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The life orientation of concavo-convex brachiopods: overturning the paradigm

Published online by Cambridge University Press:  08 February 2016

Halard L. Lescinsky*
Affiliation:
Department of Geology, University of California, Davis, California 95616

Abstract

Concavo-convex brachiopods are generally assumed to have lived with their convex valves against the sediment. This orientation is based primarily on the a priori assumption that the upturned commissure would prevent fouling of the brachiopods' mantle cavity. Here, I present epibiontic and taphonomic evidence from orthids and nonproductid strophomenids that concavo-convex brachiopods lived in the reverse life orientation, with the convex valve on top.

Ten testable epibiontic and taphonomic criteria are proposed to establish the uppermost valve during encrustation and whether epibionts encrusted primarily live or dead hosts. The criteria are evaluated for 11 collections of Ordovician and Devonian nonproductid concavo-convex brachiopods that contain a total of over 500 brachiopods and 4000 epibionts. In all cases, the results support a convex-up orientation and the encrustation of live hosts. Five criteria concern epibiont growth patterns and show that (1) epibionts predominate on convex valves, (2) epibionts are not restricted to shell margins, (3) distinct exposed and cryptic faunas, as predicted theoretically and described in previous studies, exist on convex and concave valves respectively, (4) epibiont colonies were often truncated by the commissures and hinges of live hosts, and (5) growth patterns of some epibionts indicate live hosts. Five taphonomic criteria show that (1) concavo-convex shells have greater epibiont cover than other morphologies, (2) internal surfaces of disarticulated specimens are rarely encrusted, (3) interiors of articulated specimens are rarely mud-filled, (4) standard taphofacies indicators suggest little postmortem exposure, and (5) sedimentological reconstructions suggest rapid burial.

Convex-up brachiopods suggest that the Paleozoic mud bottoms they lived on were probably firmer than usually assumed and that a more complex functional interpretation of concavo-convexity is required. Hydrodynamic stability was important to many concavo-convex, plano-convex, and unequally bi-convex brachiopods. Brachiopods with these morphologies probably lived in the more stable orientation on their concave or flat valve. Productid brachiopods, although also concavo-convex, lived convex valve down and are ecologically distinct from earlier concavo-convex taxa. Productids were anchored, semi-infaunally, by spines and were more similar to Mesozoic oysters in orientation and shell function.

Type
Articles
Copyright
Copyright © The Paleontological Society 

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References

Literature Cited

Ager, D. V. 1967. Brachiopod palaeoecology. Earth Science Reviews 3:157179.CrossRefGoogle Scholar
Alexander, R. R. 1975. Phenotypic lability of the brachiopod Rafinesquina alternata (Ordovician) and its correlation with the sedimentologic regime. Journal of Paleontology 49:607618.Google Scholar
Alexander, R. R. 1984. Comparative hydrodynamic stability of brachiopod shells on current-scoured arenaceous substrates. Lethaia 17:1732.CrossRefGoogle Scholar
Alexander, R. R. 1986. Resistance to and repair of shell breakage induced by durophages in Late Ordovician brachiopods. Journal of Paleontology 60:273285.CrossRefGoogle Scholar
Alexander, R. R. 1989. Influence of valve geometry, ornamentation, and microstructure on fractures in Late Ordovician brachiopods. Lethaia 22:133147.CrossRefGoogle Scholar
Alexander, R. R. 1990. Disarticulated shells of Late Ordovician brachiopods: inferences on strength of hinge and valve architecture. Journal of Paleontology 64:524531.CrossRefGoogle Scholar
