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Stream channel network analysis applied to colony-wide feeding structures in a Permian bryozoan from Greenland

Published online by Cambridge University Press:  08 April 2016

Marcus M. Key Jr.
Affiliation:
Department of Earth Sciences, Dickinson College, Carlisle, Pennsylvania 17013-2896. E-mail: [email protected]
Patrick N. Wyse Jackson
Affiliation:
Department of Geology, Trinity College, Dublin 2, Ireland
Louis J. Vitiello
Affiliation:
Department of Earth Sciences, Dickinson College, Carlisle, Pennsylvania 17013-2896. E-mail: [email protected]

Abstract

Colony-wide feeding currents are a common feature of many bryozoan colonies. These feeding currents are centered on excurrent macular chimneys that expel previously filtered water away from the colony surface. In some bryozoans these macular chimneys consist of a branching channel network that converges at a point in the center of the chimney. The bifurcating channels of the maculae are analogous to a stream channel network in a closed basin with centripetal drainage. The classical methods of stream channel network analysis from geomorphology are here used to quantitatively analyze the number and length of macular channels in bryozoans. This approach is applied to a giant branch of the trepostome bryozoan Tabulipora from the Early Permian Kim Fjelde Formation in North Greenland. Its large size allowed 18 serial tangential peels to be made through the 8-mm-thick exozone. The peels intersected two stellate maculae as defined by contiguous exilapores. The lengths of 1460 channels radiating from the maculae were measured and their Horton-Strahler stream order and Shreve magnitude scored.

We hypothesize that if fossil bryozoan maculae function as excurrent water chimneys, then they should conform to Horton's laws of stream networks and behave like closed basins with centripetal drainage. Results indicate that the stellate maculae in this bryozoan behaved liked stream channel networks exhibiting landscape maturation and stream capture. They conformed to the Law of Stream Number. They have a Bifurcation Ratio that falls within the range of natural stream channel networks. They showed a pattern opposite that expected by the Law of Stream Lengths in response to behavior characteristic of a centripetal drainage pattern in a closed basin. Thus, the stellate maculae in this bryozoan probably functioned as excurrent water chimneys with the radiating channels serving to efficiently collect the previously filtered water, conducting it to the central chimney for expulsion away from the colony surface.

