REPRODUCTIVE and LARVAL ECOLOGY OF MARINE BOTTOM INVERTEBRATES
Tóm tắt
1. In analysing the ecological conditions of an animal population we have above all to focus our attention upon the most sensitive stages within the life cycle of the animal, that is, the period of breeding and larval development.
2. Most animal populations on the sea bottom maintain the qualitatively composition of the species composing them, over long periods of time, though the individual species use quite different modes of reproduction and development. This shows that species producing a large number of eggs have a larger wastage of eggs and larvae than those with only a few eggs. The wastage of eggs in the sea is much larger than on the land and in fresh water.
3. In the invertebrate populations on the level sea bottom, large fluctuations in numbers from year to year indicate species with a long pelagic larval life, while a more or less constant occurrence indicates species with a very short pelagic life or a non‐pelagic development.
4. In most marine invertebrates which shed their eggs and sperm freely in the water, either
5. Embryos with a non‐pelagic development may originate
6. Three types of pelagic larvae are known:
7. To find out the factors which cause the enormous waste of eggs and larvae, we thus have to study those forms (constituting 7o% of all species of bottom invertebrates in Recent seas) which have a long planktotrophic pelagic life, as only species reproducing in this way have really large numbers of eggs.
8. The food requirements of the planktotrophic pelagic larvae are much greater than those of the adult animals at the bottom. The adaptability of the larvae to poor food conditions seems, nevertheless, to be greater than hitherto believed. The significance of starvation seems mainly to be an indirect one: poor food conditions cause slow growth, prolong larval life, and give the enemies a longer interval of time to attack and eat the larvae.
9. At the temperatures to which they are normally exposed, northern as well as tropical larvae seem on an average to spend a similar time (about 3 weeks) in the plankton. The length of the pelagic life of the individual species may, however, vary significantly in nature. In the Sound (Denmark) the larvae are never exposed to temperatures outside the range which they are able to endure. The wastage caused by temperature, like that due to starvation, seems mainly to be an indirect one: low temperatures postpone growth and metamorphosis, and give the enemies a longer time to feed on the larvae.
1o. When a larva feeding on a pure algal diet metamorphoses into a carnivorous bottom stage, a ‘physiological revolution’ occurs and a huge waste of larvae might be expected. Experiments have, however, shown that this is not the case.
11. Young pelagic larvae are photopositive and crowd near the surface; larvae about to metamorphose are photonegative. Larval polychaetes, echinoderms, and presumably also prosobranchs, may prolong their pelagic life for days or weeks until they find a suitable substratum. Forced towards the bottom by their photonegativity and transported by currents over wide bottom areas, testing the substratum at intervals, their chance of finding a suitable place for settling is much better than hitherto believed.
12. Continuous currents from the continental shelf towards the open ocean may transport larvae from the coast to the deep sea where they will perish. Such conditions may (for instance in the Gulf of Guinea) deeply influence the composition of the fauna, while in other areas (European western coast, southern California) they seem to be only of small significance.
13. The toll levied by enemies appears to be the most essential source of waste among the larvae. A list of such enemies, comprising other pelagic larvae, holoplank‐tonic animals and bottom animals, is given on p. 2o. A medium‐sized
14. Species reproducing in a vegetative way, by fission, laceration, budding, etc., might be expected to have good chances of competition in such areas where conditions for sexual reproduction are unfavourable. Nevertheless, they only supply a rather small percentage of the animal populations of all Recent seas, probably because their intensity of reproduction is low and because they are unable to spread to new areas. Most forms reproducing in a vegetative way have sexual reproduction as well.
15. Pelagic development is nearly or totally suspended in the deep sea, and is restricted to the shelf faunas. In the arctic and antarctic seas pelagic development is nearly or totally suppressed, even in the shelf faunas, but starting from here the percentage of forms with pelagic larvae gradually increases as we pass into warmer water, reaching its summit on the tropic shelves.
16. In order to survive in high arctic areas a planktotrophic, pelagic larva has to complete its development from hatching to metamorphosis within I–I ½ months (i.e. the period during which phytoplankton production takes place) at a temperature below 2–4o C. Most larvae, that is in 95% of the species, are unable to do so and have a non‐pelagic development, but if a pelagic larva is able to develop under these severe conditions the planktotrophic pelagic life seems to afford good opportunities even in the Arctic. Thus the 5 % of arctic invertebrates reproducing in this way comprise several of the species which quantitatively are most common within the area.