Alexander, R. R., and Scharpf, C. D. 1990. Epizoans on Late Ordovician brachiopods from southeastern Indiana. Historical Biology 4:179202.CrossRefGoogle Scholar
Allen, J. A. 1953. Observations on the epifauna of the deep water muds of the Clyde Sea area, with special reference to Chlamys septemradiata (Muller). Journal of Animal Ecology 22:240260.CrossRefGoogle Scholar
Alvarez, F., and Taylor, P. 1987. Epizoan ecology and interactions in the Devonian of Spain. Palaeogeography Palaeoclimatology Palaeoecology 61:1731.CrossRefGoogle Scholar
Ausich, W. I., and Gurrola, R. A. 1979. Two boring organisms in a lower Mississippian community of southern Indiana. Journal of Paleontology 53:335344.Google Scholar
Baird, G. C., Brett, C. E., and Frey, R. C. 1989. Hitchhiking epizoans on orthoconic cephalopods: preliminary review of the evidence and its implications. Senckenbergiana Lethaea 69:439465.Google Scholar
Baird, G. C., Brett, C. E., and Tomlinson, J. J. 1990. Host specific acrothoracid barnacles on Middle Devonian platyceratid gastropods. Historical Biology 4:221224.CrossRefGoogle Scholar
Barthel, K. W., and Barth, W. 1972. Paleoecologic specimens from the Devonian of Bolivia. Neues Jahrbuch für Geologie und Palaontologie Monatshefte 10:573581.Google Scholar
Bassett, M. G. 1984. Life strategies of Silurian brachiopods. Special Papers in Palaeontology 32:237263.Google Scholar
Baumiller, T. K. 1990. Non-predatory drilling of Mississippian crinoid by platyceratid gastropods. Palaeontology 33:743748.Google Scholar
Bordeaux, Y. L., and Boyajian, G. E. 1991. Simulating random encrustation patterns on a growing individual: a re-examination of epibionts as tools in autecology and taphonomy. Geological Society of America, Abstracts with Programs A166.Google Scholar
Bordeaux, Y. L., and Brett, C. E. 1990. Substrate specific associations of epibionts on Middle Devonian brachiopods: implications for paleoecology. Historical Biology 4:221224.CrossRefGoogle Scholar
Brett, C. E., and Baird, G. C. 1986. Comparative taphonomy: a key to paleoenvironmental interpretation based on fossil preservation. Palaios 1:207227.CrossRefGoogle Scholar
Brett, C. E., and Baird, G. C. 1994. Taphonomic approaches to time resolution in stratigraphy: example from Paleozoic marine mudrocks. Short Course in Paleontology 6:250274.Google Scholar
Brett, C. E., and Bordeaux, Y. L. 1991. Taphonomy of brachiopods from a Middle Devonian shell bed: implications for the genesis of skeletal accumulations. Pp. 219226in MacKinnon, D. I., Lee, D. E., and Campbell, J. D., eds. Brachiopods through time. Balkema, Rotterdam.Google Scholar
Brett, C. E., Miller, K. B., and Baird, G. C. 1990. A temporal hierarchy of paleoecologic processes within a Middle Devonian epeiric sea. Pp. 178209in Miller, W. I., ed. Paleocommunity temporal dynamics: the long-term development of multispecies assemblages. Paleontological Society Special Publication 5.Google Scholar
Brett, C. E., Boucot, A. J., and Jones, B. 1993. Absolute depths of Silurian benthic assemblages. Lethaia 26:2540.CrossRefGoogle Scholar
Brice, D., and Hou, H. F. 1992. “Blisters” in a Famennian cyrtospiriferid brachiopod from Hunan (South China). Palaeogeography Palaeoclimatology Palaeoecology 94:253260.CrossRefGoogle Scholar
Brice, D., and Mistiaen, B. 1992. Epizoaires des brachiopodes Frasniens de Ferques (Boulonnais: Nord de la France). Geobios Memoir Special 14:4558.CrossRefGoogle Scholar
Brunton, C. H. C. 1985. Growth and shell shape in Productacean brachiopods. Bulletin of the British Museum of Natural History (Geology) 38:273281.Google Scholar
Cameron, B. 1969. Paleozoic shell-boring annelids and their trace fossils. American Zoologist 9:689703.CrossRefGoogle Scholar
Chatterton, B. D. E. 1975. A commensal relationship between a small filter feeding organism and Australian Devonian spiriferid brachiopods. Paleobiology 1:371378.CrossRefGoogle Scholar
