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Metrarabdotos and Related Genera (Bryozoa: Cheilostomata) in the Late Paleogene and Neogene of Tropical America

Published online by Cambridge University Press:  11 August 2017

Alan H. Cheetham ,
Joann Sanner  and
Jeremy B. C. Jackson
Alan H. Cheetham
Affiliation:
Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, Washington, DC 20650, , Unit 647, 3101 Old Pecos Trail, Santa Fe, New Mexico, 87505
Joann Sanner
Affiliation:
Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, Washington, DC 20650, ,
Jeremy B. C. Jackson
Affiliation:
Geoscience Research Division, Scripps Institution of Oceanography, La Jolla, California 92093, Center for Tropical Ecology and Archeology, Smithsonian Tropical Research Institute, Box 2072, Balboa, Republic of Panama
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Abstract

Metrarabdotos Canu, 1914 and the related genera Escharoides Milne Edwards, 1836b, Adeonellopsis MacGillivray, 1886, and Reptadeonella Busk, 1884 were key taxa in the decline of Bryozoa with erect, arborescent colonies and concomitant increase in numbers of species with encrusting colonies in the late Paleogene and Neogene of tropical America. In particular, the abundance, continuity of occurrence, and diversity of Metrarabdotos before its decline have made this genus, over the past 20 years, a model taxon for detailed morphometric studies of fine-scale evolutionary tempo and mode. During the same period, significant new occurrences of both Metrarabdotos and its near relatives have been documented with detailed collections from tropical American areas not represented in the original studies.

In this paper we present a complete reanalysis of the original morphometric data (Cheetham, 1986a), incorporating the new tropical American material and comparative material of selected Metrarabdotos species from Europe and Africa in order to formalize the taxonomy of the genus, many species of which have been in open nomenclature since 1986, and to explore their possible phylogenetic relationships to each other and to Escharoides, Adeonellopsis, and Reptadeonella. The new analysis, with procedures slightly modified from those used in the original analysis, is based on more than twice the number of specimens and 20% more morphological characters. Although the results include changes in species assignments for 13% of the specimens in the original analysis, the pattern of intraspecific morphological stasis previously identified in the tropical American Neogene species, and thus the concomitant interpretation of evolutionary tempo and mode, are unaltered.

Cladistic analysis resulted in a single most parsimonious tree for the 22 tropical American Metrarabdotos species, arranged in two monophyletic crown groups and a paraphyletic stem group. The stem group, ranging from latest Eocene to late Early or early Middle Miocene in age, includes four species, two of which are new: M. aguilerai from Venezuela and M. hispaniolae from the Dominican Republic and Haiti. Crown group A comprises seven species occurring in deposits of Late Miocene to Recent age, four of which are new: M. arawakorum from Venezuela; M. boldi and M. saundersi from the Dominican Republic; and M. coatesi, which occurs in the Dominican Republic, Costa Rica, and Panama. Crown group B, comprising 11 species, ranges from Early Miocene to Recent and includes five new species: M. cubaguaense from Venezuela; M. vokesorum from the Dominican Republic, Costa Rica, and Panama; and M. tainorum, M. jungi, and M. lopezense from the Dominican Republic. Incorporation of six European and African (eastern Atlantic) species, ranging in age from Early Miocene to Recent, into cladograms with the tropical American species produced more variable results in terms of both numbers of alternative trees and the positions of the tropical American species in them. One of these species, M. thomseni from the Pleistocene of Greece, and possibly a second, M. cf. M. maleckii Cheetham, 1968 from the Miocene of the Czech Republic, are new. With one exception, the eastern Atlantic species were placed in more or less basal positions in one or the other of the crown groups, suggesting that, as hypothesized by Cheetham (1968), the “Old World” and “New World” species of Metrarabdotos represent largely independent radiations. The exception is the living species M. cookae Cheetham, 1968, from West Africa and the Cape Verde Islands, which is placed unequivocally in a terminal position in crown group B. Although the three groups of tropical American species correspond in part to subgenera of Metrarabdotos erected by Cheetham (1968), the paraphyly of the stem group and the unresolved identity of the type species of the subgenus apparently corresponding to crown group A suggest that formalizing the taxonomy of these groups is unwarranted at present.

A further cladistic analysis, incorporating the three groups of Metrarabdotos species and Escharoides, Adeonellopsis, and Reptadeonella, yielded two trees, both of which support Cheetham's (1968) hypothesis that Metrarabdotos is more closely related to Escharoides than to Adeonellopsis and Reptadeonella. Escharoides is represented in tropical American Neogene deposits by two species, one of which, E. guraboensis from the Dominican Republic, is new. Of the six tropical American Neogene species of Adeonellopsis, four are new: A. cribrospiramen and A. guraboensis from the Dominican Republic, A. limonensis from Costa Rica, and A. antilleana from Cuba and Jamaica. Three of the six tropical American Neogene species of Reptadeonella, R. buddae from the Dominican Republic, R. collinsae from the Dominican Republic, Panama, Costa Rica, and the southeastern United States, and R. buricaensis from Panama, are also new.