17. The antarctic shore fauna has poor conditions similar to those of the Arctic. The longest continuous periods of phytoplankton production are 2 and 3 weeks respectively, and pelagic larvae have, in order to survive, to complete their development within this short space of time at a temperature between 1 and 4o C. Accordingly, non‐pelagic development is the rule, but most arctic species are able to support their non‐pelagic development by means of much smaller eggs than the antarctic species, where brood protection and viviparity is dominant. The antarctic fauna has apparently had a longer time to develop its tendency to abandon a pelagic life. The greater the size of the individual born, the smaller its relative food requirements and the better its chance of competing under poor food conditions.
18. The relatively few data on reproduction in deep sea invertebrates point to a non‐pelagic development. The larvae of such forms, in order to develop through a planktotrophic pelagic stage, would have to rise by the aid of their own locomotory organs through a water column 2000–4000 m. high or more (often with counteracting currents) to the food producing surface layer, and to cover the same distance when descending to metamorphose and settle.
19. The ecological features common to the deep sea, the arctic and the antarctic seas, which enable the same animals to live and to reproduce there, contribute to explain the ‘equatorial submergence’ of many arctic and antarctic coastal forms.
20. In the tropical coastal zones where the percentage of species with pelagic larvae reaches its maximum, the production of food for the larvae takes place much more continuously than in temperate and arctic seas, because light conditions enable the phytoplankton to assimilate all the year round. The tropical species of marine invertebrates breed (in contrast to temperate and arctic species) within such different seasons that their larval stock, taken as a whole, is more or less equally distributed in the plankton all the year round. This makes the competition in the plankton less keen.
21. The fact that a mode of reproduction and development, well fit for an arctic area, is unfit in a temperate or tropical area of the sea is probably one of the main reasons for the restricted distribution of species.
22. In most groups of marine invertebrates the individual species have only one mode of reproduction and development, which accordingly restricts their area of distribution. In the polychaetes, however, the individual species often show an astonishing lability in their mode of reproduction and development which enables them to compete in wide areas of the sea. Thus, out of the Western European species of polychaetes, 28‐4% have been found also in the Indian Ocean, and 18%, at least, along the Californian coast, while the corresponding number of Western European echinoderms, prosobranchs and lamellibranchs found also in the Indian Ocean and California amounts to less than 2%.
23. The pelagic or non‐pelagic development of marine prosobranchs has proved to be a very fine ‘barometer’ for ecological conditions. Recent observations, still not elaborated, seem to indicate that the shape of the top whorls, the apex, of the adult shells of prosobranchs may show whether the species in question has a pelagic or a non‐pelagic development. This discovery may also give us valuable information about the larval development in fossil species, and help us to form an idea about ecological conditions in sea areas from earlier geological periods.
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Tài liệu tham khảo
Aiyar R. G., 1933, Preliminary observations on some polychaete larvae of the Madras coast and a note on the occurrence in tow‐net water of the larvae of Chaetogordius Moore, J. Madras Univ., 5, 115
Ankel W. E., 1936, Prosobranchia, Tierwelt N.- u. Ostsee, 9, 1
Aziz N. D., 1938, Polychaetes. In ‘Fauna of Karachi’, Mem. Dep. Zool. Panjab Univ., 1, 19
Balss H., 1927, Crustacea Decapoda, Kükenthals Handb. Zool. Berl., 3, 840
Batchelder C. H., 1915, Migration of Ilyanassa obsoleta, Litorina litorea, and Lit. rudis, Nautilus, 29, 43
Belding D. L., 1930, The soft‐shelled clam fishery of Massachusetts (including the natural history of the soft‐shelled clam, etc.), Mar. Fish. Ser., 65
Bernard F., 1898, Recherch. es ontogéniques et morphologiques sur la coquille des Lamellibranches, Ann. Sci. nat. Sér. 8, 8, 1
Carlgren O., 1901, Die Brutpflege der Actiniarien, Biol. Zbl., 21, 468
Carlgren O., 1924, Actiniaria from New Zealand and its subantarctic islands, Vidensk. Medd. naturh. Foren. Kbh., 77, 179
Carter G. S., 1931, The fertilisable life of the eggs of Echinus esculentus and Ech. miliaris, J. exp. Biol., 8, 194
Caullery M., 1916, Viviparité et parthénogenèse chez les Annélides Polychètes: un nouveau Syllidien vivipare (Ehlersia nepiotoca n. sp.) dans les mares a Lithothamnion de la Hague (au C Saint Martin), C. R. Acad. Sci., Paris, 163, 576
Cerruti A., 1942, Osservazioni ed esperimenti sulle cause di distruzione delle larve d'ostrica nel Mar Piccolo e nel Mar Grande di Taranto, Arch. Oceanogr. Limnol., 1, 163
Chun C., 1897, Die Beziehungen zwischen dem arktischen und antarktischen Plankton, 64
Claparede ED., 1869, Beiträge zur Kenntnis der Entwicklungsgeschichte der Chaetopoden, Z. wiss. Zool., 29, 163
Clubb J. A., 1902, Rep.‘Southern Cross’Antarctic Exp.
