Hostname: page-component-586b7cd67f-rdxmf Total loading time: 0 Render date: 2024-11-23T21:52:15.762Z Has data issue: false hasContentIssue false
  • English
  • Français

Possible propulsion modes in Graptoloidea: a new model for graptoloid locomotion

Published online by Cambridge University Press:  08 February 2016

Michael J. Melchin  and
M. Edwin DeMont
Michael J. Melchin
Affiliation:
Department of Geology, St. Francis Xavier University, Post Office Box 5000, Antigonish, Nova Scotia B2G 2W5, Canada
M. Edwin DeMont
Affiliation:
Department of Biology, St. Francis Xavier University, Post Office Box 5000, Antigonish, Nova Scotia B2G 2W5, Canada
Get access Rights & Permissions [Opens in a new window]

Abstract

The mode of locomotion of any swimming animal is constrained by its size, architecture, and phylogenetic history. Considering these factors and the range of locomotory systems used by extant zooplankton, the range of possible modes of locomotion for graptoloids can be effectively limited. Three assumptions have been made: (1) graptoloids did not use a mode of locomotion unknown among modern organisms; (2) all graptoloids employed essentially the same mode of locomotion except, possibly, in their early growth stages; and (3) graptoloids did not rely entirely on passive buoyancy—no extant zooplankton groups in the size range of the graptoloids do. Structures that increase buoyancy or drag are often found in actively swimming zooplankton. They enhance feeding efficiency and reduce sinking rates during nonswimming periods.

The modes of locomotion utilized by extant zooplankton groups are ciliary propulsion, elongate body undulation, jet propulsion, rowing with skeletonized appendages, and rowing or undulation with muscular appendages. Of these, all but the last can be rejected for the graptoloids on the basis of scale or architecture. It is concluded that graptoloids probably used a rowing or undulatory motion with muscular appendages for swimming. Using a pterobranch model for the graptoloid zooids, the lophophore is considered an unlikely propulsive structure because the design requirements would conflict with those of a ciliarly suspension-feeding organ. Winglike, lateral extensions of the muscular cephalic shield, the same structure used for creeping locomotion in the benthic pterobranchs, is regarded as the most likely propulsive organ, analogous to the pteropod swimming wings.

Type
Research Article
Information
Paleobiology , Volume 21 , Issue 1 , Winter 1995 , pp. 110 - 120
Copyright
Copyright © The Paleontological Society 

Access options

Get access to the full version of this content by using one of the access options below. (Log in options will check for institutional or personal access. Content may require purchase if you do not have access.)