Conway Morris, S. 1985. The middle Cambrian metazoan Wiwaxia corrugata (Mathew) from the Burgess Shale and Ogygopsis Shale, British Columbia, Canada. Philosophical Transactions of the Royal Society of London, Series B 307:507582.Google Scholar
Copper, P. 1967. Adaptations and life habits of Devonian atrypid brachiopods. Palaeogeography Palaeoclimatology Palaeoecology 3:363379.CrossRefGoogle Scholar
Cowen, R. 1979. Functional morphology. Pp. 487491in Fairbridge, R. W. and Jablonski, D., eds. The encyclopedia of paleontology. Dowden, Hutchinson and Ross, Stroudsburg, Penn.CrossRefGoogle Scholar
Daley, G. M. 1993. Passive deterioration of shelly material: a study of the recent eastern Pacific articulate brachiopod Terebratalia transversa Sowerby. Palaios 8:226232.CrossRefGoogle Scholar
Eckert, J. D. 1988. The ichnogenus Tremichnus in the Lower Silurian of western New York. Lethaia 21:281283.CrossRefGoogle Scholar
Emig, C. C. 1990. Examples of post-mortality alteration in Recent brachiopod shells and (paleo)ecological consequences. Marine Biology 104:233238.CrossRefGoogle Scholar
Fenton, C. L., and Fenton, M. A. 1932. Orientation and injury in the genus Atrypa. American Midland Naturalist 13:6374.CrossRefGoogle Scholar
Franzen, C. 1974. Epizoans on Silurian-Devonian crinoids. Lethaia 7:287301.CrossRefGoogle Scholar
Gill, G. A., and Coates, A. G. 1977. Mobility, growth patterns and substrate in some fossil and recent corals. Lethaia 10:119134.CrossRefGoogle Scholar
Grant, R. E. 1963. Unusual attachment of a Permian linoproductid brachiopod. Journal of Paleontology 37:134140.Google Scholar
Grant, R. E. 1975. Methods and conclusions in functional analysis: a reply. Lethaia 8:3133.CrossRefGoogle Scholar
Harvey, P. H., and Pagel, M. D. 1991. The comparative method in evolutionary biology. Oxford University Press.CrossRefGoogle Scholar
Hay, H. B. 1992. Paleogeography and paleoenvironments, Fairview through Whitewater Formations (Upper Ordovician, southeastern Indiana and southwestern Ohio). Pp. 174199in Davis, R. A. and Cuffey, R. J., eds. Sampling the layer cake that isn't: the stratigraphy and paleontology of the type-Cincinnatian. Ohio Geological Survey, Columbus.Google Scholar
Hoare, R. D., and Steller, D. L. 1967. A Devonian brachiopod with epifauna. Ohio Journal of Science 67:291297.Google Scholar
Holland, S. M. 1988. Taphonomic effects of sea-floor exposure on an Ordovician brachiopod assemblage. Palaios 3:588597.CrossRefGoogle Scholar
Holland, S. M. 1992. Sequence stratigraphy of the Cincinnatian Series (Upper Ordovician, Cincinnati Ohio region). Pp. 199220in Davis, R. A. and Cuffey, R. J., eds. Sampling the layer cake that isn't: the stratigraphy and paleontology of the type-Cincinnatian. Ohio Geological Survey, Columbus.Google Scholar
Hurst, J. M. 1974. Selective epizoan encrustation of some Silurian brachiopods from Gotland. Palaeontology 17:423429.Google Scholar
Jackson, J. B. C. 1977. Competition on marine hard substrata: the adaptive significance of solitary and colonial strategies. American Naturalist 111:743767.CrossRefGoogle Scholar
Jackson, J. B. C. 1983. Biological determinants of present and past sessile animal distributions. Pp. 39120in Tevesz, M. J. S. and McCall, P. L., eds. Biotic interactions in recent and fossil benthic communities. Plenum, New York.CrossRefGoogle Scholar
Jackson, J. B. C., and Winston, J. E. 1982. Ecology of cryptic coral reef communities. I. Distribution and abundance of major groups of encrusting organisms. Journal of Experimental Marine Biology and Ecology 57:135–47.CrossRefGoogle Scholar
Kase, T. 1986. Mode of life of the Silurian uncoiled gastropod Semitubina sakoi n. sp. from Japan. Lethaia 19:327337.CrossRefGoogle Scholar