Coe W. R., 1901, Papers from the Harriman Alaska expedition, XX. The Nemerteans, Proc. Wash. Acad. Set., 3, 1
Coe W. R., 1943, Biology of the Nemerteans of the Atlantic coast of North America, Trans. Conn. Acad. Arts Sci., 35, 129
Cole H. A., 1939, Further experiments in the breeding of oysters in tanks, Fish. Invest. Ser. II, 16, 1
Dall W. H., 1924, The value of the nuclear characters in the classification of marine gastropods, J. Wash. Acad. Sci., 14, 177
Dautzenberg PH., 1912, Pectinidés, Siboga Exped., 53, 1
Dautzenberg PH., 1925, Les mollusques marins du Finistère et en particulier de la région de Roscoff, Trav. Stat. biol. Roscoff, 3, 1
Davis F. M., 1923, Quantitative studies on the fauna of the sea bottom. No. 1. Preliminary investigation of the Dogger Bank, Fish. Invest. Ser. II, 6, 1
Dawydoff C., 1940, Quelques Véligères géantes de Prosobranches provenant de la Mer de Chine, Bull. biol., 74, 497
Day J. H., 1937, The development of Capitellides giardi Mesnil, Rep. Dove mar. Lab. (1936), ser- 3, 4, 31
Dehorne A., 1925, Observations sur la biologie de Nereis diversicolor, C.R. Acad. Sci., Paris, 180, 1441
Dehorne A., 1927, Le cycle reproducteur annuel de Dodecaceria concharum au Portel. La Schizo metamérie, C.R. Acad. Sci., Paris, 184, 547
Drasche C., 1884, Beitrage zur Entmchlung der Polychäten. I. Entmcklung von Pomatoceras triqueter, 10
Duncan F. M., 1946, Wonders of migration, 150
Ekman S., 1925, Holothurien, Further zool. Res. Swed. Antarct. Exp.
Ekman S., 1935, Tiergeographie des Meeres, xii + 542
Eliot C. N. E., 1904, On some nudibranchs from East Africa and Zanzibar. Pt. 5, Proc. zool. Soc. Lond., 83, 10.1111/j.1469-7998.1904.tb08314.x
Elsey C. R., 1936, Artificial stimulation of spawning in the Pacific oyster, Progr. Rep. Pacific Biol. Stat. Nanaimo B.C., 29, 10
Fauvel P., 1911, Annélides polychètes du Gulfe persique recueillis par M. N. Bagoyawlensky, Arch. Zool. exp. gen., Sér., 5, 6
Fauvel P., 1916, Annélides Polychètes des lies Falkland recueillies par M. Rupert Vallentin Esq. (1902–1910), Arch. Zool. exp. gin., 55, 417
Fauvel P., 1923, Polychètes errantes, Faune Fr., 5, 1
Fauvel P., 1927, Polychètes sedentaires, Faune Fr., 16, 1
Fauvel P., 1932, Annelida polychaeta of the Indian museum, Calcutta, Mem. Indian Mus., 12, 1
Fauvel P., 1939, Annelides Polychetes de l'lndochine, Commentat. Pontif. Acad. Sci., 3, 243
Fell H. B., 1945, A revision of the current theory of Echinoderm embryology, Trans. R. Soc. N.Z., 75, 73
Fisher W. K., 1940, Asteroidea, Discovery Rep., 20, 69
Fox, 1937, The activity of poikilothermal animals in different latitudes. I, Proc. zool. Soc. Lond., 945
Franc A., 1943, Études sur le développement de quelques Prosobranches méditerranéens, Thèse, Faculté Sci. Alger, 158
