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, 49:290–296.Google Scholar
Cracraft, J. 1983. Species concepts and speciation analysis. Current Ornithology, 1:159–187.Google Scholar
Cuffey, R. J. 1967. Bryozoan Tabulipora carbonaria in Wreford Megacyclothem (Lower Permian) of Kansas. The University of Kansas Paleontological Contributions Article, 1:1–96.Google Scholar
Cumings, E. R., and Galloway, J. J. 1913. The stratigraphy and paleontology of the Tanner's Creek sections of the Cincinnatian Series of Indiana. Indiana Department of Geology and Natural Resources Annual Report, 37:353–479.Google Scholar
Donoghue, M. J. 1985. A critique of the biological species concept and recommendations for a phylogenetic alternative. Bryologist, 88:172–181.Google Scholar
Ehrenberg, C. G. 1845. Neue untersuchungen das kleinste leben als geologisches moment. Bericht Akademie der Wissenschaften, p. 53–88.Google Scholar
Eldredge, N. 1993. What, if anything, is a species?, p. 3–20. In Kimbel, W. H. and Martin, L. B. (eds.), Species, Species Concepts, and Primate Evolution. Plenum, New York.Google Scholar
Eldredge, N., and Cracraft, J. 1980. Phylogenetic Patterns and the Evolutionary Process. Columbia University Press, New York, 349 p.Google Scholar
Eldredge, N., and Gould, S. J. 1972. Punctuated equilibria: an alternative to phyletic gradualism, p. 82–115. In Schopf, T. J. M. (ed.), Models in Paleobiology. Freeman, Cooper, San Francisco.Google Scholar
Erwin, D. H., and Anstey, R. L. 1995. Speciation in the fossil record, p. 11–38. In Erwin, D. H. and Anstey, R. L. (eds.), New Approaches to Speciation in the Fossil Record. Columbia University Press, New York.Google Scholar
Gauthier, J., Kluge, A., and Rowe, T. 1988. Amniote phylogeny and the importance of fossils. Cladistics, 4:105–209.Google Scholar
Gingerich, P. D. 1979. Populations, phylogeny, and classification: an example from the mammalian fossil record. Systematic Zoology, 28:451–464.Google Scholar
Gould, S. J. 2002. The Structure of Evolutionary Theory. Belknap Press of Harvard University Press, Cambridge, 1433 p.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, 33:1713–1735.Google Scholar
Hammer, O. 2003. PAST 1.15. http://folk.uio.no/ohammer/past.Google Scholar
Hennig, W. 1966. Phylogenetic Systematics. University of Illinois Press, Urbana, 263 p.Google 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, 62:180–203.Google Scholar
Jackson, J. B. C., and Cheetham, A. H. 1990. Evolutionary significance of morphospecies: a test with cheilostome Bryozoa. Science. 248:579–583.Google Scholar
James, U. P. 1884. Descriptions of four new species of fossils from the Cincinnati Group. Journal of the Cincinnati Society of Natural History, 7:137–139.Google Scholar
Karklins, O. L. 1984. Trepostome and cystoporate bryozoans from the Lexington Limestone and the Clays Ferry Formation (Middle and Upper Ordovician) of Kentucky. United States Geological Survey Professional Paper, 1066–1:1–105.Google Scholar
Kimbel, W. H., and Martin, L. B. 1993. Species and speciation. Conceptual issues and their relevance for primate evolutionary biology, p. 539–553. In Kimbel, W. H. and Martin, L. B. (eds.), Species, Species Concepts, and Primate Evolution. Plenum, New York.Google Scholar
Kimbel, W. H., and Rak, Y. 1993. The importance of species taxa in paleoanthropology and an argument for the phylogenetic concept of the species category, p. 461–484. In Kimbel, W. H. and Martin, L. B. (eds.), Species, Species Concepts, and Primate Evolution. Plenum, New York.Google Scholar