Type
Research Article
Information
Journal of Paleontology , Volume 81 , Issue S67: The Paleontological Society Memoir 67: Metrarabdotos and Related Genera (Bryozoa: Cheilostomata) in the Late Paleogene and Neogene of Tropical America , January 2007 , pp. 1 - 91
Copyright
Copyright © 2007, The Paleontological Society 

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References

Abildgaard, P. C. 1806. Descripserunt et tabulas dederunt P. C. Abildgaard, M. Vahl, J. S. Holten, J. Rathke, Volume 4, p. 1–46. In Müller, O. F. (ed.), Zoologia danica, seu Animalium Daniae et Norvegiae rariorum ac minus notorum descriptiones et historia. N. Mölleri avlae regiae typographic et filli, Havniae.Google Scholar
Banta, W. C. 1969. The body wall of cheilostome Bryozoa. II. Interzooidal communication organs. Journal of Morphology, 129:149–170.Google Scholar
Banta, W. C. 1975. Origin and early evolution of cheilostome Bryozoa, p. 565–582. In Pouyet, Simone (ed.), Bryozoa 1974. Doc. Lab. Geol. Fac. Sci. Lyon, hors-serie 3, fasc. 2.Google Scholar
Bassler, R. S. 1935. Bryozoa. Fossilium Catalogus. 1: Animalia, pars 67. W. Junk, 's-Gravenhage, 229 p.Google Scholar
Bassler, R. S. 1936. Nomenclatorial notes on fossil and recent Bryozoa. Journal of the Washington Academy of Sciences, 26:156–162.Google Scholar
Bishop, J. D. D. 1987. Type and figured material from the “Pliocene Bryozoa of the Low Countries” (Lagaaij, 1952) in the collection of the Royal Belgian Institute of Natural Sciences. Institute Royal des Sciences Naturelle de Belgique, Documents de Travail, 37:1–36.Google Scholar
Bishop, J. D. D., and Hayward, P. J. 1989. SEM atlas of type and figured material from Robert Lagaaij's “The Pliocene Bryozoa of the Low Countries” (1952). Mededelingen Rijks Geologische Dienst, 43: 1–64.Google Scholar
Blainville, H.-M. D. de. 1830. Dictionnaire des Sciences Naturelle, v. 60. Normant, Paris.Google Scholar
Boardman, R. S., and Cheetham, A. H. 1969. Skeletal growth, intracolony variation, and evolution in Bryozoa: A review. Journal of Paleontology, 43:205–233.Google Scholar
Boardman, R. S., and Cheetham, A. H. 1973. Degrees of colony dominance in stenolaemate and gymnolaemate Bryozoa, p. 121–220. In Boardman, R. S., Cheetham, A. H., and Oliver, W. A. Jr. (eds.), Animal Colonies: Development and Function Through Time. Dowden, Hutchinson, and Ross, Stroudsburg, Pennsylvania.Google Scholar
Bold, W. A. van den. 1963. Upper Miocene and Pliocene Ostracoda of Trinidad. Micropaleontology, 9:361–424.Google Scholar
Borg, F. 1940. The genus Tubiporella and on a new boring bryozoan. Zoologiska bidrag från Uppsala, 18:415–438.Google Scholar
Budd, A. F., and Coates, A. G. 1992. Nonprogressive evolution in a clade of Cretaceous Montastraea-like corals. Paleobiology, 18:425–446.Google Scholar
Buge, E., and Galopim de Carvalho, A. M. 1963. Révision du genre Metrarabdotos Canu 1914 (Bryozoa, Cheilostomata). Revista da Faculdade de Ciéncias de Lisboa, 11:137–196.Google Scholar
Buge, E., and Galopim de Carvalho, A. M. 1964. Nouvelles ovservations sur le genre Metrarabdotos Canu 1914 (Bryozoa, Cheilostomata). Garcia de Orta, 12:651–656.Google Scholar
Busk, G. 1854. Catalogue of Marine Polyzoa in the Collection of the British Museum, Pt. II, Cheilostomata (part). British Museum, London, p. 55–120.Google Scholar
Busk, G. 1859. A Monograph of the Fossil Polyzoa of the Crag. The Palaeontographical Society, London, 136 p.CrossRefGoogle Scholar
Busk, G. 1884. Report on the Polyzoa collected by H. M. S. Challenger during the years 1873–1876, Pt. I, The Cheilostomata. Report on the Scientific Results of the Voyage of H. M. S. Challenger during the years 1873–76, Zoology. Vol. 10. Pt. 30. 216 p.Google Scholar
Calvet, L. 1931. Bryozoaires provenant des campagnes du Prince Albert I de Monaco. Résultats des Campagnes scientifiques de Monaco, Number 83, p. 1–152.Google Scholar
