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The shapes of Paleozoic and modern reef-builders: a critical review

Published online by Cambridge University Press:  08 February 2016

Colin W. Stearn
Colin W. Stearn*
Affiliation:
Department of Geological Sciences, McGill University, Montreal, Quebec, H3A 2A7, Canada
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Abstract

Paleoecologists studying Paleozoic reef-builders have interpreted their growth forms as responses to conditions of depth and turbulence in reef complexes. Comparison of the shapes of Paleozoic stromatoporoids and corals with the growth forms of modern scleractinians has been used to reconstruct Paleozoic conditions. A review of shape zonation on modern reefs indicates that no general pattern is applicable to all reefs and variations in shape are the result of the interaction of many environmental factors with the genetically dictated growth pattern of the coral. In most zones of a reef a wide range of shapes co-exist. The growth forms of corals on modern reefs are not a simple vegetative response to the many environmental parameters that have been shown to influence form, but are constrained by phylogenetic and developmental influences as well as functional ones.

Interpretations of the environments of western Canadian and other mid-Paleozoic reefs have been based on the growth forms of stromatoporoids. The environmental significance of the shapes has been deduced from comparison with the shapes of modern scleractinians, functional morphology, nature of the enclosing sediment, position of growth, position within the reef, and diversity gradients. The validity of these criteria is open to question and considerable doubt remains concerning the significance of the growth forms. The shapes of reef animals are not specific guides to environments of modern reefs and should not be expected to be guides for ancient ones.

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Articles
Information
Paleobiology , Volume 8 , Issue 3 , Summer 1982 , pp. 228 - 241
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Copyright © The Paleontological Society 

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References

Literature Cited

Abbott, B. M. 1973. Terminology of stromatoporoid shapes. J. Paleontol. 47:805–806.Google Scholar
Abbott, B. M. 1976. Origin and evolution of bioherms in Wenlock limestone (Silurian) of Shropshire, England. Bull. Am. Assoc. Petrol. Geol. 60:2117–2127.Google Scholar
Adams, R. D. 1968. Leeward Island reefs of St. Vincent. J. Geol. 76:587–595.CrossRefGoogle Scholar
Alberstadt, L. P., Walker, K. R., and Zurawski, R. P. 1974. Patch reefs in the Carters Limestone (Middle Ordovician) in Tennessee and vertical zonation in Ordovician reefs. Bull. Geol. Soc. Am. 85:1171–1182.2.0.CO;2>CrossRefGoogle Scholar
Andrichuk, J. M. 1958. Stratigraphy and facies analysis of Upper Devonian reefs in Leduc, Stettler and Redwater areas, Alberta. Am. Assoc. Petrol. Geol. Bull. 42:1–93.Google Scholar
Antonius, A. 1980. Occurrence and distribution of stony corals, in the Gulf of Cariaco, Venezuela. Int. Rev. Gesamten Hydrobiol. 65:321–338.CrossRefGoogle Scholar
Bak, R. P. M. 1977. Coral reefs and their zonation in Netherlands Antilles. In: Frost, S. H., Weiss, M. P. and Saunders, J. B., eds. Reefs and Related Carbonates—Ecology and Sedimentology. Am. Assoc. Petrol. Geol. Studies Geol. 4:3–16.Google Scholar
Barnes, D. J. 1973. Growth in colonial scleractinians. Bull. Mar. Sci. 23:280–298.Google Scholar
