Aspects of Feeding Behaviour of West Indian Reef Corals
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ASPECTS OF FEEDING BEHAVIOUR OF WEST INDIAN REEF CORALS William Stephen Price Thesis submitted in partial fulfillment of the requirements for the Degree of Master of Science, March 1973. ® William Stephen Price 1973 ABSTRACT The feeding behaviour of 35 species of West Indian hermatypic corals was observed underwater, on the reefs off Bellairs Institute, Barbados, West Indies. Observations were supplemented by laboratory study. Food varied from large zooplankton to fine suspended particulate material, which was secured and ingested by tentacular activity and mucus secretions. Long 'sweeper tentacles', up to 80 mm in length, were observed in the polyps of several species. Ciliary currents did not reverse in any of the species and did not aid directly in feeding. Extracoelenteric activity of mesenterial filaments is not considered normal behaviour. William Stephen Price Marine Sciences Centre McGill University Résumé Le comportement nutritif de 35 espèces de coraux hermatypiques des Antille fut observé sous l'eau, sur les récifs au large de l'Institut Bellairs, à la Barbade, aux Petites Antilles. Les observations furent complétées par des études en laboratoire. La nourriture est saisie et absorbée grâce à l'activité des tentacules et à sécrétions mugueuses. Sa grosseur varie entre le zooplancton de grande taille et la matiére particulaire en suspension. Chez quelques espèces, de longues "tentacules balayeuses" atteignant jusqu'à 80 mIn de longueur furent observées dans les polypes. Les courants ciliares n'ont pas été inversés chez aucune des espèces et n'ont pas aidé directement à la nutrition. L'activité extracoelentérique des filaments mésentériques n'est pas considérée comme un comportement normal. Stephen William Price Marine Sciences Centre NcGill University i TABLE OF CONTENTS Introduction Page l Materia1s and Methods 10 Resu1ts 13 Group l 15 Group II 53 Group III 76 Group IV 103 Discussion 110 Sunnnary 117 Literature Cited 120 ii ACKNOWLEDGEMENTS l would like to express my appreciation to those people who have made this project possible: Dr. John Wells, Cornell University, Ithaca, New York, who identified several coral specimens; Mt. Bruce Ott and Mt. Alex Urban who took part in many hours of diving with me; and Dr. John Lewis, to whom l wish to express special thanks, as my supervisor who supplied both academic and financial assistance, and suggested the project. Hi LIST OF TABLES Table 1. Systematic list of Barbados hermatypic coral species examined. Page 8 Table 2. Division of the 35 species of hermatypic corals into four feeding groups. 13 iv LIST OF FIGURES Figure 1. Map of Barbados showing location of reefs and study sites. Page vii 2. Porites porites - fully expanded 19 3. P. porites - ciliary currents 20 4. P. porites - tentacular feeding 20 5. 1. astreoides - fully expanded 22 6. P. astreoides - mucus sheath 22 7. Madracis mirabilis - fully expanded 27 8. M. mirabilis - ciliary currents 28 9. M. mirabilis - tentacular feeding 28 10. M. mirabilis - 'sweeper tentacle' 29 11. M. mirabilis - ingestion of organic particles 29 12. Eusmilia fastigiata - fully expanded 31 13. E. fastigiata - branching tentacles 32 14. E. fastigiata - ciliary currents 32 15. Montastrea cavernosa - fully expanded 35 16. M. cavernosa - 'sweeper tentacles' 36 17. M. cavernosa - tentacular feeding 36 18. M. cavernosa - mucus strand feeding 37 19. M. cavernosa - feeding with polyps contracted 37 20. Mussa angulosa - fully expanded 39 21. M. angulosa - ciliary currents 40 22. M. angulosa - tentacular feeding 40 23. Isophyllia multiflora - fully expanded 42 24. I. multiflora - tentacular feeding 42 25. I. multiflora - ingestion of organic particles 43 26. Dichocoenia stokesi - fully expanded 45 27. ~. stokesi - partially expanded [,.5 28. Favia fragum - fully expanded 48 29. K. fragum - partially expanded 49 30. F. fragum - ciliary currents 50 31. F. fragum - ejection of mucus bolus 50 v Figure 32. Stephanocoenia michelini - fully expanded Page 52 33. Colpophyllia sp. - fully expanded 59 34. Colpophyllia sp. - ciliary currents on periphery of colony 60 35. Colpophyllia sp. - ciliary currents 60 36. Colpophyllia sp. - excretion of fecal bolus 61 37. Colpophyllia sp. - mucus strand feeding 61 38. Colpr~hyllia sp. - 'zipper action' of tentacles 62 39. Colpophyllia sp. - 'zipper action', sequence of movements 63 40. Colpophyllia sp. - capture of small zooplankton 64 41. Colpophyllia sp. - baffle or 'snow fence' effect 64 42. Diploria clivosa - fully expanded 66 43. D. strigosa - fully expanded 68 44. D. labyrinthiformis - fully expanded 70 45. D. labyrinthiformis - 'zipper action' 71 46. MYcetophyllia lamarckiana - expanded soft tissue 73 47. M. lamarckiana - solitary tentacle structure 