Keough, M. J. 1984. Dynamics of the epifauna of the bivalve Pinna bicolor: interactions among recruitment, predation and competition. Ecology 65:677688.CrossRefGoogle Scholar
Kershaw, S. 1980. Cavities and cryptic faunas beneath non-reef stromatoporoids. Lethaia 13:327338.CrossRefGoogle Scholar
Kesling, R. V., and Chilman, R. B. 1975. Strata and megafossils of the Middle Devonian Silica Formation. Papers on Paleontology 8:1408.Google Scholar
Kesling, R. V., Hoare, R. D., and Sparks, D. K. 1980. Epizoans of the Middle Devonian brachiopod Paraspirifer bownockeri: their relationships to one another and to their host. Journal of Paleontology 54:11411154.Google Scholar
Kidwell, S. M., and Jablonski, D. 1983. Taphonomic feedback: ecological consequences of shell accumulation. Pp. 195250in Tevesz, M. J. S. and McCall, P. L., eds. Biotic interactions in recent and fossil communities. Plenum, New York.CrossRefGoogle Scholar
Kidwell, S. M., and Jablonski, D. 1990. Phanerozoic evolution of macroinvertebrate shell accumulations: preliminary data from the Jurassic of Britain. Pp. 309327in Miller, W. III, ed. Paleocommunity temporal dynamics: the long-term development of multispecies assemblages. The Paleontological Society Special Publication 5.Google Scholar
Kobluk, D. R., and James, N. P. 1979. Cavity-dwelling organisms in Lower Cambrian patch reefs from southern Labrador. Lethaia 12:193218.CrossRefGoogle Scholar
Koch, W. F. 1973. Quantitative study of relationships between host brachiopods and epizoans in the Middle Devonian Silica Formation. M.S. thesis. University of Michigan.Google Scholar
LaBarbera, M. 1981. The ecology of Mesozoic Gryphaea, Exogyra, and llymatogyra (Bivalvia: Mollusca) in a modern ocean. Paleobiology 7:510526.CrossRefGoogle Scholar
Lamont, A. 1934. Lower Paleozoic brachiopods of the Girvan District, with suggestions on morphology in relation to environment. Annals and Magazine of Natural History, Series 10 14:161184.CrossRefGoogle Scholar
Lescinsky, H. L. 1993. Taphonomy and paleoecology of epibionts on the scallops Chlamys hastata (Sowerby 1843) and Chlamys rubida (Hinds 1845). Palaios 8:267277.CrossRefGoogle Scholar
Lescinsky, H. L., and Benninger, L. 1994. Pseudo-borings and predator traces: artifacts of pressure dissolution in fossiliferous shales. Palaios 9:599604.CrossRefGoogle Scholar
Lohse, D. P. 1993. The importance of secondary substratum in a rocky intertidal community. Journal of Experimental Marine Biology and Ecology 166:117.CrossRefGoogle Scholar
Martindale, W. 1992. Calcified epibionts as palaeoecological tools: examples from the Recent and Pleistocene reefs of Barbados. Coral Reefs 11:167–77.CrossRefGoogle Scholar
McKinney, F. K., and McKinney, M. J. 1993. Larval behaviour and choice of settlement site: correlation with environmental distribution pattern in an erect bryozoan. Facies 29:119132.CrossRefGoogle Scholar
McKinney, F. K., Broadhead, T. W., and Gibson, M. A. 1990. Coral-bryozoan mutualism: structural innovation and greater resource exploitation. Science 248:466467.CrossRefGoogle Scholar
Menard, H. W., and Boucot, A. J. 1951. Experiments on the movement of shells by water. American Journal of Science 249:131151.CrossRefGoogle Scholar
Meyer, D. L. 1990. Population paleoecology and comparative taphonomy of two edrioasteroid (Echinodermata) pavements: Upper Ordovician of Kentucky and Ohio. Historical Biology 4:155179.CrossRefGoogle Scholar
Meyer, D. L., Wahlman, G. P., and Elias, R. J. 1978. Paleoecology of an Upper Ordovician edrioasteroid bed. Geological Society of America Abstracts with Programs 10:278.Google Scholar
Morris, R. W., and Rollins, H. B. 1971. The distribution and paleoecological interpretation of Cornulites in the Waynesville Formation (Upper Ordovician) of southwestern Ohio. Ohio Journal of Science 71:159170.Google Scholar