Canu, F. 1907. Les Bryozoaires des terrains tertiaires des environs de Paris. Annales de Paléontologie, 2:137–160.Google Scholar
Canu, F. 1914. Les Bryozoaires fossils des terrains du Sud-Ouest de la France. Bulletin de la Société Géologique de France, série 4, 14:465–474.Google Scholar
Canu, F., and Bassler, R. S. 1919. Fossil Bryozoa of the West Indies, p. 73–102. In Vaughan, T. W. (ed.), Contributions to the Geology and Paleontology of the West Indies. Carnegie Institution of Washington Publication, 291.Google Scholar
Canu, F., and Bassler, R. S. 1920. North American Early Tertiary Bryozoa. United States National Museum Bulletin, 106, 879 p.Google Scholar
Canu, F., and Bassler, R. S. 1923. North American Later Tertiary and Quaternary Bryozoa. United States National Museum Bulletin, 125, 302 p.Google Scholar
Canu, F., and Bassler, R. S. 1927. Classification of the cheilostomatous Bryozoa. Proceedings of the United States National Museum, 69(14), 42 p.Google Scholar
Canu, F., and Bassler, R. S. 1928a. Fossil and Recent Bryozoa of the Gulf of Mexico region. Proceedings of the United States National Museum, 72(14), 199 p.Google Scholar
Canu, F., and Bassler, R. S. 1928b. Bryozoaires du Brésil. Bulletin de la Société des Sciences de Seine-et-Oise, 9:58–100.Google Scholar
Canu, F., and Bassler, R. S. 1929. Bryozoa of the Philippine Region. United States National Museum Bulletin, 100, 685 p.Google Scholar
Canu, F., and Bassler, R. S. 1930. The bryozoan fauna of the Galapagos Islands. Proceedings of the United States National Museum, 76(13), 78 p.Google Scholar
Cheetham, A. H. 1954. A new Early Cretaceous bryozoan from Texas. Journal of Paleontology, 28:177–184.Google Scholar
Cheetham, A. H. 1957. Eocene-Oligocene boundary, eastern Gulf Coast region. Transactions of the Gulf Coast Association of Geological Societies, 7:89–97.Google Scholar
Cheetham, A. H. 1963. Late Eocene Zoogeography of the Eastern Gulf Coast Region. Geological Society of America Memoir, 91, 113 p.Google Scholar
Cheetham, A. H. 1966. Cheilostomatous Polyzoa from the Upper Bracklesham Beds (Eocene) of Sussex. Bulletin of the British Museum (Natural History), Geology, 13(1), 115 p.Google Scholar
Cheetham, A. H. 1967. Paleoclimatic significance of the bryozoan Metrarabdotos . Transactions of the Gulf Coast Association of Geological Societies, 17:400–407.Google Scholar
Cheetham, A. H. 1968. Morphology and Systematics of the Bryozoan Genus Metrarabdotos . Smithsonian Miscellaneous Collections, 153(1), 121 p.Google Scholar
Cheetham, A. H. 1975. Taxonomic significance of autozooid size and shape in some early multiserial cheilostomes from the Gulf Coast of the U. S. A., p. 547–564. In Pouyet, Simone (ed.), Bryozoa 1974. Doc. Lab. Geol. Fac. Sci. Lyon, hors-serie 3, fasc. 2.Google Scholar
Cheetham, A. H. 1986a. Tempo of evolution in a Neogene bryozoan: Rates of morphologic change within and across species boundaries. Paleobiology, 12:190–202.Google Scholar
Cheetham, A. H. 1986b. Branching, biomechanics, and bryozoan evolution. Proceedings of the Royal Society, London, B, 228:151–171.Google Scholar
Cheetham, A. H. 1987. Tempo of evolution in a Neogene bryozoan: Are trends in single morphologic characters misleading? Paleobiology, 13: 286–296.Google Scholar
Cheetham, A. H. 2001a. 2.1.3 Evolutionary stasis versus change, p. 137–142. In Briggs, D. E. G. and Crowther, P. R. (eds.), Paleobiology II. Blackwell Science, Oxford.Google Scholar
Cheetham, A. H. 2001b. Evolutionary stasis and taxonomic stability: The bryozoan Metrarabdotos in tropical America. Program and Abstracts, North American Paleontological Convention 2001, Berkeley, California, p. 41.Google Scholar
Cheetham, A. H. 2002. Asexual propagation in the cheilostomes Metrarabdotos and Coscinopleura: Effects on genetic variation and larval productivity, p. 73–80. In Wyse Jackson, P. N., Buttler, C. J., and Spencer Jones, M. E. (eds.), Bryozoan Studies 2001, Proceedings of the Twelfth International Bryozoology Association Conference. A. A. Balkema, Lisse.Google Scholar