Galtsoff P. S., 1932, Spawning reactions of three species of oysters, J. Wash. Acad. Sci., 22, 65
Galtsoff P. S., 1939, Natural history and method of controlling the starfish (Asterias forbesi Desor), Bull. U.S. Bur. Fish., 49, 73
Gårder F., 1932, Hydrographisch‐biochemische Untersuchungen in norwegischen Austern‐Pollen, Bergens Mus. Aarb., 1, 1
Gerould S. H., 1906, The development of Phascolosoma. (Studies on the embryology of the Sipunculidae, II), Zool. Jrb. (Abt. 2), 23, 77
Giard A., 1904, Rep. 6. Congr. international de Zool
Grave B. H., 1932, La ponte et l'incubation chez les Annélides Polychètes, Ann. Sci. nat., 6
Gravter CH., 1923, La ponte et l'incubation chez les Annelides Polychetes, Ann. Sci. Nat., 6
Guberlet J. E., 1934, Observations on the spawning and development of some Pacific annelids, Proc. Fifth Pacific Sci. Congr., 5, 4213
Gutsell J. S., 1930, Natural history of the bay‐scallop (Pecten irradians), Bull. U.S. Bur. Fish., 46, 569
Hagmeier A., 1930, Abderhalden's Handb. biol. Arbeitsmethoden, Abt. 9, Teil 5, 465
Hart T. J., 1934, On the phytoplankton of the south‐west Atlantic and the Bellinghausen sea, 1929‐31, Discovery Rep., 8, 1
Heath H., 1905, The breeding habits of Chitons of the Californian coast, Zool. Anz., 29, 390
Hempelmann FR., 1911, Zur Naturgeschichte von Nereis dumerilii Aud. & Edw, Zoologica, Stuttgart, 62, 1
Hentschel E., 1936, Allgemeine Biologie des Südatlantischen Ozeans, Wiss. Ergebn. dtsch. atlant. Exp. ‘Meteor’ 1925–7, 1
Herpin R., 1923, Un essaimage en plein jour d'une Annélide polychète, Pionosyllis lamelligera, C.R. Acad. Sci., Paris, 177, 355
Herpin R., 1924, Essaimage et développement d'un Eunicien et d'un Syllidien, C.R. Acad. Sci., Paris, 179, 1431
Herpin R., 1925, Recherches biologiques sur la reproduction et le développement de quelques Annélides Polychètes, Bull. Soc. Sci. Nat. Ouest, sér. 4, 5, 1
Hesse R., 1924, Tiergeographie auf ökologischer Grundlage, 613
Hornell J., 1894, On the method of dispersion and fertilization of ova in some Sabellids, J. Mar. Zool., 1, 13
Hörstadius SV., 1925, Entwicklungsmechanische Studien an Holothuria Poli D. Ch., Ark. Zool., 17, 1
HÖRSTADIUS SV., 1926, Ueber die Entwicklung von Astropecten aurantiacus L, Ark. Zool., 18, 1
Izuka A., 1912, The errantiate Polychaeta of Japan, J. Coll. Sci. Tokyo, 30, 1
JÄGERSTEN G., 1940, Die Abhängigkeit der Metamorphose vom Substrat des Biotops bei Protodrilus, Ark. Zool., 32, 1
Jeffreys GW., 1863, British Conchology, 2, 1863
Johnson M. W., 1939, The correlation of water movements and dispersal of pelagic larval stages of certain littoral animals, especially the sand crab, Emerita, J. Mar. Res., 236
Jorgensen C.Barker(1946).Lamellibranch larvae. See Thorson (1946).