Morris, P. J., Linsley, R. M., and Cottrell, J. F. 1991. A middle Devonian symbiotic relationship involving a gastropod: a trepostomatous bryozoan, and an inferred secondary occupant. Lethaia 24:5567.CrossRefGoogle Scholar
Morton, B. 1981. The anomalodesmata. Malacologia 21:3560.Google Scholar
Muir-Wood, H., and Cooper, G. A. 1960. Morphology, classification and life habits of the Productoidea (Brachiopoda). Geological Society of America Memoir 81:1447.CrossRefGoogle Scholar
Nagle, J. S. 1967. Wave and current orientation of shells. Journal of Sedimentary Petrology 37:11241138.Google Scholar
Nield, E. W. 1986. Noncryptic encrustation and preburial fracturing in stromatoporoids from the Upper Visby Beds of Gotland, Sweden. Palaeogeography Palaeoclimatology Palaeoecology 55:3544.CrossRefGoogle Scholar
Parsons, K. M., Brett, C. E., and Miller, K. B. 1988. Taphonomy and depositional dynamics of Devonian shell-rich mudstones. Palaeogeography Palaeoclimatology Palaeoecology 63:109139.CrossRefGoogle Scholar
Paul, C. 1980. The natural history of fossils. Weidenfeld and Nicolson, London.Google Scholar
Pickerill, R. K. 1976. Vermiforichnus borings from the Ordovician of central Wales. Geological Magazine 113:159164.CrossRefGoogle Scholar
Pitrat, C., and Rogers, F. 1978. Spinocyrtia and its epibionts in the Traverse Group (Devonian) of Michigan. Journal of Paleontology. 52:13151324.Google Scholar
Pope, J. K. 1976. Comparative morphology and shell histology of the Ordovician Strophomenacea (Brachiopoda). Palaeontographica Americana. 49:129213.Google Scholar
Powers, B. G., and Ausich, W. I. 1990. Epizoan associations in a Lower Mississippian paleocommunity (Borden Group: Indiana: United States of America). Historical Biology 4:245265.CrossRefGoogle Scholar
Reid, S. R., and Broadhead, T. W. 1983. Epibiont encrustations as indicators of life and hydrodynamic orientation of Leptaena (Brachiopoda: Strophomenida). Geological Society of America Abstracts with Programs 15:669.Google Scholar
Richards, R. P. 1972. Autecology of Richmondian brachiopods (Late Ordovician of Indiana and Ohio). Journal of Paleontology 46:386405.Google Scholar
Richards, R. P. 1974. Ecology of the Cornulitidae. Journal of Paleontology 48:514523.Google Scholar
Rodriguez, J., and Gutschick, R. C. 1977. Barnacle borings in live and dead hosts from the Louisiana Limestone (Famennian) of Missouri. Journal of Paleontology 51:718724.Google Scholar
Rudwick, M. J. S. 1964. The inference of function from structure in fossils. British Journal of Philosophy of Science 15:2740.CrossRefGoogle Scholar
Rudwick, M. J. S. 1970. Fossil and living brachiopods. Hutchinson, London.Google Scholar
Savarese, M. 1994. Taphonomic and paleoecologic implications of flow-induced forces on concavo-convex articulate brachiopods: an experimental approach. Lethaia 27:301312.CrossRefGoogle Scholar
Schumann, D. 1967. Die lebensweise von Mucrospirifer Grabau 1931 (Brachiopoda). Palaeogeography Palaeoclimatology Palaeoecology 3:381392.CrossRefGoogle Scholar
Seilacher, A. 1968. Swimming habits of belemnites—recorded by boring barnacles. Palaeogeography Palaeoclimatology Palaeoecology 4:279285.CrossRefGoogle Scholar
Seilacher, A. 1969. Paleoecology of boring barnacles. American Zoologist 9:705719.CrossRefGoogle Scholar
Sheehan, P. M. 1978. The hinging mechanisms of brachiopods—taphonomic considerations. Journal of Paleontology 52:748.Google Scholar
Solle, G. 1968. Hederelloidea (Cyclostomata) und einige ctenostome Bryozoen aus dem Rheinischen Devon. Hessisches Landesamt fur Bodenforschung Abhandlungen 54:140.Google Scholar
Speyer, S. E., and Brett, C. E. 1991. Background and episodic processes in fossil assemblage preservation. Pp. 502545in Allison, P. A. and Briggs, D. E. G., eds. Taphonomy: releasing the data locked in the fossil record. Plenum, New York.Google Scholar