Barnes, J., Bellamy, D. J., Jones, D. J., Whitton, B. A., Drew, E. A., Kenyon, L., Lythgoe, J. N., and Rosen, B. R. 1971. Morphology of the reef front of Aldabra. In: Stoddard, D. R. and Yonge, C. M., eds. Regional Variation in Indian Ocean Coral Reefs. Zool. Soc. London Symp. 28:87–114.Google Scholar
Bottjer, D. S. 1980. Branching morphology of the reef coral Acropora cervicornis in different hydraulic regimes. J. Paleontol. 54:1102–1107.Google Scholar
Brakel, W. H. 1976. The ecology of coral shapes, microhabitat variation in the colony form and corallite structure of Porites on a Jamaican reef. Ph.D. Thesis Yale Univ.Google Scholar
Brakel, W. H. 1977. Corallite variation in Porites and the species problems in corals. 3rd Int. Coral Reef Symp. Proc. 1:457–462.Google Scholar
Broadhurst, F. M. 1966. Growth forms of stromatoporoids in the Silurian of southern Norway. Nor. Geol. Tidssk. Oslo. 46:401–404.Google Scholar
Burchette, T. P. 1981. European Devonian reefs: a review of current concepts and models. In: Toomey, D. F., ed. European Reef Models. Soc. Econ. Paleontol. Mineral. Spec. Publ. 30:85–142.Google Scholar
Chamberlain, J. A. 1978. Mechanical properties of coral skeleton: compressive strength and its adaptive significance. Paleobiology. 4:419–435.CrossRefGoogle Scholar
Chappell, J. 1980. Coral morphology, diversity and reef growth. Nature. 286:249–252.CrossRefGoogle Scholar
Copper, P. and Grawbarger, D. J. 1978. Paleoecological succession leading to a late Ordovician biostrome on Manitoulin Island, Ontario. Can. J. Earth Sci. 15:1987–2005.CrossRefGoogle Scholar
Crossland, C. (Totton, A. K., ed.). 1954. Madreporaria, Hydrocorallinacea, Heliopora. Great Barrier Reef Exped. 1928–1929 Sci. Reps. 6:1–257.Google Scholar
Davies, P. S., Stoddart, D. R., and Sigee, D. C. 1971. Reef forms of Addu Atoll, Maldive Islands. In: Stoddart, D. R. and Yonge, M., eds. Regional Variation in Indian Ocean Coral Reefs. Zool. Soc. London Symp. 28:217–259.Google Scholar
Duchassaing de Frombressin, P. and Michelotti, J. 1864. Supplement au Mémoire sur les coralliares des Antilles. Mém. R. Acad. Sci. Torino. 2(23):97–206.Google Scholar
Dustan, P. 1975a. Gene ecological differentiation in the reef-building coral Montastrea annularis. Ph.D. Thesis, Yale Univ.Google Scholar
Dustan, P. 1975b. Growth and form in the reef-building coral Montastrea annularis. Mar. Biol. 33:101–107.CrossRefGoogle Scholar
Embry, A. F. and Klovan, J. E. 1972. Absolute water depth limits of Late Devonian paleoecologic zones. Geol. Rundschau. 61:672–686.CrossRefGoogle Scholar
Fischbuch, N. R. 1962. Stromatoporoid zones of the Kaybob Reef, Alberta. J. Alberta Soc. Petrol. Geol. 8:62–72.Google Scholar
Fischbuch, N. R. 1968. Stratigraphy Devonian Swan Hills reef complexes of central Alberta. Bull. Can. Petrol. Geol. 16:446–587.Google Scholar
Flügel, H. W. 1976. Ein Spongienmodell für Favositidae. Lethaia. 9:405–419.CrossRefGoogle Scholar
Foster, A. B. 1976. Intraspecific morphologic variation in scleractinian corals: a key to life habitat interpretation. Geol. Soc. Am. Abstr. with Programs. 6:872–873.Google Scholar
Foster, A. B. 1977. Patterns of small scale variation in skeletal morphology within scleractinian corals Montastrea annularis and Siderastrea siderea. 3rd Int. Coral Reef Symp. Miami Proc. 2:409–415.Google Scholar
Foster, A. B. 1979a. Environmental variation in a fossil scleractinian coral. Lethaia. 12:245–264.CrossRefGoogle Scholar