Kirk H. B., 1909, Two species of Actiniaria from Campbell Island, The subantarctic islands of New Zealand, 1, 384
Knudsen J., 1949, Egg‐capsules and development of some marine prosobranchs from tropical West‐Africa, Atlantide-Report, 1
Koehler R., 1904, Ophiures de l'Expédition du Siboga, ie Partie, Ophiures de Mer Profonde, Siboga Exped, 45, 1
Koehler R., 1905, Ophiures de l'Expédition du Siboga, 2e Partie, Ophiures littorales, Siboga Exped., 45, 1
Koehler R., 1910, Asteries du Musée de Calcutta. II. Les Asteries littorales. Echinoderms of the Indian Museum, Publ. Trustees Ind. Mus., Calcutta, 6, 192
Koehler R., 1921, Echinodermes, Faune Fr., 1, 1
Koehler R., 1924, Les Echinodermes des mers d'Europe, 1, 1
Koehler R., 1910, Echinoderma of the Indian Museum, 54
Korringa P., 1940, Experiments and observations on swarming pelagic life, and setting in the European flat oyster, Ostrea edulis L, Arch, neerl. Zool., 5, 1
Lamy ED., 1923, Revision des Saxicavidae vivants du museum national d'histoire naturelle de Paris, J. Conchy Hot., 68, 218
Lamy ED., 1928, La ponte chez les gastéropodes prosobranches, J. Conchyliol. ser. 4, 72, 25
Lang K., 1939, Ober die Entwicklung von Priapulus caudatus Lam, K. fysiogr. Sällsk. Lund Forh., 9, 1
Lemche H., 1948, Northern and arctic Tectibranch Gastropods. I. The larval shells. II. A revision of the cephalaspid species, K. danske vidensk. Selsk. Skr. (Biol.), 5, 1
Loeb J., 1901, On the prolongation of the life of unfertilised eggs of the sea‐urchin by potassium cyanide, Amer. J. Physiol., 6, 305, 10.1152/ajplegacy.1902.6.5.305
Ludwig H., 1904, Brutpflege bei Echinodermen, Zool. J.
Lynge H., 1909, Marine Larnellibranchiata. In The Danish Expedition to Siam. IV, K. danske vidensk. Selsk. Skr. ser. 7, 5, 100
McBride E. W., 1920, Echinoderma (Part II) and Enteropneusta. Larvae of Echinoderma and Enteropneusta, Rep. Brit. Antarct. Exp. 1910, Zoology, 4, 83
McBride E. W., 1908, Echinoderma. II. Echinoderm‐larvae, Nation. Antarct. Exp. 1901-4, Nat. Hist., 4, 9
McIntosh N. A., 1934, Distribution of the Macroplankton in the Atlantic sector of the Antarctic, Discovery Rep., 9, 69
Mann G.(1948).Biologia de la Antarctica Suramericana. Inst. Geogr. Univ. Chile Rep. no. 517 364pp. Chile.
Marshall SH. M., 1933, The breeding of reef animals. Part I. The corals, Great Barrier Reef Exp. 1928–29, Sci. Rep., 3, 219
Medem F., 1942, Beiträge zur Frage der Befruchtungsstoffe bei marinen Mollusken, Biol. Zbl., 62, 431
Meijere J. C. H., 1904, Die Echinoidea der Siboga‐Expedition, Siboga Exped., 43, 1
Melvill J. C., 1901, The mollusca of the Persian Gulf, Gulf of Oman, and Arabian sea, as evidenced mainly through the collections of F. W. Townsend, 1893–1906. Parts I and II, Proc. Zool. Soc. Land., 1901, 327
Mesnil F., 1898, Sur la viviparité d'une Annélide Polychète (Dodecaceria concharum Oerst. forme A), C.R. Soc. Biol., Paris, 5, 905
Mesnil F., 1898, Formes épitoques et polymorphisme évolutif chez une Annélide du groupe des Cirratuliens (Dodecaceria concharum), C.R. Soc. Biol., Paris, 5, 620
Mesnil F., 1917, Un nouveau type de dimorphisme évolutif chez une Annélide Polychète (Spio martinensis), C.R. Acad. Sci., Paris, 165, 646
Mesnil F., 1939, Polychètes sédentaires de l'Expédtion du Siboga, Maldanidae, Cirratulidae, Capitellidae, Sabellidae et Serpulidae, Siboga Exped., 133, 1
Metcalf M. M., 1893, Contribution to the embryology of Chiton, Stud. Biol. Lab. Johns Hopkins Univ., 5, 249