Type
Articles
Copyright
Copyright © The Paleontological Society 

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References

Literature Cited

Andah, K., Rosso, R., and Taramasso, A. C. 1987. The role of quantitative geomorphology in the hydrological response of river networks. In Rodda, J. C. and Matalas, N. C., eds. Water for the future: hydrology in perspective. International Association of Hydrological Sciences Publication 164:93110.Google Scholar
Anstey, R. L. 1981. Zooid orientation structures and water flow patterns in Paleozoic bryozoan colonies. Lethaia 14:287302.Google Scholar
Anstey, R. L. 1987. Colony patterning and functional morphology of water flow in Paleozoic stenolaemate bryozoans. Pp. 18 in Ross, 1987.Google Scholar
Anstey, R. L., and Delmet, D. A. 1972. Genetic meaning of zooecial chamber shapes in fossil bryozoans. Science 177:10001002.CrossRefGoogle ScholarPubMed
Anstey, R. L., and Pachut, J. F. 1976. Functional morphology of animal colonies by comparison to sand dune paradigms. Geological Society of America Abstracts with Programs 8:124125.Google Scholar
Anstey, R. L., and Pachut, J. F. 1980. Fourier packing ordinate: a univariate size-independent measurement of the polygonal packing variation in Paleozoic bryozoans. Mathematical Geology 12:139156.Google Scholar
Anstey, R. L., Pachut, J. F., and Prezbindowski, D. R. 1976. Morphogenetic gradients in Paleozoic bryozoan colonies. Paleobiology 2:131146.Google Scholar
Astrova, G. G. 1973. Polymorphism and its development in the trepostomatous Bryozoa. Pp. 110 in Larwood, G. P., ed. Living and fossil Bryozoa. Academic Press, London.Google Scholar
Banta, W. C., McKinney, F. K., and Zimmer, R. L. 1974. Bryozoan monticules: excurrent water outlets? Science 185:783784.Google Scholar
Barker, S. B., Cumming, G., and Horsfield, K. 1973. Quantitative morphometry of the branching structure of trees. Journal of Theoretical Biology 40:3343.Google Scholar
Baugh, N. F., and Brown, R. H. 2008. Channel length, stream order and channel network integration on Titan. Lunar and Planetary Science 39:1943.Google Scholar
Bejan, A. 2000. Shape and structure from engineering to nature. Cambridge University Press, Cambridge, United Kingdom. Google Scholar
Bejan, A., and Lorente, S. 2008. Design with constructal theory. Wiley, Hoboken, N.J. Google Scholar
Bishop, J. W., and Bahr, L. M. 1973. Effects of colony size on feeding by Lophopodella carteri (Hyatt). Pp. 433437 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, PA. Google Scholar
Boardman, R. S. 1983. General features of the class Stenolaemata. Pp. 49137 in Boardman, et al., 1983b.Google Scholar
Boardman, R. S., and Buttler, C. J. 2005. Zooids and extrazooidal skeleton in the Order Trepostomata (Bryozoa). Journal of Paleontology 79:10881104.Google Scholar
Boardman, R. S., and Cheetham, A. H. 1983. Glossary of morphological terms. Pp. 304320 in Boardman, et al., 1983b.Google Scholar
Boardman, R. S., Cheetham, A. H., and Cook, P. L. 1983a. Introduction to the Bryozoa. Pp. 348 in Boardman, et al., 1983b.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. 1983b. Bryozoa 1 revised, Part G of Robison, R. A., ed. Treatise on invertebrate paleontology. Geological Society of America, Boulder, Colo., and University of Kansas, Lawrence.Google Scholar
Boyajian, G. E., and LaBarbera, M. C. 1987. Biomechanical analysis of passive flow of stromatoporoids-morphologic, paleoecologic, and systematic implications. Lethaia 20:223229.Google Scholar
Buss, L. W. 1979. Habitat selection, directional growth and spatial refuges: why colonial animals have more hiding places. Pp. 459497 in Larwood, G. and Rosen, B. R., eds. Biology and systematics of colonial organisms. Academic Press, New York.Google Scholar
Buss, L. W. 1980. Bryozoan overgrowth interactions — the interdependence of competition for space and food. Nature 281:475477.Google Scholar
Buss, L. W. 1981. Mechanisms of competition between Onychocella alula (Hastings) and Antropora tincta (Hastings) on an eastern Pacific rocky shoreline. Pp. 3949 in Larwood, G. P. and Nielsen, C., eds. Recent and fossil Bryozoa. Olsen and Olsen, Fredensborg, Denmark. Google Scholar
Buss, L. W. 2001. Growth by intussusception in hydractiniid hydroids. Pp. 326 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
Chorley, R. J. 1969. Water, earth, and man, a synthesis of hydrology, geomorphology, and socio-economic geography. Methuen, London.Google Scholar
Cook, P. L. 1977. Colony-wide water currents in living bryozoans. Cahiers de Biologie Marine 18:3147.Google Scholar