Foster, A. B. 1979b. Phenotypic plasticity in the reef corals Montastrea annularis (Ellis and Solander) and Siderastrea siderea (Ellis and Solander). J. Exp. Mar. Biol. Ecol. 39:25–54.CrossRefGoogle Scholar
Foster, A. B. 1980. Environmental variation in skeletal morphology he Caribbean reef corals Montastrea annularis and Siderastrea siderea. Bull. Mar. Sci. 30:678–709.Google Scholar
Frydl, P. 1979. The effect of parrot fish (Scaridae) on coral in Barbados, W. I. Int. Rev. Gesamt. Hydrobiol. 64:737–748.CrossRefGoogle Scholar
Geister, J. 1977. The influence of wave exposure on the ecological zonation of Caribbean coral reefs. 3rd Int. Coral Reef Symp. Miami, Proc. 1:23–30.Google Scholar
Gladfelter, E. H. and Monahan, R. K. 1977. Primary production and carbonate deposition rates in Acropora palmata from different positions on the reef. 3rd Int. Coral Reef Symp. Miami Proc. 2:389–394.Google Scholar
Goreau, T. F. 1959a. The physiology of skeleton formation in corals. I. A method for measuring the rate of calcium deposition by corals under different conditions. Biol. Bull. 116:59–75.CrossRefGoogle Scholar
Goreau, T. F. 1959b. The ecology of Jamaican coral reefs. I. Species composition and zonation. Ecology. 40:67–90.CrossRefGoogle Scholar
Goreau, T. F. 1963. Calcium carbonate deposition by coralline algae and hermatypic corals in relation to their role as reef builders. Ann. N.Y. Acad. Sci. 109:127–167.CrossRefGoogle Scholar
Goreau, T. F. and Goreau, N. I. 1973. Ecology of Jamaican coral reefs: II Geomorphology, zonation, and sedimentary phases. Bull. Mar. Sci. 23:399–464.Google Scholar
Goreau, T. F. and Hartman, W. D. 1963. Boring sponges as controlling factors in the formation and maintenance of coral reefs. In: Sognnaes, F., ed. Mechanisms of Hard Tissue Destruction. Am. Assoc. Adv. Sci. Publ. 75:25–54.Google Scholar
Goreau, T. F. and Wells, J. W. 1967. The shallow water scleractinia of Jamaica: revised list of species and their vertical distribution range. Bull. Mar. Sci. 17:442–453.Google Scholar
Gould, S. J. 1980. The promise of paleobiology as a nomothetic, evolutionary discipline. Paleobiology 6:96–118.CrossRefGoogle Scholar
Gould, S. J. and Lewontin, R. C. 1979. The spandrels of San Marco and the Panglossian paradigm: a critique of the adaptationist programme. Proc. R. Soc. London. B205:147–164.Google Scholar
Graus, R. R. 1977. Investigation of coral growth adaptations using computer modeling. 3rd Int. Coral Reef Symp. Miami Proc. 2:463–469.Google Scholar
Graus, R. R., Chamberlain, J. A., and Boker, A. M. 1977. Structural modification of corals in relation to waves and currents. In: Frost, S. H., Weiss, M. P. and Saunders, J. B., eds. Reefs and Related Carbonates—Ecology and Sedimentology. Am. Assoc. Petrol. Geol. Studies Geol. 4:135–153.Google Scholar
Graus, R. R. and Macintyre, I. G. 1976. Light control of growth form in colonial reef corals: computer simulation. Science. 193:895–897.CrossRefGoogle ScholarPubMed
Graus, R. R. and Macintyre, I. G. 1982. Variation in growth form of the reef coral Montastrea annularis (Ellis and Solander): a quantitative evaluation of growth response to light distribution using computer simulation. In: Rutzler, K. and Macintyre, I. G., eds. The Atlantic Barrier Reef Ecosystem at Carrie Bow Cay, Belize, 1. Smithsonian Contrib. Mar. Sci. 12:441–464.Google Scholar
Harper, J. D. 1970. Trends of faunal morphologic variation and their environmental significance: key to paleoecologic analysis. (Abstr.). Am. Assoc. Petrol. Geol. Bull. 54:850.Google Scholar
