ABSTRACT a Study of the Life History of a Population of Nereis Virensat
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· '.' -\ ABSTRACT A study of the life history of a population of Nereis virensat Brandy Cove, St. Andrews, New Brunswick, indica-6es that these worms may live for 12-15 years, maturing at the earliest in their fourth year. The majority, however, do not mature until their fifth or sixth year. At the onset of maturity 'gonadal' clumps are found either floating free in the coelom or embedded in the parenchymal tissue at the bases of the parapodia. Eggs were observed to arise froIn these 'gonadal' clumps during every month of theyear and took 1-2 years to mature. 'Sperm plates' were only produced froID 'gonadal' clUlnps at the end of July to the beginning of' August and mature sperrn \'lere observed by the follovIing May. The ratio of males: females in the spawning population was found to be 3:1. It was also observed that this population did not undergo extensive epi tol;:al metamorphosis and that the worms "l:;herefore spawned in an atokous .condi tian. These worms did no-t; show an extensive swarming behaviour at the sea surface p in fact only the males were observed swimrning close to the surface of the mud p on the incomming tide, releasing a continuous stream of sperm froIn thelr pygidial papillae. What the females do in the field i8 still uncertain. Larval develop ment followed very closely that described for other nereids an0. was observed to be non-pelagic. · ",' History of Nereis virens at Brandy Cove, St. Andrews, N.B. Doreen Snow ',\ Some Aspects of the Life History of the Nereid Worm. Nereis virens (Sars), on an Inter tidal Mudflat at Brandy Cove, St. Andrews, N.B. by Doreen Rosemary Snow, B.Sc. (Hons.) A thesis submitted in partial fulfilment of the requirements for the degree of Master of Science Department of Biology McGill University lVIarch 1972 @ Doreen Rosemary Snm'l 1972 · '.' Frontispiece 1 - Size range in the N. virens population at Brandy Cove, st Andrews, N.B. a) This worm is about O.5gm. in weight and according to this study, about one year old. b) These worms are about 25-)Ograms. • ~. t '.\ . 1111 ~IIIIIIII li IllIlllllllll,l, Il,llll J 111'l111~111 1111111~11 l Il.1 ~~TlMETE~S ' ., : ,', '.' 1 • 1 ' . 1 1111 1 )':NTlMETERS 2' '~, '. 4. 5,111 III 1 1III · ,.1 Frontispiece 2 - Colour differentiation in N. virens. The upper individual is immature with the typical orange-reddish-green colour, whereas the bottom individual has green eggs about 100-120u in the coelom which make the bases of the parapodia green. ".' ..' '.' · '.' Frontispiece 3 - Pale green 3-segment larvae of N. virens (approximately 330u) with the red pigment spots at the sides of the head. • t~ f , . ',' ",,{. TABLE OF CONTENTS ACKNOWLEDGElV.lENTS. • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • i LIST OF TABLES ••••••• 0 • 0 • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • ii LIST OF FIGURES ••••••••••••••••• ~ ••••••••••••••••••••••••• iii CIASSIFICATION •••••••••••••••••••••••••••••••••••••••••••• vii ll'ITRODUCTI ON •••••••••••••••••••••••••••••••••••••••••••••• 1 lV.lETHODS AND lVlATERIALS ••••••••••••••••••••••••••••••••••••• 6 Samplillg' Area •••••••.••. ., 0 •••••••••••••••••••••••••• 6 Samplillg' technique •••••••••••••••••••••••••••••••••• 6 Maintenance in the laboratory....................... 13 Measurement of worm size ••••• .. 14 Examination of Coelomic Fluid •••••••••••••••••• ..... 15 Histological treatment.............................. 16 Observations on Spawning............................ 16 Artificial Fertilizations and Rearillg' of Larvae..... 16 Collections of Larvae in the Field.................. 17 Photographs •••••.•••.••..••... 0 •••••••••••• QI • • • • • • • • 18' SIZE DISTRIBUTION Growth Pattern••••••• •••••••••••••••••••••• 0 ........ .. 19 Age Cl.asses •••••••••• 0 •••••• 0 • • • • • • • • • • • • • • • • • • • • • • • 22 Age at Maturity..................................... 36 REPRODUCTION Development of the Coelomic Fluid ••••••••••••••••••• 47 "'~' TABLE OF CONTENTS (Cont'd) 1) Small c oe lomocyte S·y •••••••••••••••••••••• 0 • 2) Mature trephocytes and the formation of 'parenchymal' tissue........................ 48 J) Gonadal clumps............................. 49 4) Phagocytic cells and later changes......... 51 Maturation and Structure of the Oocytes............ 52 Maturation and Structure of the Spermatozoan....... 62 Differentiation of the Sexes....................... 70 Sex Ratio.......................................... 71 Spawning 1) Field observations ••••••••••••••••••••••••• 7J 2) Laboratoryobservations •••••••••••••••••••• 74 J) Release of Spermatozoa ••••••••••••••••• ~ ••• 76 4) Release of oocytes ••••••••••••••••••••••••• 79 5) Conditions governing the time of spawning •• 80 URVAL DEVELOPMENT Early Development.................................. 