Cook, P. L., and Chimonides, P. J. 1980. Further observations on water current patterns in living Bryozoa. Cahiers de Biologie Marine 21:393402.Google Scholar
Delmet, D. A., and Anstey, R. L. 1974. Fourier analysis of morphological plasticity within an Ordovician bryozoan colony. Journal of Paleontology 48:217226.Google Scholar
Dick, M. H. 1987. A proposed mechanism for chimney formation in encrusting bryozoan colonies. Pp. 7380 in Ross, 1987.Google Scholar
Dodds, P. S., and Rothman, D. H. 2000. Scaling, universality, and geomorphology. Annual Review of Earth and Planetary Sciences 28:571610.Google Scholar
Eckman, J. E., and Okamura, B. 1998. A model of particle capture by bryozoans in turbulent flow: significance of colony form. American Naturalist 152:861880.Google Scholar
Folk, R. L. 1971. Genesis of longitudinal and oghurd dunes elucidated by rolling upon grease. Geological Society of America Bulletin 82:34613468.Google Scholar
Grünbaum, D. 1995. A model of feeding currents in encrusting bryozoans shows interference between zooids within a colony. Journal of Theoretical Biology 174:409425.CrossRefGoogle Scholar
Grünbaum, D. 1997. Hydromechanical mechanisms of colony organization and cost of defense in an encrusting bryozoan, Membranipora membranacea . Limnology and Oceanography 42:741752.Google Scholar
Håkansson, E. 1979. Carboniferous to Tertiary development of the Wandel Sea Basin, Peary Land, eastern North Greenland. Rapport Gr⊘nlands Geologiske Unders⊘gelse 88:7383.Google Scholar
Håkansson, E., Heinberg, C., and Stemmerik, L. 1981. The Wandel Sea Basin from Holm Land to Lockwood Ø, eastern North Greenland. Rapport Gr⊘nlands Geologiske Unders⊘gelse 106:4763.Google Scholar
Håkansson, E., and Madsen, L. 1991. Symbiosis—a plausible explanation of gigantism in Permian trepostome bryozoans. Pp. 151159 in Bigey, F. P. and d'Hondt, J.-L., eds. Bryozoa: living and fossil. Société des Sciences Naturelles de l'Ouest de la France, Mémoire hors série, Nantes, France. Google Scholar
Horsfield, K., and Cumming, G. 1976. Morphology of the bronchial tree in the dog. Respiration Physiology 26:173182.CrossRefGoogle ScholarPubMed
Horsfield, K., Relea, F. G., and Cumming, G. 1976. Diameter, length and branching ratios in the bronchial tree. Respiration Physiology 26:351356.Google Scholar
Horton, R. E. 1945. Erosional development of streams and their drainage basins; hydrophysical approach to quantitative morphology. Geological Society America Bulletin 56:275370.CrossRefGoogle Scholar
J⊘rgensen, C. B. 1966. Biology of suspension feeding. Pergamon, Oxford.Google Scholar
Key, M. M. Jr., Thrane, L., and Collins, J. A. 2001. Space-filling problems in ramose trepostome bryozoans as exemplified in a giant colony from the Permian of Greenland. Lethaia 34:125135.Google Scholar
Key, M. M. Jr., Thrane, L., and Collins, J. A. 2002. Functional morphology of maculae in a giant ramose bryozoan from the Permian of Greenland. Pp. 163170 in Jackson, P. N. Wyse, Buttler, C. J., and Jones, M. E. Spencer, eds. Bryozoan studies 2001. Balkema, Lisse, The Netherlands. Google Scholar
Key, M. M. Jr., Jackson, P. N. Wyse, Håkansson, E., Patterson, W. P., and Moore, M. D. 2005. Gigantism in Permian trepostomes from Greenland: testing the algal symbiosis hypothesis using δ13C and δ18O values. Pp. 141151 in Moyano, H. I. G., Cancino, J. M., and Jackson, P. N. Wyse, eds. Bryozoan studies 2004. Balkema, Leiden, The Netherlands. Google Scholar
Kirchner, J. W. 1993. Statistical inevitability of Horton's laws and the apparent randomness of stream channel networks. Geology 21:591594.Google Scholar
LaBarbera, M. C., and Boyajian, G. E. 1991. The function of astrorhizae in stromatoporoids: quantitative tests. Paleobiology 17:121132.Google Scholar
Langille, B. L. 1995. Blood flow-induced remodeling of the artery wall. Pp. 277299 in Bevan, J. A., Kaley, G. and Rubanyi, G. M., eds. Flow-dependent regulation of vascular function. Oxford University Press, New York.Google Scholar
Larsen, P. S., and Riisgård, H. U. 2001. Chimney spacing in encrusting bryozoan colonies (Membranipora membranacea): video observations and hydrodynamic modeling. Ophelia 54:167176.Google Scholar
Leopold, L. B. 1971. Trees and streams: the efficiency of branching patterns. Journal of Theoretical Biology 31:339354.Google Scholar
Leopold, L. B., Wolman, M. G., and Miller, J. P. 1964. Fluvial processes in geomorphology. W. H. Freeman, San Francisco.Google Scholar
Lidgard, S. 1981. Water flow, feeding, and colony form in an encrusting cheilostome. Pp. 135142 in Larwood, G. P. and Nielsen, C., eds. Recent and fossil Bryozoa. Olsen and Olsen, Fredensborg, Denmark. Google Scholar