Cheetham, A. H., and Cook, P. L. 1983. General features of the class Gymnolaemata, p. 138–207. In Robison, R. A. (ed.), Treatise on Invertebrate Paleontology, Pt. G, Bryozoa (revised). Geological Society of America and University of Kansas Press, Lawrence.Google Scholar
Cheetham, A. H., and Deboo, P. B. 1963. A numerical index for biostratigraphic zonation in the mid-Tertiary of the eastern Gulf. Transactions of the Gulf Coast Association of Geological Societies, 13:139–147.Google Scholar
Cheetham, A. H., and Hayek, L. C. 1983. Geometric consequences of branching growth in adeoniform Bryozoa. Paleobiology, 9:240–260.Google Scholar
Cheetham, A. H., and Hayek, L. C. 1988. Phylogeny reconstruction in the Neogene bryozoan Metrarabdotos: A paleontologic evaluation of methodology. Historical Biology, 1:65–83.CrossRefGoogle Scholar
Cheetham, A. H., and Jackson, J. B. C. 1995. Process from pattern: Tests for selection versus random change in punctuated bryozoan speciation, p. 184–207. In Erwin, D. H. and Anstey, R. L. (eds.), New Approaches to Speciation in the Fossil Record. Columbia University Press, New York.Google Scholar
Cheetham, A. H., and Jackson, J. B. C. 1996. Speciation, extinction, and the decline of arborescent growth in Neogene and Quaternary Bryozoa of tropical America, p. 205–233. In Jackson, J. B. C., Budd, A. F., and Coates, A. G. (eds.), Evolution and Environment in Tropical America. University of Chicago Press, Chicago.Google Scholar
Cheetham, A. H., and Jackson, J. B. C. 1998. The fossil record of cheilostome Bryozoa in the Neogene and Quaternary of tropical America: Adequacy for phylogenetic and evolutionary studies, p. 227–242. In Donovan, S. K. and Paul, C. R. C. (eds.), The Adequacy of the Fossil Record. John Wiley and Sons, Chicester, England.Google Scholar
Cheetham, A. H., and Jackson, J. B. C. 2000. Neogene history of cheilostome Bryozoa in tropical America, p. 1–16. In Herrera, A. and Jackson, J. B. C. (eds.), Proceedings of the 11th International Bryozoology Association Conference. Allen Press, Lawrence, Kansas.Google Scholar
Cheetham, A. H., and Lorenz, D. M. 1976. A vector approach to size and shape comparisons among zooids in cheilostome bryozoans. Smithsonian Contributions to Paleobiology, number 29, 55 p.Google Scholar
Cheetham, A. H., and Sandberg, P. A. 1964. Quaternary Bryozoa from Louisiana mudlumps. Journal of Paleontology, 38:1013–1046.Google Scholar
Cheetham, A. H., and Thomsen, E. 1981. Functional morphology of arborescent animals: Strength and design of cheilostome bryozoan skeletons. Paleobiology, 7:355–383.Google Scholar
Cheetham, A. H., Hayek, L. C., and Thomsen, E. 1980. Branching structure in arborescent animals: Models of relative growth. Journal of Theoretical Biology, 85:335–369.Google Scholar
Cheetham, A. H., Hayek, L. C., and Thomsen, E. 1981. Growth models in fossil arborescent cheilostome bryozoans. Paleobiology, 7:68–86.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, 47:1526–1538.Google Scholar
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, 48:360–375.Google Scholar
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, 49:290–296.Google Scholar
Cheetham, A. H., Jackson, J. B. C., and Sanner, J. 2001. Evolutionary significance of sexual and asexual modes of propagation in Neogene species of the bryozoan Metrarabdotos in tropical America. Journal of Paleontology, 75:564–577.Google Scholar
Cheetham, A. H., Rucker, J. B., and Carver, R. E. 1969. Wall structure and mineralogy of the cheilostome bryozoan Metrarabdotos . Journal of Paleontology, 43:129–135.Google Scholar
Cheetham, A. H., Jackson, J. B. C., Sanner, J., and Ventocilla, Y. 1999. Neogene cheilostome Bryozoa of tropical America: Comparison and contrast between the Central American isthmus (Panama, Costa Rica) and the north-central Caribbean (Dominican Republic), p. 159–192. In Collins, L. S. and Coates, A. G. (eds.), A Paleobiotic Survey of the Caribbean Faunas from the Neogene of the Isthmus of Panama. Bulletins of American Paleontology, Number 357.Google Scholar