Knowlton, N., and Budd, A. F. 2001. Recognizing coral species present and past, p. 97–119. 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
Krishtalka, L. 1993. Anagenetic angst. Species boundaries in Eocene primates, p. 331–344. In Kimbel, W. H. and Martin, L. B. (eds.), Species, Species Concepts, and Primate Evolution. Plenum, New York.Google Scholar
Mayr, E. 1969. Principles of Systematic Zoology. McGraw-Hill, New York, 428 p.Google Scholar
Mayr, E. 2000. The biological species concept, p. 17–29, 93–100, 161–166. In Wheeler, Q. D. and Meier, R. (eds.), Species Concepts and Phylogenetic Theory. A Debate. Columbia University Press, New York.Google Scholar
Meier, R., and Willman, R. 2000. The Hennigian species concept, p. 30–43, 101–118, 167–178. In Wheeler, Q. D. and Meier, R. (eds.), Species Concepts and Phylogenetic Theory. A Debate. Columbia University Press, New York.Google Scholar
Mishler, B. D., and Theriot, E. C. 2000. The phylogenetic species concept (sensu Mishler and Theriot): monophyly, apomorphy, and phylogenetic species concepts, p. 44–54, 119–132, 179–184. In Wheeler, Q. D. and Meier, R. (eds.), Species Concepts and Phylogenetic Theory. A Debate. Columbia University Press, New York.Google Scholar
Nicholson, H. A. 1881. On the Structure and Affinities of the Genus Monticulipora and Its Subgenera. William Blackwood and Sons, Edinburgh, 235 p.Google Scholar
Nickles, J. M. 1905. The Upper Ordovician rocks of Kentucky and their Bryozoa. Kentucky Geological Survey Bulletin, 5:1–64.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, 56:703–716.Google Scholar
Pachut, J. F. 1989. Heritability and intraspecific heterochrony in Ordovician bryozoans from environments differing in diversity. Journal of Paleontology, 63:182–194.Google Scholar
Pachut, J. F., and Anstey, R. L. 1984. The relative information content of Fourier-structural, binary (presence-absence) and combined data sets: a test using the H. A. Nicholson Collection of Paleozoic stenolaemate bryozoans. Journal of Paleontology, 58:1296–1311.Google Scholar
Pachut, J. F., and Anstey, R. L. 2002. Phylogeny, systematics, and biostratigraphy of the Ordovician bryozoan genus Peronopora. Journal of Paleontology, 76:607–637.Google Scholar
Pachut, J. F., Anstey, R. L., and Horowitz, A. S. 1994. The H. A. Nicholson Collection of Paleozoic stenolaemate bryozoans: comparison of cladistic and phenetic classifications. Journal of Paleontology, 68:978–994.Google Scholar
Page, R. D. M. 2001. TREEVIEW (WIN32) 1.6.6. http://taxonomy.zoology.gla.ac.uk/rod/rod.html.Google Scholar
Rominger, C. 1866. Observations on Chaetetes and some related genera, in regard to their systematic position; with an appended description of some new species. Proceedings of the National Academy of Sciences (Philadelphia), 1866:113–123.Google Scholar
Rose, K. D., and Brown, T. M. 1993. Species concepts and species recognition in Eocene primates, p. 299–330. In Kimbel, W. H. and Martin, L. B. (eds.), Species, Species Concepts, and Primate Evolution. Plenum, New York.Google Scholar
Savage, T. E. 1924. Richmond rocks of Iowa and Illinois. American Journal of Science, 8:411–427.Google Scholar
Savage, T. E. 1925. Correlation of the Maquoketa and Richmond rocks of Iowa and Illinois. Illinois Academy of Science Transactions, 17:233–247.Google Scholar
Simpson, G. G. 1951. The species concept. Evolution, 5:285–298.Google Scholar