Lin, H., Kang, S., Gburek, W. J., Folmar, G. J., Sharpley, A. N. 2003. Stream order and fractal as scaling factors for watershed water quality. Eos 84:F717.Google Scholar
Madsen, L. 1987. Growth and polypide morphology in some ramose trepostome bryozoans from the Permo-Carboniferous of the Arctic. Pp. 169176 in Ross, 1987.Google Scholar
Madsen, L. 1994. Bryozoans from the Upper Palaeozoic sequence in the Wandel Sea Basin, North Greenland. Wandel Sea Basin: Basin Analysis Scientific Report No. 6. University of Copenhagen, Denmark.Google Scholar
Madsen, L., and Håkansson, E. 1989. Upper Paleozoic bryozoans from the Wandel Sea Basin, North Greenland. Rapport Gr⊘nlands Geologiske Unders⊘gelse 144:4352.CrossRefGoogle Scholar
McKinney, F. K. 1986a. Historical record of erect bryozoan growth forms. Proceedings of the Royal Society of London B 228:133148.Google Scholar
McKinney, F. K. 1986b. Evolution of erect marine bryozoan faunas: repeated success of unilaminate species. American Naturalist 128:795809.Google Scholar
McKinney, F. K. 1988. Elevation of lophophores by exposed introverts in Bryozoa: a gymnolaemate character recorded in some stenolaemate species. Bulletin of Marine Science 43:317322.Google Scholar
McKinney, F. K. 1989. Two patterns of colonial water flow in an erect bilaminate bryozoan, the cheilostome Schizotheca serratimargo (Hincks, 1886). Cahiers de Biologie Marine 30:3548.Google Scholar
McKinney, F. K. 1990. Feeding and associated colonial morphology in marine bryozoans. Critical Reviews in Aquatic Sciences 2:255280.Google Scholar
McKinney, F. K. 1992. Competitive interactions between related clades: evolutionary implications of overgrowth interactions between encrusting cyclostome and cheilostomes bryozoans. Marine Biology 114:645652.Google Scholar
McKinney, F. K., and Jackson, J. B. C. 1989. Bryozoan evolution. Unwin-Hyman, London.Google Scholar
McShea, D. W., and Venit, E. P. 2002. Testing for bias in the evolution of coloniality: a demonstration in cyclostome bryozoans. Paleobiology 28:308327.Google Scholar
Morisawa, M. E. 1957. Accuracy of determination of stream lengths from topographic maps. Transactions of the American Geophysical Union 38:8688.Google Scholar
Morisawa, M. E. 1968. Streams, their dynamics and morphology. McGraw-Hill, New York.Google Scholar
Nakagaki, T., Yamada, H., and Tóth, Á. 2001. Path finding by tube morphogenesis in an amoeboid organism. Biophysical Chemistry 92:4752.Google Scholar
Nielson, J., and Kocurek, G. 1987. Surface processes, deposits, and development of star dunes: Dumont dune field, California. Geological Society of America Bulletin 99:177186.Google Scholar
Okamura, B., and Partridge, J. C. 1999. Suspension feeding adaptations to extreme flow environments in a marine bryozoan. Biological Bulletin 196:205215.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.Google 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.Google Scholar
Patzkowsky, M. E. 1987. Inferred water flow patterns in the fossil Fistulipora M'Coy (Cystoporata, Bryozoa). Pp. 213219 in Ross, 1987.Google Scholar
Paul, C. R. C. 1975. A reappraisal of the paradigm method of functional analysis in fossils. Lethaia 7:1521.Google Scholar
Pelletier, J. D. 1999. Self-organization and scaling relationships of evolving river networks. Journal of Geophysical Research 104:73597375.Google Scholar
Pilgrim, D. H. 1983. Some problems in transferring hydrological relationships between small and large drainage basins and between regions. Journal of Hydrology 65:4972.Google Scholar
Pilgrim, D. H., Cordery, I., and Baron, B. C. 1982. Effects of catchment size on runoff relationships. Journal of Hydrology 58:205221.CrossRefGoogle Scholar
Podell, M. E., and Anstey, R. L. 1979. The interrelationship of early colony development, monticules, and branches in Palaeozoic bryozoans. Paleontology 22:965982.Google Scholar
Pratt, M. C. 2004. Effect of zooid spacing on bryozoan feeding success: is competition or facilitation more important? Biological Bulletin 207:1727.Google Scholar
Rasmussen, J. A., and Håkansson, E. 1996. First Permo-Carboniferous conodonts from North Greenland. Geological Magazine 133:553564.Google Scholar
Ross, J. R. P. 1987. Bryozoa: present and past. Western Washington University, Bellingham.Google Scholar
Ross, J. P., and Ross, C. A. 1962. Faunas and correlation of the late Paleozoic rocks of northeast Greenland, part IV, Bryozoa. Meddelelser Om Gr⊘nland 167:165.Google Scholar
Rudwick, M. J. S. 1964. The inference of function from structure in fossils. British Journal for the Philosophy of Science 15:2740.CrossRefGoogle Scholar