Cheetham, A. H., Sanner, J., Taylor, P. D., and Ostrovsky, A. N. 2006. Morphological differentiation of avicularia and the proliferation of species in mid-Cretaceous Wilbertopora Cheetham, 1954 (Bryozoa: Cheilostomata). Journal of Paleontology, 80:49–71.CrossRefGoogle Scholar
Coates, A. G. 1999a. Maps, p. 287–298. In Collins, L. S. and Coates, A. G. (eds.), A Paleobiotic Survey of the Caribbean Faunas from the Neogene of the Isthmus of Panama. Bulletins of American Paleontology, Number 357.Google Scholar
Coates, A. G. 1999b. Stratigraphic sections, p. 300–348. In Collins, L. S. and Coates, A. G. (eds.), A Paleobiotic Survey of the Caribbean Faunas from the Neogene of the Isthmus of Panama. Bulletins of American Paleontology, Number 357.Google Scholar
Collins, L. S. 2005. Panama Paleontology Project. $lthttp://www.fiu.edu/~collinsl/pppimagemapnew.htm$gtGoogle Scholar
Collins, L. S., and Coates, A. G. (eds.). 1999. A paleobiotic survey of Caribbean faunas from the Neogene of the Isthmus of Panama. Bulletins of American Paleontology, Number 357, 351 p.Google Scholar
Cook, P. L. 1967. Polyzoa (Bryozoa) from West Africa, the Pseudostega, the Cribrimorpha, and some Ascophora Imperfecta. Bulletin of the British Museum (Natural History), Zoology, 15:321–351.Google Scholar
Cook, P. L. 1968. Bryozoa (Polyzoa) from the coasts of tropical West Africa. Atlantide Report, number 10:115–262.Google Scholar
Cook, P. L. 1973a. Settlement and early colony development in some Cheilostomata, p. 65–71. In Larwood, G. P. (ed.), Living and Fossil Bryozoa. Academic Press, London.Google Scholar
Cook, P. L. 1973b. Preliminary notes on the ontogeny of the frontal body wall in the Adeonidae and Adeonellidae (Bryozoa, Cheilostomata). Bulletin of the British Museum (Natural History), Zoology, 25:245–263.Google Scholar
Cook, P. L. 1979. Some problems in interpretation of heteromorphy and colony integration in Bryozoa, p. 193–210. In Larwood, G. P. and Rosen, B. R. (eds.), Biology and Systematics of Colonial Organisms. Academic Press, London.Google Scholar
Cook, P. L. 1985. Bryozoa from Ghana, A preliminary survey. Annalen Zoologische Wetenschappen, 238: 315 p.Google Scholar
Cook, P. L., and Chimonides, P. J. 1981. Morphology and systematics of some interior-walled cheilostome Bryozoa. Bulletin of the British Museum (Natural History), Zoology, 41:53–89.Google Scholar
Dick, M. H., and Mawatari, S. F. 2004. Resolving taxonomic problems of north Pacific bryozoans, p. 67–74. In Mawatari, S. F. and Okada, H. (eds.), Neo-Science of Natural History: Integration of Geoscience and Biodiversity Studies. Proceedings of International Symposium on “Dawn of a New Natural History—Integration of Geoscience and Diversity Studies,” Hokkaido University, Sapporo, Japan.Google Scholar
Dick, M. H., and Mawatari, S. F. 2005. Morphological and molecular concordance of Rhynchozoon clades (Bryozoa, Cheilostomata) from Alaska. Invertebrate Biology, 124:344–354.Google Scholar
Dick, M. H., Herrera-Cubilla, A., and Jackson, J. B. C. 2003. Molecular phylogeny and phylogeography of free-living Bryozoa (Cupuladriidae) from both sides of the Isthmus of Panama. Molecular Phylogenetics and Evolution, 26:355–371.CrossRefGoogle Scholar
d'Orbigny, A. 1852. Paléontologie Française. Description des Animaux Invertébrés. Terrain Crétacé. 5, Bryozoaires. Masson, Paris, p. 189–472.Google Scholar
Foote, M. 1996. On the probability of ancestors in the fossil record. Paleobiology, 22:141–151.CrossRefGoogle Scholar
Foote, M., and Raup, D. M. 1996. Fossil preservation and the stratigraphic ranges of taxa. Paleobiology, 22:121–140.CrossRefGoogle ScholarPubMed
Gabb, W. M., and Horn, G. H. 1862. Monograph of the fossil Polyzoa of the Secondary and Tertiary formations of North America. Journal of the Academy of Natural Sciences of Philadelphia, 5(2):111–179.Google Scholar