Hartman, W. D. and Goreau, T. F. 1970. Jamaican coralline sponges: their morphology, ecology and fossil relatives. In: Fry, W. G., ed. Biology of the Porifera. Zool. Soc. London Symp. 25:205–243.Google Scholar
Hubbard, J. A. E. B. and Pocock, Y. P. 1972. Sediment rejection by recent scleractinian corals—a key to paleoenvironmental reconstruction. Geol. Rundschau. 61:598–626.CrossRefGoogle Scholar
Jackson, J. B. C. 1979. Morphological strategies of sessile animals. In: Larwood, G. and Rosen, B., eds. Biology and Systematics of Colonial Organisms. Syst. Assoc. Spec. Publ. 11:499–555.Google Scholar
James, N. P. and Ginsburg, R. N. 1979. The deep seaward margin of the Belize barrier and atoll reefs. Int. Assoc. Sedimentologists Spec. Publ. 3:1–191.Google Scholar
Jamieson, E. R. 1971. Paleoecology of reefs in western Canada. North Am. Paleontol. Convention, Chicago, Proc. 2:1300–1340.Google Scholar
Jenik, A. J. and Lerbekmo, J. F. 1968. Facies and geometry of Swan Hills reef member of Beaverhill Lake Formation (Upper Devonian), Goose River Field, Alberta, Canada. Bull. Am. Assoc. Petrol. Geol. 52:21–56.Google Scholar
Kapp, U. S. 1974. Mode of growth of middle Chazyan (Ordovician) stromatoporoids, Vermont. J. Paleontol. 48:1235–1240.Google Scholar
Kapp, U. S. 1975. Paleoecology of Middle Ordovician stromatoporoid mounds in Vermont. Lethaia. 8:195–207.CrossRefGoogle Scholar
Kawaguti, S. 1937. On the physiology of reef corals. 2. The effects of light on the color and form of reef corals. Palao Tropical Biol. Stn. Stud. 1:199–208.Google Scholar
Kershaw, S. 1981. Stromatoporoid growth form and taxonomy in a Silurian bioherm, Gotland. J. Paleontol. 55:1284–1295.Google Scholar
Kershaw, S. and Riding, R. 1980. Stromatoporoid morphotypes of the Middle Devonian Torbay reef complex at Long Quarry Point, Devon. Proc. Ussher Soc. 5:13–23.Google Scholar
Klovan, J. E. 1964. Facies analysis of the Redwater reef complex, Alberta, Canada. Bull. Can. Petrol. Geol. 12:1–100.Google Scholar
Kobluk, D. 1975. Stromatoporoid paleoecology of the southeast margin of the Miette carbonate complex, Jasper Park, Alberta. Bull. Can. Petrol. Geol. 26:218–236.Google Scholar
Kobluk, D. 1978. Reef stromatoporoid morphologies as dynamic populations: Application of field data to a model and the reconstruction of an Upper Devonian reef. Bull. Can. Petrol. Geol. 26:218–236.Google Scholar
Krebs, W. 1974. Devonian carbonate complexes of central Europe. In: Laporte, L. F., ed. Reefs in Time and Space. Soc. Econ. Paleontol. Mineral. Spec. Publ. 18:155–208.CrossRefGoogle Scholar
Lang, J. C. 1973. Interspecific aggression by scleractinian corals. II. Why the race is not only to the swift. Bull. Mar. Sci. 23:260–279.Google Scholar
Langton, J. C. and Chin, G. E. 1968. Rainbow member facies and related reservoir properties, Rainbow Lake, Alberta. Bull. Can. Petrol. Geol. 16:104–143.Google Scholar
Leavitt, E. M. 1968. Petrology, paleontology, Carson Creek north reef complex, Alberta. Bull. Can. Petrol. Geol. 16:298–413.Google Scholar
Lecompte, M. 1958. Les récifs paleozoique en Belge. Geol. Rundschau. 47:384–401.CrossRefGoogle Scholar
Lecompte, M. 1959. Certain data on the genesis and ecologic character of Frasnian reefs of the Ardennes. Int. Geol. Rev. 1(7):1–23.CrossRefGoogle Scholar
Lewis, J. B. 1960. The coral reefs and coral communities of Barbados. W.I. Can. J. Zool. 38:1133–1145.CrossRefGoogle Scholar
Logan, B. W. 1969. Carbonate sediments and reefs, Yucatan shelf, Mexico. Am. Assoc. Petrol. Geol. Mem. 11:144–152.Google Scholar