Mohr J. C., 1941, A note on two species of Malaysian King Crabs (Xiphosura), Treubia, 18, 201
Monro C. C. A., 1937, Polychaeta. The John Murray Exp. 1933‐4, Sci. Rep., 4, 243
Mortensen TH., 1910, Echinoidea, Swedish South Polar Exp., 6, 1
Mortensen TH., 1913, Echinodermenlarven. Deutsche Südpolar‐Exped., Zoologie, 6, 67
Mortensen TH., 1920, Notes on the development and the larval forms of some Scandinavian Echinoderms, Vidensk. Medd. naturh. Foren. Kbh., 71, 133
Mortensen TH., 1920, Studies in the development of Crinoids, Pap. Tortugas Lab., 16, 1
Mortensen TH., 1931, Contributions to the study of the development and larval forms of Echinoderms. I–II, K. danske vidensk. Selsk. Skr., Naturv. Math. Afd., Raekke 9, 4, 1
Mortensen TH., 1937, Contributions to the study of the development and larval forms of Echinoderms. III, K. danske vidensk. Selsk. Skr., Naturv. Math. Afd., Raekke 9, 7, 1
Mortensen TH., 1938, Contribution to the study of the development and larval forms of Echinoderms. IV, K. danske vidensk. Selsk. Skr., Naturv. Math. Afd., Raekke 9, 7, 1
Mortensen TH., 1940, Echinoderms from the Iranian Gulf. Asteroidea, Ophiuroidea and Echinoidea.‐Crinoidea by T. Gislén, Danish Sci. Invest, in Iran, 2, 55
Murayama, 1935, On the development of the Japanese Abalone, Haliotis gigantea, J. Coll. Agric. Tokyo, 13, 227
Murray J., 1895, The general conditions of existence and distribution of marine organisms, C.R. Séances je Congr. Intern, Zool., 99
Murray J., 1898, The scientific advantages of an Antarctic Expedition, Proc. roy. Soc. B, 57, 424
Nelson J.(1908).Experimental studies on oyster propagation. Rep. Biol. Dep. N.Y. Agric. Coll. Exp. Sta. for 1907.
Nelson TH. C.(1928b). Rep. Biol. Dep. N.Y. Agric. Coll. Exp. Sta. for 1927.
Nielsen, 1937, Medd. Komm. Danm. Fiskeriog Havunders., ser. Plankton, 3, 37
Nobre A., 1930, Echinodermes de Portugal, 176, 14
Nobre A., 1932, Moluscos Marinhos de Portugal, 1
Nordgaard O., 1912, Faunistiske og biologiske Iakttagelser ved den Biologiske Station i Bergen, K. norske vidensk. Selsk. Skr., 1911, 1
Nyholm K. G., 1943, Zur Entwicklung und Entwicklungsbiologie der Ceriantharien und Aktinien, Zool. Bidr. Uppsala, 22, 85
Okuda S., 1938, Notes on the spawning habits of Arenicola claparedii Levinsen, Annot. zool. jap., 17, 577
Olsen H., 1942, The development of the brittle‐star Ophiopholis aculeata (O. Fr. Müller), with a short report on the outer hyaline layer, Bergens Mus. Aarb., 1
Onoda K., 1936, Notes on the development of some Japanese Echinoids with special reference to the structure of the larval body, Jap. J. Zool., 6, 637
Panikkar N. K., 1939, Observations on breeding in brackish water animals off Madras, Proc. Indian Acad. Set. Ser. B., 9, 343, 10.1007/BF03048507
Panning A., 1937, Über die Wanderungen der Wollhandkrabbe. Markierungsversuche, Mitt. Hamburg. Zool. Mus., 47, 32
Peach C. W., 1858, A few remarks on Lamellaria tentaculata and its nidification etc., as observed at Wick, Proc. R. phys. Soc. Edinb., 1, 455
Pelseneer P., 1901, Sur le degré d'eurythermie de certaines larves marines, Bull. Acad. Belg. bl. Sci., 279
Pelseneer P., 1902, Sur l'exagération du dimorphisme sexuel chez un Gastéropode marin, J. Conchyliol., 50, 41
Pelseneer P., 1903, Mollusques, Prés. Voy. Belgica, 85
Percival E., 1944, A contribution to the life history of the Brachiopod, Terebratella inconspicua Sowerby, Trans. R. Soc. N.Z., 74, 1