Scheidegger, A. E. 1966. Statistical description of river networks. Water Resources Research 2:785790.CrossRefGoogle Scholar
Shreve, R. L. 1966. Statistical law of stream numbers. Journal of Geology 74:1737.CrossRefGoogle Scholar
Shreve, R. L. 1967. Infinite typologically random channel networks. Journal of Geology 75:178186.Google Scholar
Shunatova, N. N., and Ostrovsky, A. N. 2002. Group autozooidal behavior and chimneys in marine bryozoans. Marine Biology 140:503518.Google Scholar
Slaymaker, O. 2006. Towards the identification of scaling relations in drainage basin sediment budgets. Geomorphology 80:819.Google Scholar
Stemmerik, L. 1997. Permian (Artinskian-Kazanian) cool-water carbonates in North Greenland, Svalbard and the western Barents Sea. In James, N. P. and Clarke, J. A. D., eds. Cool-water carbonates. Society for Sedimentary Geology Special Publication 55:349364.Google Scholar
Stemmerik, L., and Håkansson, E. 1989. Stratigraphy and depositional history of the Upper Palaeozoic and Triassic sediments in the Wandel Sea Basin, central and eastern North Greenland. Rapport Gr⊘nlands Geologiske Unders⊘gelse 143:2145.Google Scholar
Stemmerik, L., Håkansson, E., Madsen, L., Nilsson, I. Piasecki, S., Pinard, S., and Rasmussen, J. A. 1996. Stratigraphy and depositional evolution of the Upper Palaeozoic sedimentary succession in eastern Peary Land, North Greenland. Bulletin Gr⊘nlands Geologiske Unders⊘gelse 171:4571.Google Scholar
Stepinski, T. F., and Collier, M. L. 2003. Drainage densities of computationally extracted Martian drainage basins. International Conference on Mars 6:3100.Google Scholar
Strahler, A. N. 1952. Hypsometric (area-altitude) analysis of erosional topography. Geological Society of America Bulletin 63:11171142.Google Scholar
Strahler, A. N. 1957. Quantitative analysis of watershed geomorphology. American Geophysical Union Transactions 38:913920.Google Scholar
Taylor, P. D. 1975. Monticules in a Jurassic cyclostomatous bryozoan. Geological Magazine 112:601606.Google Scholar
Taylor, P. D. 1979. The inference of extrazooidal feeding currents in fossil bryozoan colonies. Lethaia 12:4756.Google Scholar
Taylor, P. D. 1999. Bryozoans. Pp. 623645 in Savazzi, E., ed. Functional morphology of the invertebrate skeleton. Wiley, New York.Google Scholar
Taylor, P. D., and Voigt, E. 1999. An unusually large cyclostome bryozoan (Pennipora anomalopora) from the Upper Cretaceous of Maastricht. Bulletin de l'Institut Royal des Sciences Naturelles de Belgique, Sciences de la Terre 69:165171.Google Scholar
Thorpe, J. P., and Ryland, J. S. 1987. Some theoretical limitations on the arrangement of zooids in encrusting Bryozoa. Pp. 276283 in Ross, 1987.Google Scholar
Thurlbeck, A., and Horsfield, K. 1980. Branching angles in the bronchial tree related to order of branching. Bulletin of Mathematical Biology 41:173181.Google Scholar
Troutman, B. M., and Karlinger, M. R. 1994. Comment on “Statistical inevitability of Horton's laws and the apparent randomness of stream channel networks.” Geology 22:573574.Google Scholar
Turner, D. 2000. The functions of fossils: inference and explanation in functional morphology. Studies in History and Philosophy of Science C 31:193212.Google Scholar
Ulrich, E. O. 1890. Paleozoic Bryozoa. Pp. 283688 in Lindahl, J., ed. Geology and paleontology, Vol. 8. Geological Survey of Illinois, Springfield.Google Scholar
Utgaard, J. 1983. Paleobiology and taxonomy of the order Cystoporata. Pp. 327357 in Boardman, et al., 1983b.Google Scholar
Vogel, S. 1981. Life in moving fluids. Willard Grant, Boston.Google Scholar
Von Dassow, M. 2005a. Flow and conduit formation in the external fluid-transport system of a suspension feeder. Journal of Experimental Biology 208:29312938.Google Scholar
Von Dassow, M. 2005b. Effects of ambient flow and injury on the morphology of a fluid transport system in a bryozoan. Biological Bulletin 208:4759.Google Scholar
Von Dassow, M. 2006. Function-dependent development in a colonial animal. Biological Bulletin 211:7682.Google Scholar
Winston, J. E. 1978. Polypide morphology and feeding behavior in marine ectoprocts. Bulletin of Marine Science 28:131.Google Scholar
Winston, J. E. 1979. Current-related morphology and behavior in some Pacific Coast bryozoans. Pp. 247268 in Larwood, G. P. and Abbott, M. B., eds. Advances in bryozoology. Academic Press, London.Google Scholar
Winston, J. E. 1981. Feeding behavior of modern bryozoans. In Dutro, J. T. and Boardman, R. S., eds. Lophophorates: notes for a short course. Studies in Geology 5:121. University of Tennessee, Knoxville.Google Scholar