Gontar, V. I. 2002. New species and new genera of Cheilostomata from the Weddell Sea, Antarctica (Bryozoa). Zoosystematica Rossica, 10: 285–292.Google Scholar
Gordon, D. P. 1993. Bryozoan frontal shields: Studies on umbonulomorphs and impacts on classification. Zoologica Scripta, 22:203–221.Google Scholar
Gould, S. J. 2001. The interrelationship of speciation and punctuated equilibrium, p. 196–217. 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
Gould, S. J. 2002. The Structure of Evolutionary Theory. Belknap Press of Harvard University Press, Cambridge, Massachusetts, 1433 p.Google Scholar
Grischenko, A. V., and Mawatari, S. F. 2002. Kubaninella: A new genus of Adeonidae (Bryozoa: Cheilostomata) from the Western Kamchatka shelf of the Sea of Okhotsk, p. 125–130. In Wyse Jackson, P. N., Buttler, C. J., and Spencer Jones, M. E. (eds.), Bryozoan Studies 2001, Proceedings of the Twelfth International Bryozoology Association Conference. A. A. Balkema, Lisse.Google Scholar
Hageman, S. J., Bayers, 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
Harmer, S. F. 1957. The Polyzoa of the Siboga Expedition, Pt. IV, Cheilostomata Ascophora, II. Siboga-Expeditie, Monographic 28d, p. 641–1147.Google Scholar
Hayward, P. J., and Ryland, J. S. 1979. British Ascophoran Bryozoans: Keys and Notes for the Identification of Species. Academic Press, London, 312 p.Google Scholar
Hayward, P. J., and Thorpe, J. P. 1987. The systematic position of Smittia inclusa Waters, an endemic Antarctic bryozoan. Journal of Natural History, 21:1469–1476.Google Scholar
Herrera-Cubilla, A., Dick, M. H., Sanner, J., and Jackson, J. B. C. 2006. Neogene Cupuladriidae of tropical America, I: Taxonomy of Recent Cupuladria from opposite sides of the Isthmus of Panama. Journal of Paleontology, 80:245–263.Google Scholar
Hincks, T. 1877. On British Polyzoa, Pt. II, Classification. Annals and Magazine of Natural History, (4) 20:520–532.Google Scholar
Hincks, T. 1895. Contributions Towards a General History of the Marine Polyzoa. Privately printed, 213 p.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
Jackson, J. B. C., and Cheetham, A. H. 1994. Phylogeny reconstruction and the tempo of speciation in cheilostome Bryozoa. Paleobiology, 20:407–423.Google Scholar
Jackson, J. B. C., and Cheetham, A. H. 1999. Tempo and mode of speciation in the sea. Trends in Ecology and Evolution, 14:72–77.Google Scholar
Jackson, J. B. C., and McKinney, F. K. 1990. Ecological processes and progressive macroevolution of marine clonal benthos, p. 173–209. In Ross, R. M. and Allmon, W. D. (eds.), Causes of Evolution: A Paleontological Perspective. University of Chicago Press, Chicago.Google Scholar
Johnston, G. 1847. A History of the British Zoophytes. John Van Voorst, London, 488 p.Google Scholar
Jullien, J. 1882. Dragages du “Travailleur.” Bryozoaires, espèces draguées dans l'Ocean Atlantique en 1881. Bulletin de la Societé Zoologique de la France, 7:497–529.Google Scholar
Jullien, J. 1903. Pages 1–188. In Jullien, J. and Calvet, L. (eds.), Bryozoaires Provenant des Campagnes de l'Hirondelle (1886–1888). Résultats des Campagnes Scientifiques Accomplies sur son Yacht par Albert Ier Prince Souverain de Monaco. Fasc. 23.Google Scholar
Kornicker, L. S., Bonnet, F., Cann, R., and Hoskins, C. M. 1959. Alacran Reef, Campeche Bank, Mexico. Institute of Marine Science, 6:1–22.Google Scholar
Labracherie, M. 1975. Description des Bryozoaires cheilostomes d'âge Eocène inférieur du site 246 (croisière 25, Deep Sea Drilling Project). Bulletin de l'Institut de Géologie du Bassin d'Aquitaine, 18:149–202.Google Scholar
Lagaaij, R. 1952. The Pliocene Bryozoa of the Low Countries and Their Bearing on the Marine Stratigraphy of the North Sea Region. Ernst van Aelst, Maastricht, The Netherlands, 233 p.Google Scholar
