All living crinoids appear to be passive suspension feeders; they do not generate their own filtration current, but rely on extrinsic water movement to bring food particles to them. Until the 1970’s, understanding of their ecology was both sketchy and fanciful. Crinoids were long thought to spread their arms in an upturned bowl, subsisting on a slow rain of detritus from above (Hyman 1955, Nichols 1962). Fell (1966) reported that they entangled prey with rhythmic scooping motions of their arms. However, SCUBA, ROV and submersible observations have largely disproved both views (e.g., Meyer 1973, Macurda & Meyer, 1974) and have also shown that crinoid feeding is not completely passive. Crinoids are active participants to the extent that they modify arm and pinnule postures (and mobile comatulids and isocrinids seek preferred feeding stations) to take best advantage of prevailing and changing flow patterns and velocities (Meyer 1982a, Meyer et al. 1984a, Baumiller 1997). Although crinoids range from rheophilic "current-lovers" to species that prefer weak flow environments (rheophobic), even abyssal forms appear to depend on horizontal water movements for food rather than a rain of detrital particles.

Many ecological studies of living crinoids have been descriptions of feeding postures and strategies, because the food-gathering apparatus of arms and pinnules comprises such a large proportion of the animal's structure. However, in the comatulids and stalked isocrinids, which account for almost 90% of living species, arms and pinnules also function in locomotion, a dual obligation that requires a skeleton rigid enough to stand erect against a passing current, yet flexible enough to permit movement (Lawrence 1987, Messing & Dearborn 1990). The muscular articulations of arms and pinnules fit this requirement as follows: crinoid ligaments consist of mutable collagenous tissue (or catch connective tissue), the uniquely echinodermal material capable of altering between flaccid and stiffened states (Motokawa 1985, Wilkie & Emson 1988). Contraction of muscles on the ambulacral side of the fulcral ridge curls or rolls an arm inward toward the mouth and flexes pinnules toward the arm axis. When the muscles relax, the elasticity of the large antagonistic ligament on the aboral side of the ridge extends arms and pinnules outward. Once extended, stiffened ligaments allow the arm and pinnules to maintain an extended posture passively against a current for food gathering. Individual articulations have limited scope but an arm of over 200 segments or a pinnule with more than 50 may have great flexibility (see also Baumiller 1997).

Comatulids are the most mobile of extant crinoids and are active arm crawlers, in many cases creeping from cryptic daytime retreats to exposed nocturnal feeding perches (Meyer et al. 1984a, Lawrence 1987, Vail 1987). Also, at least some colobometrids, antedonids, atelecrinids and thalassometrids can swim with graceful and coordinated arm undulations although they appear to do so only rarely (Macurda 1973, Macurda & Meyer 1983, Shaw & Fontaine 1990). Such mobility likely contributed to the shallow-water survival of comatulids in the face of the late-Mesozoic radiation of durophagous predators that drove stalked crinoids into deeper water (Meyer & Macurda 1977, Meyer 1985, Schneider 1988, Oji 1996) (see "Predation" below).

The basic feeding mechanism is well known although few species have been closely examined and several significant details remain to be worked out. Much of the following is taken from Meyer’s (1982a) and Lawrence’s (1987) thorough discussions of crinoid feeding plus information published subsequently. Baumiller (1997) provides a thorough review of feeding and filtration from a biomechanical standpoint.

Fine fingerlike podia (or tube feet), the terminal branches of the water vascular system, occur in groups of three (triads) along both sides of the pinnular ambulacra. Each triad consists of a long, medium and short (or primary, secondary and tertiary) podion. In the photo at left, only the primary podia are visible, as rows of fine short threads along the pinnules. A similar arrangement across five families examined so far suggests that the arrangement is common to all comatulids, and probably to all living crinoids (Nichols 1960, Meyer 1979, Byrne & Fontaine 1981 LaHaye & Jangoux 1985, Holland et al. 1986). The primary podia, 0.43-0.85 mm in length (Meyer 1979, Byrne & Fontaine 1981), alternate with flap-like lappets along the ambulacral margin and, when extended, project almost at right angles outward from the central groove. The bases of secondary podia are fused to the inner surface of the lappets; their contraction pulls the lappets inward, covering the groove. In Antedon bifida and Florometra serratissima, secondary podia curl outward at an angle over the lappets (Lawrence 1987), while in Oligometra serripinna, they project upward from the groove (Holland et al. 1986). The short tertiaries extend vertically from the groove margins. Like ophiuroids, but unlike most other extant echinoderms, crinoid tube feet lack a terminal sucker. They bear papillae tipped with cilia (ostensibly sensory) and containing mucus-producing cells (Holland 1969, McKenzie 1992).