Simpson, G. G. 1961. Principles of Animal Taxonomy. Columbia University Press, New York, 247 p.Google Scholar
Singh, R. J. 1979. Trepostomatous bryozoan fauna from the Bellevue Limestone, Upper Ordovician, in the tri-state area of Ohio, Indiana, and Kentucky. Bulletins of American Paleontology, 76:159–288.Google Scholar
Smith, A. B. 1994. Systematics and the Fossil Record: Documenting Evolutionary Patterns. Blackwell Scientific Publications, Oxford, 223 p.Google Scholar
Smith, A. B. 2000. Stratigraphy in phylogeny reconstruction. Journal of Paleontology, 74:763–766.Google Scholar
Sumrall, C. D., and Brochu, C. A. 2003. Resolution, sampling, higher taxa and assumptions in stratocladistic analysis. Journal of Paleontology, 77:189–194.Google Scholar
Sweet, W. 1984. Graphic correlation of upper Middle and Upper Ordovician rocks, North American Midcontinent Province, U.S.A., p. 23–36. In Bruton, D. L. (ed.), Aspects of the Ordovician System. Universitetsforlaget, Oslo.Google Scholar
Swofford, D. L. 2000. PAUP∗. Phylogenetic Analysis Using Parsimony (∗ and Other Methods). Version 4.0b4a. Sinauer Associates, Sunderland, Massachusetts.Google Scholar
SYSTAT 8.0. 1998. SPSS, Inc., Chicago.Google Scholar
Theriot, E. 1992. Clusters, species concepts, and morphological evolution of diatoms. Systematic Biology, 41:141–157.Google Scholar
Ulrich, E. O. 1879. Descriptions of new genera and species of fossils from the Lower Silurian about Cincinnati. Journal of the Cincinnati Society of Natural History, 2:8–30.Google Scholar
Ulrich, E. O. 1882. American Paleozoic Bryozoa. Journal of the Cincinnati Society of Natural History, 5:121–175.Google Scholar
Ulrich, E. O. 1888. A correlation of the Lower Silurian horizons of Tennessee and of the Ohio and Mississippi Valleys with those of New York and Canada. American Geologist, 2:39–44.Google Scholar
Utgaard, J., and Perry, T. G. 1964. Trepostomatous bryozoan fauna of the upper part of the Whitewater Formation (Cincinnatian) of eastern Indiana and western Ohio. Indiana Department of Conservation Geological Survey Bulletin, 33:1–111.Google Scholar
Vrana, P., and Wheeler, W. 1992. Individual organisms as terminal entities: laying the species problem to rest. Cladistics, 8:67–72.Google Scholar
de Regny, P. Vinassa 1920. Sulla classificazione dei Treptostomidi. Atti della Societa Italiana di Scienze Naturali, 59:212–231.Google Scholar
Wheeler, Q. D., and Platnick, N. I. 2000. The phylogenetic species concept (sensu Wheeler and Platnick), p. 55–69, 133–145, 185–197. In Wheeler, Q. D. and Meier, R. (eds.), Species Concepts and Phylogenetic Theory. A Debate. Columbia University Press, New York.Google Scholar
Wiley, E. O., and Mayden, R. L. 2000. The evolutionary species concept, p. 70–92, 146–160, 198–208. In Wheeler, Q. D. and Meier, R. (eds.), Species Concepts and Phylogenetic Theory. A Debate. Columbia University Press, New York.Google Scholar
Willman, R. 1981. Evolution, systematik und stratigraphische bedeutung der neogenen susswassergastropoden von Rhodos und Kos/Agais. Palaeontographica, Abteilung A, 174:10–235.Google Scholar
Willman, R. 1985. Responses of the Plio-Pleistocene freshwater gastropods of Kos (Greece, Aegean Sea) to environmental changes, p. 295–321. In Bayer, U. and Seilacher, A. (eds.), Sedimentary and Evolutionary Cycles. Springer, New York.Google Scholar
Zechman, F. W., Zimmer, E. A., and Theriot, E. C. 1994. Use of ribosomal DNA internal transcribed spaces for phylogenetic studies in diatoms. Journal of Phycology, 30:507–512.Google Scholar