Loya, Y. and Slobodkin, L. B. 1971. The coral reefs of Eilat (Gulf of Eilat, Red Sea). In: Stoddard, D. R. and Yonge, C. M., eds. Regional Variations in Indian Ocean Coral Reefs. Zool. Soc. London Symp. 28:87–114.Google Scholar
Manten, A. A. 1971. Silurian Reefs of Gotland. 539 pp. Elsevier Publishing Co.; Amsterdam.Google Scholar
Manton, S. M. 1935. Ecological survey of coral reefs. In: Scientific Reports of the Great Barrier Reef Exped. 1928–29. 3:273–312.Google Scholar
Mayall, M. J. 1979. Facies and sedimentology of part of the Middle Devonian limestones of Brixham, South Devon, England. Proc. Geol. Assoc. 90:171–179.CrossRefGoogle Scholar
Mori, K. 1970. Stromatoporoids from the Silurian of Gotland, Part 2. Stockholm Contrib. Geol. 22:1–152.Google Scholar
Mountjoy, E. W. and Jull, R. K. 1978. Fore-reef carbonate mud bioherms and associated reef margin, Upper Devonian, Ancient Wall reef complex, Alberta. Can. J. Earth Sci. 15:1304–1325.CrossRefGoogle Scholar
Murray, J. W. 1966. An oil producing, reef fringed carbonate bank in the Upper Devonian Swan Hills Member, Judy Creek, Alberta. Bull. Can. Petrol. Geol. 14:1–103.Google Scholar
Noble, J. P. A. 1970. Biofacies analysis, Cairn Formation of Miette Reef Complex (Upper Devonian), Jasper National Park, Alberta Bull. Can. Petrol. Geol. 18:493–534.Google Scholar
Odum, E. P. 1971. Fundamentals of Ecology. 574 pp. W. B. Saunders Co.; Philadelphia.Google Scholar
Pitcher, M. 1964. Evolution of Chazyan (Ordovician) reefs of eastern United States and Canada. Bull. Can. Petrol. Geol. 12:932–961.Google Scholar
Porter, J. W. 1972. Patterns of species diversity in Caribbean reef corals. Ecology. 53:745–748.CrossRefGoogle Scholar
Porter, J. W. 1974. Community structure of coral reefs on opposite sides of the isthmus of Panama. Science. 186:543–545.CrossRefGoogle Scholar
Potts, D. C. 1978. Differentiation of coral reef populations. Atoll Res. Bull. 220:55–74.CrossRefGoogle Scholar
Riding, R. and Kershaw, S. 1978. Parameterization of stromatoporoid shape. Lethaia. 11:233–242.Google Scholar
Rosen, B. R. 1971. Principle features of coral reef ecology in shallow water environments of Mahé, Seychelles. In: Stoddard, D. R. and Yonge, C. M., eds. Regional Variation in Indian Ocean Coral Reefs. Zool. Soc. London Symp. 28:163–183.Google Scholar
St. Jean, J. Jr. 1971. Paleobiologic considerations of reef stromatoporoids. North Am. Paleontol. Convent. Chicago Proc. 2:1389–1429.Google Scholar
Scatterday, J. W. 1974. Reefs and associated coral assemblages off Bonaire, Netherlands Antilles, and their bearing on Pleistocene and recent reef models. 2nd Int. Coral Reef Symp. Brisbane Proc. 2:85–106.Google Scholar
Scatterday, J. W. 1977. Low-water emergence of Caribbean reefs and effect of exposure upon coral diversity—observations off Bonaire, Netherlands Antilles. In: Frost, S. H., Weiss, M. P., and Saunders, J. B., eds. Reef and Related Carbonates—Ecology and Sedimentology. Am. Assoc. Petrol. Geol. Studies Geol. 4:155–169.Google Scholar
Schlager, W. 1981. The paradox of drowned reefs and carbonate platforms. Bull. Geol. Soc. Am. Pt. 1. 92:197–211.2.0.CO;2>CrossRefGoogle Scholar
Schopf, T. J. M., Collier, K. O., and Bach, B. O. 1980. Relationship of the morphology of stick-like bryozoan at Friday Harbour, Washington to bottom currents, suspended matter, and depth. Paleobiology. 6:466–476.CrossRefGoogle Scholar
Scoffin, T. P., Stearn, C. W., Boucher, D., Frydl, P., Hawkins, C. M., Hunter, I. G., and Macgeachy, J. K. 1980. Calcium carbonate budget of a fringing reef on the west coast of Barbados. Part II—Erosion, sediments and internal structure. Bull. Mar. Sci. 30:475–508.Google Scholar