Lagaaij, R. 1973. Shallow-water Bryozoa from deep-ea sands of the Principe Channel, Gulf of Guinea, p. 139–151. In Larwood, G. P. (ed.), Living and Fossil Bryozoa, Recent Advances in Research. Academic Press, London and New York.Google Scholar
Levinsen, G. M. R. 1909. Morphologic and Systematic Studies on the Cheilostomatous Bryozoa. Nationale Forfatteres Forlag, Copenhagen, 427 p.Google Scholar
Levinton, J. 1988. Genetics, Paleontology, and Macroevolution. Cambridge University Press, Cambridge, England, 637 p.Google Scholar
Lidgard, S. 1996. Zooid skeletal morphogenesis of some Australian and New Zealand Adeonellopsis (Cheilostomatida), p. 167–177. In Gordon, D. P., Smith, A. M., and Grant-Mackie, J. A. (eds.), Bryozoans in Space and Time. NIWA, Wellington, New Zealand.Google Scholar
Lidgard, S., and Buckley, G. A. 1992. Toward a morphological species concept in cheilostomates: Phenotypic variation in Adeonellopsis yarraensis (Waters), p. 101–105. In Hayward, P. J., Ryland, J. S., and Taylor, P. D. (eds.), Biology and Paleobiology of Bryozoans. Olsen and Olsen, Fredensborg, Denmark.Google Scholar
MacGillivray, P. H. 1886. Descriptions on new, or little known, Polyzoa, Pt. IX. Transactions and Proceedings of the Royal Society of Victoria, 22:128–139.Google Scholar
Marcus, E. 1938. Bryozoarios marinhos brasileiros II. Universidad de Sao Paulo, Boletins da Faculdade de Philosophia, Sciências e Letras IV, Zoologia, 2, 196 p.Google Scholar
Marcus, E. 1949. Some Bryozoa from the Brazilian coast. Communicaciones Zoologicas del Museo de Historia Natural de Montevideo, 3(53), 33 p.Google Scholar
Marcus, E. 1955. Notas sôbre Briozoos marinhos brasileiros. Arquivos do Museu Nacional, 42:273–342.Google Scholar
Marshall, C. R. 1990. Confidence intervals on stratigraphic ranges. Paleobiology, 16:1–10.Google Scholar
Maturo, F. J. S. 1968. The distributional pattern of the Bryozoa of the east coast of the United States exclusive of New England, p. 261–284. In Annoscia, E. (ed.), Proceedings of the First International Conference on Bryozoa. Atti della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano, 108.Google Scholar
Milne Edwards, H. 1836a. Observations sur les Polypiers fossiles du genre Eschare. Annales des Sciences Naturelles (Zoologie), Paris, série 2, 6:321–345.Google Scholar
Milne Edwards, H. 1836b. II. Histoire des polypes, p. 1–683. In de Lamarck, J. B. P. A., Histoire Naturelle des Animaux sans Vertèbres (second edition). Maisson, Paris.Google Scholar
Nielsen, C. 1981. On morphology and reproduction of “Hippodiplosia” insculpta and Fenestrulina malusii (Bryozoa, Cheilostomata). Ophelia, 20:91–125.Google Scholar
Norusis, M. J. 1994a. SPSS Base System, 6.1. SPSS, Inc., Chicago, 941 p.Google Scholar
Norusis, M. J. 1994b. SPSS Professional Statistics, 6.1. SPSS, Inc., Chicago, 385 p.Google Scholar
Osburn, R. C. 1914. The Bryozoa of the Tortugas Islands, Florida. Carnegie Institution of Washington Publication, 182:181–222.Google Scholar
Osburn, R. C. 1940. Bryozoa of Porto Rico with a resume of the West Indian bryozoan fauna. Scientific Survey of Porto Rico and the Virgin Islands, 16:321–486.Google Scholar
Osburn, R. C. 1952. Bryozoa of the Pacific Coast of America, Pt. 2, Cheilostomata-Ascophora. Allan Hancock Pacific Expeditions, 14:271–611.Google Scholar
Ostrovsky, A. N., and Taylor, P. D. 2005. Ovicell development in the early calloporid Wilbertopora Cheetham (Bryozoa: Cheilostomata) from the mid-Cretaceous of the USA, p. 223–230. In Moyano, H. I., Cancino, J. M., and Wyse Jackson, P. N. (eds.), Bryozoan Studies, 2004. A. A. Balkema Publishers, Leiden.Google Scholar
Ostrovsky, A. N., Grischenko, A. V., Taylor, P. D., Bock, P., and Mawatari, S. F. 2006. Comparative anatomical study of internal brooding in three anascan bryozoans (Cheilostomata) and its taxonomic and evolutionary implications. Journal of Morphology, 267:739–749.Google Scholar
Page, R. D. M. 1996. Treeview, an application to display phylogenetic trees on personal computers. Computer Applications in the Biosciences, 12:357–358.Google Scholar