When a suspended food particle comes in contact with a primary podion, the podion flicks, bends or curls rapidly inward, forcing the particle into the food groove. The shorter podia and lappets vary somewhat in function among species. In A. bifida and F. serratissima, the shorter podia (and, in F. serratissima, the lappets as well) scrape particles off the primaries and retain them in the groove; in A. bifida, secondary podia can also capture food (Nichols 1960, Byrne & Fontaine 1981, LaHaye & Jangoux 1985, Lawrence 1987). By contrast, the primary podia of O. serripinna perform all "conspicuous small-scale feeding acts" unassisted by secondary podia (Holland et al., 1986). Byrne & Fontaine (1981), Holland et al. (1986) and Leonard (1989) also describe the coordinated activity of multiple adjacent podia associated with capture of larger or mobile particles.

The primary podia at least are adhesive, but the role of mucus in food capture apparently varies among species. Several authors (Magnus 1963, Rutman & Fishelson 1969, Nichols 1960) describe food capture via entanglement in mucous threads and strands. Nichols (1960) describes the podia of A. bifida as forcibly ejecting mucous strands when contacted by a food particle (not a preformed mucous web), but La Touche contends that mucous threads "apparently do not occur in A. bifida" (personal communication in Byrne & Fontaine 1981, p. 17). F. serratissima produces mucous threads, but does not shoot them out upon particle contact. Although Byrne & Fontaine (1981) suggest that food collection via threads could be an important resource, perhaps during dense plankton blooms, they consider that direct impingement of particles on adhesive primary podia is the typical collection method, a conclusion also reached by Meyer (1982a) and Holland et al. (1986) for all comatulids. The latter authors found no mucous threads in O. serripinna.

Wiping of podia against each other and against the current generated by ciliary tracts on the groove floor wraps captured particles in mucous secretions and forms them into boluses which are then transported mouthward by the cilia (Nichols 1960, Byrne & Fontaine 1981, LaHaye & Jangoux 1985, Lawrence 1987). Holland et al. (1986) discuss three possible mechanisms of particle transport via ciliary action. Pinnule grooves run into arm grooves which converge like tributary rivers on the mouth.

The photo at right shows the oral surface, or disk, of a typical comatulid with ambulacral food grooves (AFGs) converging on the central mouth (M) and with the anal papilla (AP) off to one side.

[Modified from Messing (1997)]

References

Baumiller, T.K. 1997. Crinoid functional morphology. Pp. 45-68. IN: Waters, J. A. & Maples, C. G. (eds.) Geobiology of Echinoderms. Paleontological Society Papers 3.

Byrne, M. & Fontaine, A.R. 1981. The feeding behavior of Florometra serratissima (Echinodermata: Crinoidea). Canadian Journal of Zoology 59(1):11-18.

Fell, H.B. 1966. Ecology of crinoids. Pp. 49-62. IN: Boolootian, R. A. (ed.) Physiology of Echinodermata. Wiley-Interscience, NY.

Holland, N.D. 1969. An electron microscope study of the papillae of crinoid tube feet. Pubblicazione Stazione Zoologica di Napoli 37:575-580.

Holland, N.D., Strickler, J.R. & Leonard, A.B. 1986. Particle interception, transport and rejection by the feather star Oligometra serripinna (Echinodermata: Crinoidea), studied by frame analysis of videotapes. Marine Biology 93:111-126.

Hyman, L.H. 1955. The Invertebrates, vol. 4: Echinodermata. McGraw-Hill, New York. vii + 763 p.

Lahaye, M.C. & Jangoux, M. 1985. Functional morphology of the podia and ambulacral grooves of the comatulid crinoid Antedon bifida (Echinodermata). Marine Biology 86:307-318.

Lawrence, J. 1987. A Functional Biology of Echinoderms. Johns Hopkins Press, Baltimore. 340 p.

Leonard, A.B. 1989. Functional response in Antedon mediterranea (Lamarck) (Echinodermata: Crinoidea): the interaction of prey concentration and current velocity on a passive suspension-feeder. Journal of Experimental Marine Biology and Ecology 127:81-103.

Macurda, D.B., Jr. 1973. Ecology of comatulid crinoids at Grand Bahama Island. Hydro-Lab Journal 2:9-24.