85 Larvae with J-Chaetigerous Segments................ 98 Later Larval Stages................................ 106 GENERAL DISCUSSION... • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • 121 CONCLUSIONS •• 0 • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • 124 REFERANCES CITED APPENDIX · '.' -i- ACKNOWLEDGEMENTS l am deeply indebted to the Fisheries Research Board of Canada, Biological Station, St. Andrews, New Brunswick, who graciously provided facilities, such as laboratory space and equipment, in conjunction with the Huntsman Marine Laboratory; and to aIl members of the Biological Station for their kind co-operation and advice. l am grateful to my supervisor, Dr. Joan Marsden, for her support during this project; to Dr. Dorothy Pocock for her help and use fuI advice, especially at the beginning of this project; to John Patterson who aided in collecting the worms; to A~ Sreeharen for help with the statistics involved in this project; to Bill McMullon and Frank Cunningham of the Biological Station, F.R.B., for their help with the photographie work; to Dr. F. A. Aldrich who offered advice during the writing of this thesis; to my brother Hugh, who stuck on the stacks of photographs; to Miss Cynthia Long who helped edit the thesis and to my father who helped to type the final copy. Finally, l wish to thank the National Research Council of Canada, for their support during this project. ".~' . -ii- LIST OF TABLES TABLE I. Time and duration of each collection period, showing as well the number of worms collected in each •••••..•...•.•••••..•.•...•...••.......•..•. 12 TABLE II The mean weight for each possible age class for each month, as determined from figure 5 •... showing some approximate ages, in terms of years, for worms of a certain weight •••••••••••••••••••• 32 TABLE III. Coelomic fluid categories for N. virens....... 38 TABLE IV. Ratio of males to females from January to May 1969 .•.•..••...•••..••.•.••••..•.••...••.. & • • 72 .-.... -iii- LIST OF FIGURES FIGURE 1. Map o~ St. Croix River and Passamaquoddy Bay showing the positions of Oak Bay, Brandy Cove, Bocabec Bay and St. Andrews Point ••••••••••• 8 FIGURE 2. Map of Brandy Cove, New Brunswick showing the collection site, Area A ••••••••••••••••••••••• 10 FIGURE 3. a) Relationship between segment number and weight in M. virens. b) Relationship between length and weight in N. virens. c) Relationship between width and weight in N. virens •...... I!' ••••••••••••••••••••••••••••••••• 21 FIGURE 4. Weight/frequency diagrams ~or Mereis virens ~rom September 1968 to September 1969 ••••••••••••• 25 FIGURE 5. Graphs o~ the logarithmic dif~erences of the class ~requencies plotted against the Iudpoint o~ the class ~or N. virens from September 1968 to September 1969 (excluding Dec. 1968) ••••••••••• 28-30 FIGURE 6. Weight/frequency distributions for the total sample of N. virens in each coelomic fluid category from September 1968 to September 1969.... 40 FIGURE 7. Coelomic fluid of M. virens with only small coelomocytes, Coelomocytes I ••••••• o............. 42 FIGURE 8. Coelomic fluid of N. virens with both small coelomocytes and large coelomocytes or parenchymal cells, Coelomocytes II.oooooooooooooo 42 FIGURE 9. Coelomic fluid o~ N. virens with small coelomocytes, parenchymal cells and gonadal clumps, Gonadal Clumps ••••••••• o................. 44 FIGURE 10. Coelomic fluid of a spent male, with sperm and phagocytic cells (cells, 15-20~. containing green crystalloid granules)....................... 44 FIGURE 11. Coelomic fluid o~ N. virens with small coelomocytes, parenchymal cells, gonadal clumps and small eggs, Females I ••• o.. o................. 55 -iv- FIGURE 12. Mean oocyte diameters for each female examined over the period of September 1968 to September 1969................................. 57 FIGURE 13. Coelomic fluid of N. virens during the early stages of Females II. Eggs containing numerous oil droplets and measuring 120-14o~............... 60 FIGURE 14. Coelomic fluid of N. virens during the late stage of Females II. Eggs very dense, pale green in colour and measuring 180-220~............ 60 FIGURE 15. Coelomic fluid of male N. virens in November and October, containing 'sperm plates', 50-90p.... 64 FIGURE 16. Coelomic fluid of male N. virens with 'sperm plates' breaking up into secondary spermatocytes which then undergo meiosis to produce spermatids.. 64 FIGURE 17. Coelomic fluid of male N. virens with spermatids breaking up into sperm............................ 67 FIGURE 18. Coelomic fluid of male N. virens in May with mature sperm possessing an acrosome, a nucleus, mitochrondrial spheres and a tail................. 67 FIGURE 19. Average sea water temperatures for Brandy Cove,