Spjeldnaes, N. 1975. Silurian bryozoans which grew in the shade, Pp. 415424in Pouyet, S., ed. Bryozoa 1974. Proceedings of the Third Conference, International Bryozoology Association. Universite Claude Bernard, Lyon.Google Scholar
Spjeldnaes, N. 1984. Epifauna as a tool in autecological analysis of Silurian brachiopods. Special Papers in Palaeontology 32:225235.Google Scholar
Stanley, S. M. 1968. Post-Paleozoic adaptive radiation of infaunal bivalve molluscs—a consequence of mantle fusion and siphon formation. Journal of Paleontology 42:214229.Google Scholar
Stanley, S. M. 1970. Relation of shell form to life habits of the Bivalvia (Mollusca). Geological Society of America Memoir 125.CrossRefGoogle Scholar
Steller, D. L., 1965. The epifaunal elements of the Brachiopoda of the Silica Formation. M.A. thesis. Bowling Green University, Ohio.Google Scholar
Takemura, Y., and Okutani, T. 1955. Notes on animals attached to the shells of the Silver-Lip pearl oyster, Pinctada maxima (Jameson), collected from the “East” fishing ground of the Arafura Sea. Bulletin of the Japanese Society of Scientific Fisheries 21:92101.CrossRefGoogle Scholar
Thayer, C. W. 1974. Substrate specificity of Devonian epizoans. Journal of Paleontology 48:881894.Google Scholar
Thayer, C. W. 1975a. Morphologic adaptations of benthic invertebrates to soft substrata. Journal of Marine Research 33:177189.Google Scholar
Thayer, C. W. 1975b. Diductor muscles of brachiopods: active or passive? Paleobiology 1:4447.CrossRefGoogle Scholar
Thayer, C. W. 1983. Sediment-mediated biological disturbance and the evolution of marine benthos. Pp. 480626in Tevesz, M. J. S. and McCall, P. L., eds. Biotic interactions in recent and fossil communities. Plenum, New York.Google Scholar
Titus, R. 1992. Clinal variation, heterochrony, and facies in the Trentonian Sowerbyella lineage (Ordovician, New York State). Journal of Paleontology 66:758771.CrossRefGoogle Scholar
Unklesby, A. G., and Niewoehover, W. B. 1959. Attachment loops on infant brachiopods from the Louisiana Limestone in Missouri. Journal of Paleontology 33:547549.Google Scholar
Velbel, D. B. 1985. Ichnologic, taphonomic, and sedimentologic clues to the deposition of Cincinnatian shales (Upper Ordovician), Ohio, United States of America Society of Economic Paleontologists and Mineralogists Special Publication 35:299307.Google Scholar
Vogel, K., Golubic, S., and Brett, C. E. 1987. Endolith associations and their relation to facies distribution in the Middle Devonian of New York State. Lethaia 20:263290.CrossRefGoogle Scholar
Walker, K. R., and Diehl, W. W. 1985. The role of marine cementation in the preservation of Lower Palaeozoic assemblages. Philosophical Transactions of the Royal Society of London B311:143–53.Google Scholar
Walker, S. E. 1992. Criteria for recognizing marine hermit crabs in the fossil record using gastropod shells. Journal of Paleontology 66:535558.CrossRefGoogle Scholar
Ward, M. A., and Thorpe, J. P. 1989. Assessment of space utilization in a subtidal temperate bryozoan community. Marine Biology 103:215224.CrossRefGoogle Scholar
Watkins, R. 1981. Epizoan ecology of the type Ludlow Series (Upper Silurian), England. Journal of Paleontology 55:2932.Google Scholar
Wilson, M. A. 1987. Ecological dynamics on pebbles, cobbles and boulders. Palaios 2:594599.CrossRefGoogle Scholar
Wood, R., Dickson, J., and Kirkland, G. B. 1994. Turning the Capitan reef upside down—a new appraisal of the ecology of the Permian Capitan Reef, Guadalupe Mountains, Texas and New Mexico. Palaios 9:422427.CrossRefGoogle Scholar
Yund, P. O., and Parker, H. M. 1989. Population structure of the colonial hydroid Hydractinia sp. nov. C. in the Gulf of Maine. Journal of Experimental Marine Biology and Ecology 125:6382.CrossRefGoogle Scholar
Zumwalt, G. S., and DeLaca, T. E. 1980. Utilization of brachiopod feeding currents by epizoic foraminifera. Journal of Paleontology 54:477484.Google Scholar