Macurda, D.B., Jr. & Meyer, D.L. 1974. Feeding posture of modern stalked crinoids. Nature 247(5440):394-396.

Macurda, D.B., Jr. & Meyer, D.L. 1983. Sea lilies and feather stars. American Scientist 71:354-365.

Magnus, D.B.E. 1963. Der federstern Heterometra savignyi im Roten Meer. Natur Museum, Frankfurt 93:355-368.

McKenzie, J.D. 1992. Comparative morphology of crinoid tube feet. Pp. 73-79. IN: Scalera-Liaci, L. & Canicatti, C. (eds.) Echinoderm Research 1991. Balkema, Rotterdam.

Messing, C.G. 1997. Living Comatulids. Pp. 3-30 IN: Waters, J.A. & Maples, C.G. (eds.) Geobiology of Echinoderms. Paleontological Society Papers 3.

Messing, C.G. & Dearborn, J.H. 1990. Marine Flora and Fauna of the Northeastern United States, Echinodermata: Crinoidea. NOAA Technical Report NMFS 91.

Meyer, D.L. 1973. Feeding behavior and ecology of shallow-water unstalked crinoids (Echinodermata) in the Caribbean Sea. Marine Biology 22(2):105-129.

Meyer, D. L. 1979. Length and spacing of the tube feet in crinoids (Echinodermata) and their role in suspension-feeding. Marine Biology 51:361-369.

Meyer, D.L. 1982. Food and feeding mechanisms: Crinozoa. Pp. 25-42. IN: Jangoux, M. and Lawrence, J. M. (eds.) Echinoderm Nutrition. Balkema, Rotterdam.

Meyer, D.L. 1985. Evolutionary implications of predation on Recent comatulid crinoids from the Great Barrier Reef. Paleobiology 11(2):154-164.

Meyer, D.L. and Macurda, D.B., Jr. 1977. Adaptive radiation of the comatulid crinoids. Paleobiology 3:74-82.

Meyer, D.L. & Macurda, D.B., Jr. 1980. Ecology and distribution of shallow-water crinoids of Palau and Guam. Micronesica 16(1):59-99.

Meyer, D.L., LaHaye, C.A., Holland, N.D., Arneson, A.C. & Strickler, J.R. 1984a. Time-lapse cinematography of feather stars (Echinodermata: Crinoidea) on the Great Barrier Reef, Australia: demonstrations of posture changes, locomotion, spawning and possible predation by fish. Marine Biology 78:179-184.

Motokawa, T. 1985. Catch connective tissue: the connective tissue with adjustable mechanical properties, pp. 69-73. In Keegan, B.F. and O’Connor, B. D. S. (eds.) Echinodermata. Proceedings of the fifth International Conference, Galway. Balkema, Rotterdam.

Nichols, D. 1960. The histology and activities of the tube feet of Antedon bifida. Quarterly Journal of Microscopial Science 101:105-117.

Nichols, D. 1962. Echinoderms. Hutchinson University Library, London 192 p.

Oji, T. 1996. Is predation intensity reduced with increasing depth? Evidence from the west Atlantic stalked crinoid Endoxocrinus parrae (Gervais) and implications for the Mesozoic marine revolution. Paleobiology 22(3):339-351.

Rutman, J. & Fishelson, L. 1969. Food composition and feeding behavior of shallow-water crinoids at Eilat (Red Sea). Marine Biology 3:46-57.

Schneider, J.A. 1988. Frequency of arm regeneration of comatulid crinoids in relation to life habit. Pp. 531-538. IN: Burke, R. D., Mladenov, P. V., Lambert, P. and Parsley, R. L. (eds.) Echinoderm Biology. Balkema, Rotterdam.

Shaw, G.D. & Fontaine, A.R. 1990. The locomotion of the comatulid Florometra serratissima (Echinodermata: Crinoidea) and its adaptive significance. Canadian Journal of Zoology 68:942-950.

Vail, L. 1987. Diel patterns of emergence of crinoids (Echinodermata) from within a reef at Lizard Island, Great Barrier Reef, Australia. Marine Biology 93:551-560.

Willkie, I.C. and Emson, R.H. 1988. Mutable collagenous tissues and their significance for echinoderm palaeontology and phylogeny. Pp. 311-330. IN: Paul, C. R. C. and Smith, A. B. (eds.) Echinoderm phylogeny and evolutionary biology. Clarendon Press, Oxford.