References

Literature Cited

Bates, D. E. B. 1987. The density of graptoloid skeletal tissue, and its implication for the volume and density of the soft tissue. Lethaia 20:149–156.CrossRefGoogle Scholar
Bates, D. E. B., and Kirk, N. H. 1984. Autecology of Silurian graptoloids. Special Papers in Palaeontology 32:121–139.Google Scholar
Bates, D. E. B., and Kirk, N. H. 1985. Graptolites, a fossil case-history of evolution from sessile, colonial animals to automobile superindividuals. Proceedings of the Royal Society of London B228:207–224.Google Scholar
Berry, W. B. N., and Boucot, A. J. 1972. Silurian graptolite depth zonation. Proceedings of the 24th International Geological Congress, Montreal 7:59–65.Google Scholar
Berry, W. B. N., Wilde, P., and Quinby-Hunt, M. S. 1987. The oceanic nonsulphide oxygen minimum zone: a habitat for graptolites? Bulletin of the Geological Society of Denmark 35:103–114.CrossRefGoogle Scholar
Bjerreskov, M. 1978. Discoveries on graptolites by x-ray studies. Acta Palaeontologica Polonica 23:364–376.Google Scholar
Bone, Q., and Trueman, E. R. 1982. Jet propulsion of the calycophoran siphonophores Chelophyes and Alypopsis. Journal of the Marine Biological Association of the United Kingdom 62:263–276.CrossRefGoogle Scholar
Bone, Q., and Trueman, E. R. 1983. Jet propulsion in salps (Tunicata: Thaliacea). Journal of Zoology, London 201:481–506.CrossRefGoogle Scholar
Chia, F.-S., Buckland-Nicks, J., and Young, C. M. 1984. Locomotion of marine invertebrate larvae: a review. Canadian Journal of Zoology 62:1205–1222.CrossRefGoogle Scholar
Clark, T. H. 1972. Stratigraphy and structure of the St. Lawrence Lowland of Quebec. 24th International Geological Congress, Montreal. Guidebook to Excursion 52.Google Scholar
Cooper, R. A., Fortey, R. A., and Lindholm, K. 1991. Latitudinal and depth zonation of early Ordovician graptolites. Lethaia 24:199–218.CrossRefGoogle Scholar
Crowther, P. R. 1981. Fine structure of the graptolite periderm. Special Papers in Palaeontology 26.Google Scholar
Daniel, T., Jordan, C., and Grunbaum, D. 1992. Hydromechanics of swimming. Pp. 17–49in Alexander, R. M., ed. Advances in comparative and environmental physiology, 11. Mechanics of animal locomotion. Springer, Berlin.CrossRefGoogle Scholar
Davenport, J., and Bebbington, A. 1990. Observations on the swimming and buoyancy of some thesosomatous pteropod gastropods. Journal of Molluscan Studies 56:487–497.CrossRefGoogle Scholar
DeMont, M. E. 1992. Locomotion of soft bodied animals. Pp. 167–190in Alexander, R. M., ed. Advances in comparative and environmental physiology, 11. Mechanics of animal locomotion. Springer, Berlin.CrossRefGoogle Scholar
DeMont, M. E., and Hokkanen, E. I. 1992. Hydrodynamics of animal movement. Pp. 263–284in Biewener, A. A., ed. Biomechanics—structures and systems. A practical approach. Oxford University Press.CrossRefGoogle Scholar
Dilly, P. N. 1985. The habitat and behavior of Cephalodiscus gracilis (Pterobranchia: Hemichordata). Journal of Zoology, London 207:223–239.CrossRefGoogle Scholar
Dilly, P. N. 1986. Modern pterobranchs: observations on their behavior and tube building. Pp 261–269in Hughes, C. P. and Rickards, R. B., eds. Palaeoecology and biostratigraphy of graptolites. Geological Society Special Publication 20.Google Scholar
Dilly, P. N. 1993. Cephalodiscus graptolitoides sp. nov. a probable extant graptolite. Journal of Zoology, London 229:69–78.CrossRefGoogle Scholar
Emlet, R. B., and Strathmann, R. R. 1985. Gravity, drag and feeding currents of small zooplankton. Science 228:1016–1017.CrossRefGoogle ScholarPubMed
Erdtmann, B.-D. 1973. Life forms and feeding habits of graptolites. Pp. 523–532in Boardman, R. S., Cheetham, A. H., and Oliver, W. A., eds. Animal colonies, development and function through time. Downden, Hutchinson and Ross, Stroudsberg, Penn.Google Scholar
Erdtmann, B.-D. 1976. Ecostratigraphy of Ordovician graptoloids. Pp. 621–643in Bassett, M. G., ed. The Ordovician system. University of Wales Press, Cardiff.Google Scholar
Finney, S. C. 1979. Mode of life of planktonic graptolites: flotation structure in Ordovician Decellograptus sp. Paleobiology 5:31–39.CrossRefGoogle Scholar
Finney, S. C. 1985. Paired pleural disks in Dicaulograptus cumdiscus n. sp. Lethaia 18:361–368.CrossRefGoogle Scholar
Finney, S. C., and Jacobson, S. R. 1985. Flotation devices in planktic graptolites. Lethaia 18:349–359.CrossRefGoogle Scholar
Gladfelter, W. G. 1973. A comparative analysis of the locomotory systems of medusoid Cnidaria. Helgoländer wiss Meeresunters 25:228–272.CrossRefGoogle Scholar
Goldman, D., and Mitchell, C. E. 1994. Three-dimensional graptolites from the upper Middle Ordovician Neuville Formation, Quebec. New York State Museum, Bulletin 481:87–100.Google Scholar