Scrutton, C. T. 1977. Facies variations in the Devonian limestones of eastern South Devon. Geol. Mag. 114:165–248.CrossRefGoogle Scholar
Seilacher, A. 1970. Arbeitskonzept sur konstruktionsmorphologie. Lethaia. 3:393–396.CrossRefGoogle Scholar
Stearn, C. W. 1972. The relationship of the stromatoporoids to the sclerosponges. Lethaia. 5:369–388.CrossRefGoogle Scholar
Stearn, C. W. 1975. Stromatoporoid assemblages: ancient Wall reef complex, (Devonian), Alberta. Can. J. Earth Sci. 12:1631–1667.CrossRefGoogle Scholar
Stearn, C. W. and Riding, R. 1973. Forms of the hydrozoan Millepora on a recent coral reef. Lethaia. 6:187–200.CrossRefGoogle Scholar
Stearn, C. W., Scoffin, T. P., and Martindale, W. 1977. Calcium carbonate budget of a fringing reef on the west coast of Barbados. Pt. I—Zonation and productivity. Bull. Mar. Sci. 27:479–510.Google Scholar
Stephenson, T. A. and Stephenson, A. 1933. Growth and asexual reproduction in corals Great Barrier Reef Exped., 1928–29. 3:167–217.Google Scholar
Stock, C. W. 1979. Upper Silurian (Pridoli) stromatoporoidea of New York. Bull. Am. Paleontol. 76(308):293–389.Google Scholar
Stoddart, D. R. 1969. Ecology and morphology of Recent coral reefs. Biol. Rev. 44:433–498.CrossRefGoogle Scholar
Storr, J. F. 1964. Ecology and oceanography of the coral reef tract, Abaco Island, Bahamas. Geol. Soc. Am. Spec. Paper 79:1–98.Google Scholar
Taylor, J. D. 1968. Coral reefs and associated invertebrate communities (mainly molluscan) around Mahé, Seychelles. Phil. Trans. R. Soc. London B254:129–206.Google Scholar
Thomas, R. D. K. 1975. Functional morphology, ecology and evolutionary conservatism in Glycimeridae (Bivalvia). Palaeontology. 18:217–254.Google Scholar
Thomas, R. D. K. 1979. Morphology, constructional. Pp. 482–487. In: Fairbridge, R. W. and Jablonski, D., eds. The Encyclopedia of Paleontology. Dowden, Hutchison and Ross; Stroudsburg.CrossRefGoogle Scholar
Tsien, H. H. 1974. Paleoecology of Middle Devonian and Frasnian in Belgium. Int. Symp. Belge Micropaleontol. Limits, Sept. 1974. 12:1–53.Google Scholar
Vopni, L. K. and Lerbekmo, J. F. 1972. Sedimentology and ecology of the Horn Plateau Formation, a Middle Devonian coral reef, Northwest Territories. Canada. Geol. Rundschau. 61:626–646.CrossRefGoogle Scholar
Walker, K. R. and Alberstadt, L. P. 1975. Ecological succession as an aspect of structure in fossil communities. Paleobiology. 1:238–257.CrossRefGoogle Scholar
Weber, J. N., Weber, P. H., and Boker, P. A. 1977. A new theory for the spatial distribution of reef corals. Geol. Soc. Am. Abstr. with Programs. 7:1313.Google Scholar
Wells, J. W. 1954. Recent corals of the Marshall Islands. Bikini and nearby Atolls, Part 2: Oceanography (Biologic). U.S. Geol. Survey Prof. Pap. 206-I:385–486.Google Scholar
Wethey, D. S. and Porter, J. W. 1976. Sun and shade differences in productivity of reef corals. Nature. 262:281–282.CrossRefGoogle Scholar
Wijsman-Best, M. 1972. Habitat modification of reef corals (Faviidae) and its consequences for taxonomy. 2nd Int. Coral Reef Sympos. Brisbane Proc. 2:217–228.Google Scholar
Wood-Jones, F. 1910. On the growth forms and supposed species in corals. Proc. Zool. Soc. London. Pp. 518–556.Google Scholar
Yonge, C. M. 1973. The nature of reef-building (Hermatypic) corals. Bull. Mar. Sci. 23:1–16.Google Scholar
Zankl, H. and Schroeder, J. H. 1972. Interaction of genetic processes in Holocene reefs off N. Eleuthera Island, Bahamas. Geol. Rundschau. 61:520–541.CrossRefGoogle Scholar