74 48. M. lamarckiana - tentacular feeding 74 49. M. lamarckiana - juvenile form 75 50. Siderastrea siderea - tentacular arrangement 79 51. S. siderea - collection detritus 80 52. S. siderea - mucus strand feeding 80 53. S. siderea - tentacu1ar feeding 81 54. ~. siderea - excretion of fecal pellet 81 55. S. radians - ciliary currents and tentacular arrangement 83 56. S. radians - ingestion of organic particles 83 57. Agaricia agaricites - fully expanded 85 58. A. agaricites - 'sweeper tentacles' 86 59. A. agaricites - mucus strand feeding 86 60. A. lamarcki - fully expanded soft tissue 88 61. Helioseris cuculata - portion of colony 90 62. ~ cuculata - ciliary currents 91 vi Figure 63. wycetophy11ia danaana - expanded co1ony with mucus strings Page 93 64. ~etophy11ia sp. - fu11y expanded 93 65. M. ferox - fu11y expanded 95 66. Montastrea annu1aris - fu11y expanded 98 67. M. annu1aris - po1yps in different postures 98 68. Acropora pa1mata - expanded apical polyps 100 69. A. pa1mata - expanded po1yps showing tentacu1ar arrangement 100 70. A. cervicornis - fu11y expanded 102 71. Dendrogyra cy1indrus - fu11y expanded 105 72. Meandrina lliE:!mdrites, f. meandrites, f. danae - fu11y expanded 108 73. M. meandrites - partia11y expanded 108 74. M. meandrites , f. danae - expanded soft tissue 109 75. M. meandrites - mucus string feeding 109 vii N Bellairs Site 1. ---~,~. Site 2.---f Bath -----Site 3, Bridgetown 5 miles Figure 1. Map of Barbados showing location of reefs and study sites . ... Active1y growing reef. 1 INTRODUC TION Unti1 recent1y reef cora1s were regarded as being primari1y carnivores feeding on zoop1ankton (Yonge, 1940). However, single ce11ed a1gae, the zooxanthe11ae present in the endoderm of a11 reef building cora1s (Yonge, 1963), are a1so being considered as an important source of nutrition (Franzisket, 1969; Johannes, Co1es and Kuenze1, 1970). It is genera11y assumed that the zooxanthe11ae and the coral form a symbiotic re1ationship. The princip1e photosynthetic product re1eased by the symbiotic a1gae after isolation from hosts is a soluble carbohydrate. "It is 1ike1y that glycero1 is the carbohydrate form trans10cated in associations invo1ving zooxanthe11ae, since the rate of 1iberation of glycero1 'in vitro' is stimu1ated by the presence of some compone~~ of the host tissue and in the intact associations, such as cora1s, the animal tissues are rich in 1abe11ed glycero1 derivatives, such as lipid • • ." (Smith et al., 1969, page 27). In genera1, the detai1s and the biochemica1 significance of the coe1enterate-zooxanthe11ae association is uncertain. There have been, however, assumptions concerning the genera1 metabo1ic characteristics. Geddes (1882) suggested that the uptake of carbon dioxide and the production of oxygen during the photosynthesis wou1d faci1itate the anima1's respiration and gas exchange. The assimilation by the a1gae of inorganic waste products of the host was postu1ated by Yonge and Nicho11s (1931). The 1ink has a1so been considered to encompass the incorp9ration of photosynthetic materia1 into the organic matrix of the animal (Muscatine and 2 Rand, 1958). The presence of zooxanthellae in the corals has been demonstrated to increase the rate of calcification (Goreau, 1959; Goreau and Goreau, 1959). Yonge (1963) postulated that the algae remove excretory waste such as ammonia and phosphates. It was concluded by Smith ~ al. (1965, page 33) that "thesE: interactions . undoubtedly take place to some extent in some associations but as yet there is little direct evidence that any of them are essential to the host." Recent speculations about other possible food sources for corals concerns suspended organic matter, organic aggregates, and dissolved substances circulating in the boundary water of the reef (Goreau et al., 1971). Marshall (1965) found that there was much fine organic material in the water not collected by fine plankton nets. Large amounts of suspended calcareous material are formed by coral-browsing acanthurid and scarid fish (Bardach, 1961). Surge currents and wave turbulence also stir up the leptopel from the bottom sediment, making it available to the benthonic fauna. Baylor and Sutcliffe (1963) have shown that colloidal and dissolved organic mattet' may aggregate into large particles on the surface of bubbles stirred up by the surge. They have suggested that this is a possible source of nutrition for intertidal filter feeders and copepods. Several small organic molecules of biological significance, such as glucose and amine acids, have also been found to be removed from dilute solutions by the coral Fungia scutaria (Stephens, 1962). Bacteria function primarily as nutrient regeneration agents in the sea, because they possess the ability to convert dissolved organic matter into cell substances and to chemo-synthesize protoplasm