Harbison, G. R. 1992. Observations on the swimming and buoyancy of Cymbulia peroni (Gastropoda: Thecosomata) made from a submersible. Journal of the Marine Biological Association of the United Kingdom 72:435–446.CrossRefGoogle Scholar
Jordan, C. E. 1991. A model of rapid-start swimming at intermediate Reynolds number: undulatory locomotion in the chaetognath Sagitta elegans. Journal of Experimental Biology 163:119–137.CrossRefGoogle Scholar
Kirk, N. H. 1978. Mode of life of graptolites. Acta Palaeontologica Polonica 23:427–448.Google Scholar
Kirk, N. H. 1990. Juvenile sessility, vertical automobility, and passive lateral transport as factors in graptoloid evolution. Modern Geology 14:153–187.Google Scholar
Kirk, N. H. 1991. Construction, form and function in the Graptolithina: a review. Modern Geology 15:287–311.Google Scholar
Kozlowski, R. 1971. Early development stages and mode of life of graptolites. Acta Palaeontologica Polonica 16:313–343.Google Scholar
Lalli, C. M., and Gilmer, R. W. 1989. Pelagic snails. The biology of holoplanktic gastropod molluscs. Stanford University Press, Stanford, Calif.CrossRefGoogle Scholar
Lester, S. M. 1985. Cephalodiscus sp. x(Hemichordata: Pterobranchia): observations of functional morphology, behavior and occurrence in shallow water around Bermuda. Marine Biology 85:263–268.CrossRefGoogle Scholar
Lighthill, M. J. 1969. Hydromechanics of aquatic animal propulsion. Annual Review of Fluid Mechanics 1:413–446.CrossRefGoogle Scholar
Madin, L. P. 1990. Aspects of jet propulsion in salps. Canadian Journal of Zoology 68:765–777.CrossRefGoogle Scholar
Mileikovsky, S. A. 1973. Speed of active movement of pelagic larvae of marine bottom invertebrates and their ability to regulate their vertical position. Marine Biology 23:11–17.CrossRefGoogle Scholar
Mitchell, C. E., and Carle, K. J. 1986. The nematularium of Pseudoclimacograptus scharenbergi (Lapworth) and its secretion. Palaeontology 29:373–390.Google Scholar
Raymont, J. E. G. 1983. Plankton and productivity in the oceans, Vol. 2, Zooplankton. Pergamon, Oxford.Google Scholar
Rickards, R. B. 1975. Palaeoecology of the Graptolithina, an extinct class of the Phylum Hemichordata. Biological Reviews 50:397–436.CrossRefGoogle Scholar
Rickards, R. B., and Stait, B. A. 1984. Psigraptus, its classification, evolution and zooid. Alcheringa 8:101–111.CrossRefGoogle Scholar
Rigby, S. 1991. Feeding strategies in graptoloids. Palaeontology 34:797–813.Google Scholar
Rigby, S. 1992. Graptoloid feeding efficiency, rotation and astogeny. Lethaia 25:51–68.CrossRefGoogle ScholarPubMed
Rigby, S. 1993a. Graptolite functional morphology: a discussion and critique. Modern Geology 17:271–287.Google Scholar
Rigby, S. 1993b. Population analysis and orientation studies of graptoloids from the Middle Ordovician Utica Shale, Quebec. Palaeontology 36:267–282.Google Scholar
Rigby, S., and Dilly, P. N. 1993. Growth rates of pterobranchs and the lifespan of graptolites. Paleobiology 19:459–475.CrossRefGoogle Scholar
Rigby, S., and Rickards, R. B. 1989. New evidence for the life habit of graptoloids from physical modelling. Paleobiology 15:402–413.CrossRefGoogle Scholar
Ruedemann, R. 1947. Graptolites of North America. Geological Society of America Memoir 19.CrossRefGoogle Scholar
Sleigh, M. A., and Blake, J. R. 1977. Methods of ciliary propulsion and their size implications Pp. 243–256in Pedley, T. J., ed. Scale effects in animal locomotion. Academic Press, London.Google Scholar
Stebbing, A. R. D., and Dilly, P. N. 1972. Some observations on living Rhabdopleura compacta (Hemichordata). Journal of the Marine Biological Association of the United Kingdom 52:443–448.CrossRefGoogle Scholar
Strickler, J. R. 1982. Calanoid copepods, feeding currents and the role of gravity. Science 218:158–160.CrossRefGoogle ScholarPubMed
Strickler, J. R. 1985. Gravity, drag and feeding currents of small zooplankton—discussion. Science 228:1017.CrossRefGoogle Scholar
Sudbury, M. 1991. On the dimension of the graptolite zooid. Geological Magazine 128:381–384.CrossRefGoogle Scholar
Underwood, C. J. 1993. The position of graptolites within Lower Paleozoic planktic ecosystems. Lethaia 26:189–202.CrossRefGoogle Scholar
Urbanek, A. 1987. The phylogenetic debate on the affinities of graptolites. An attempt at methodological analysis. Bulletin of the Geological Society of Denmark 35:223–230.CrossRefGoogle Scholar
Urbanek, A., and Towe, K. M. 1975. Ultrastructural studies on graptolites. 2. The periderm and its derivatives in the Graptoloidea. Smithsonian Contributions to Paleobiology 22:1–24.CrossRefGoogle Scholar
Vogel, S. 1981. Life in moving fluids. Princeton University Press, N. J.Google Scholar
Vogel, S. 1988. Life's devices. The physical world of animals and plants. Princeton University Press, N. J.Google Scholar