Redier, L. 1965. Hydraires et Bryozoaires du Golfe de Guinée. Bulletin du Muséum National d'Histoire Naturelle, série 2, 37:367–394.Google Scholar
Reguant, S. 1990. Metrarabdotos oriense n. sp. (Bryozoa Cheilostomata Ascophora) del Eoceno Medio de Vic (Barcelona): Una forma ramosa singula. Revista Española de Paleontologia, 5:71–76.Google Scholar
Reuss, A. 1847. Die fossilen Polyparien des Wiener Tertiärbeckens. Haidingers naturwissenschaftliche Abhandlungen, Wien, 2, 109 p.Google Scholar
Roger, J., and Buge, E. 1947. Les Bryozoaires du Redonien. Bulletin de la Société Geologique de France, série 5, 16:217–230.Google Scholar
Rosso, A. 2005. Metrarabdotos (Bryozoa, Cheilostomatida) from Plio-Pleistocene of southern Italy, with description of new species. Bollettino della Società Paleontologica Italiana, 44:11–24.Google Scholar
Sandberg, P. A. 1977. Ultrastructure, mineralogy, and development of bryozoan skeletons, p. 144–181. In Woollacott, R. M. and Zimmer, R. L. (eds.), Biology of Bryozoans. Academic Press, New York.Google Scholar
Sandberg, P. A. 1983. Ultrastructure and skeletal development in cheilostomate Bryozoa, p. 238–286. In Robison, R. A. (ed.), Treatise on Invertebrate Paleontology, Pt. G, Bryozoa (revised). Geological Society of America and University of Kansas Press, Lawrence.Google Scholar
Saunders, J. B., Jung, P., and Biju-Duval, B. 1986. Neogene Paleontology in the northern Dominican Republic. 1. Field Surveys, Lithology, Environment, and Age. Bulletins of American Paleontology, number 323, 79 p.Google Scholar
Schopf, T. J. M. 1973. Ergonomics of polymorphism: Its relation to the colony as the unit of natural selection in species of the phylum Ectoprocta, p. 247–294. In Boardman, R. S., Cheetham, A. H., and Oliver, W. A. Jr. (eds.), Animal Colonies: Development and Function Through Time. Dowden, Hutchinson, and Ross, Stroudsburg, Pennsylvania.Google Scholar
Schwaninger, H. R. 1999. Population structure of the widely dispersing marine bryozoan Membranipora membranacea (Cheilostomata): Implications for population history, biogeography, and taxonomy. Marine Biology, 135:411–423.CrossRefGoogle Scholar
Smitt, F. 1873. Floridan Bryozoa, collected by Count L.F. de Pourtales, Pt. II. Kongliga Svenska Vetenskaps-Akademiens Handlingar, 11(4):3–83.Google Scholar
Swofford, D. L. 2000. PAUP*, Phylogenetic Analysis Using Parsimony (*and Other Methods), Version 4. Sinauer Associates, Sunderland, Massachusetts.Google Scholar
Taylor, P. D., and Foster, T. S. 1998. Bryozoans from the Pliocene Bowden shell bed of Jamaica. Contributions to Tertiary and Quaternary Geology, 35(1–4):63–83.Google Scholar
Vigneaux, M. 1949. Révision des Bryozoaires néogènes du Bassin d'Aquitaine. Mémoires de la Société Géologique de France, n. s., 28, 155 p.Google Scholar
Wagner, P. J. 1998. Phylogenetic analyses and the quality of the fossil record, p. 165–187. In Donovan, S. K. and Paul, C. R. C. (eds.), The Adequacy of the Fossil Record. Wiley and Sons, Chicester, England Google Scholar
Waters, A. W. 1904. Bryozoa. Expedition antarctique Beige. Resultats du voyage du S. Y. Belgica en 1897–1898–1899 sous le commandement de A. de Gerlache de Gomery. Rapports Scientifique, Zoologie, 7(7), 113 p.Google Scholar
Winston, J. E. 1982. Marine bryozoans (Ectoprocta) of the Indian River area (Florida). Bulletin of the American Museum of Natural History, 173:99–176.Google Scholar
Winston, J. E. 1984. Shallow-water bryozoans of Carrie Bow Cay, Belize. American Museum Novitates, number 2799, 38 p.Google Scholar
Winston, J. E. 1986. An annotated checklist of coral-associated bryozoans. American Museum Novitates, number 2859, 39 p.Google Scholar
Winston, J. E. 2005. Re-description and revision of Smitt's “Floridan Bryozoa” in the collection of the Museum of Comparative Zoology, Harvard University. Virginia Museum of Natural History Memoir, 7, 152 p.Google Scholar