1931, Plymouth marine fauna, 371
Prashad B., 1932, Pelecypoda of the Siboga Expedition, Siboga Exped., 53, 1
Risbec J., 1935, Biologie et ponte de mollusques gastéropodes Néo‐Calédoniens, Bull. Soc. Zool. Fr., 60, 387
Robert A., 1902, Recherches sur le développement des Troques, Arch. zool. exp. gén. 3 sér., 10, 269
Runnstrom, 1924, Leptosynapta inhaerens, en Holothurie med forkortad utveckling, Bergens Mus. Aarb., 1
Runnström SVEN, 1927, Über die Thermopathie der Fortpflanzung und Entwicklung mariner Tiere in Beziehung zu ihrer geographischen Verbreitung, Bergens Mus. Aarb. Naturv. Raekke, 1
Runnström, 1927, Über die Entwicklung von Leptosynapta inhaerens (O. F. Müller), Bergens Mus. Aarb., Naturv. Rekke, 1
Runnström, 1919, Über die Entwicklung von Cucumaria frondosa Gunnerus und Psoitis phantapus Strussenf, Bergens Mus. Aarb. Naturv. Raekke, 1
Sars G. O., 1912, On the genera Cryptocheles and Bythocaris G. O. Sars, Arch. Math. Naturv., 32, 1
Sauvage H. E., 1873, Catalogue des Nudibranches des côtes du Boulonnais dressé d'après les notes de Bouchard‐Chanteroux, J. Conchyliol., 21, 27
Schepman M. N., 1908, The Prosobranchia of the Siboga Expedition, pts. 1–6, Siboga Exped., 49
Schmidt G. A., 1932, Dimorphisme embryonnaire de Lineusgesserensis‐ruber et ses relations avec les formes adultes, Ann. Inst. Oceanogr., 12, 65
Schmidt G. A., 1934, Ein zweiter Entwicklungstypus von Lineus gesserensis‐ruber O. F. Müller (Nemertini), Zool. Jrb. (Abt. 2), 58, 607
Seeliger O., 1892, Studien zur Entwicklungsgeschichte der Crinoiden (Antedon rosacea), Zool. Jrb. (Abt. 2), 6, 161
Selenka E., 1876, Zur Entwicklung der Holothurien (Holothuria tubulosa und Cucumaria doliolum), Z. wiss. Zool., 27, 155
Semper K., 1885, Animal life, Intern. Sci. Ser
Shepard F. P., 1941, Submarine topography off the California coast; canyons and tectonic interpretations, Geol. Soc. Amer., Spec. Paper, 31, 171
Simroth H., 1911, Gastropodenlaicheund Gastropodenlarven, Deutsche Tiefsee-Expedition, 1898–9, 9, 377
Sluiter C. PH., 1901, Die Holothurien der Siboga‐Expedition, Siboga Expedit., 44, 142
Soderström A., 1920, Studien über die Polychaetenfamilie Spionidae, 286
SPÅRCK R., 1926, Physiol. Papers dedic. to Prof. Aug. Krogh, 268
Spårck R., 1936, On the relation between metabolism and temperature in some marine lamelli branchs and its ecological and zoogeographical importance, K. danske vidensk. Selsk., Biol. Medd., 13, 1
Stephenson, 1934, The breeding of Reef Animals, part II, Invertebrates other than Corals, Great Barrier Reef Exp. 1928-9, Sci. Rep., 3, 247
Stephenson T. A., 1928, The British sea‐anemones, 1
Stephenson T. A., 1935, The British sea‐anemones 2, 426
Stott F. C., 1931, The spawning of Echinus esculentus and some changes in gonad composition, J. exp. Biol., 8, 133, 10.1242/jeb.8.2.133
Takatsdki, 1929, The heart pulsation of oysters in tropical seas compared with that of those living in seas of the temperate zone, Rec. oceanogr. Whs. Jap., 1, 102
Thorson G., 1935, Studies on the egg‐capsules and development of Arctic marine Prosobranchs, Medd. Grenland, 100, 1
Thorson G., 1936, The larval development, growth and metabolism of Arctic marine bottom invertebrates, etc, Medd. Grenland, 100, 1
Thorson G., 1940, Notes on the egg‐capsules of some North‐Atlantic Prosobranchs of the genus Troschelia, Chrysodomus, Volutopsis, Sipho and Trophon, Vidensk. Medd. naturh. Foren. Kbh., 104, 251
Thorson G., 1940, Studies on the egg‐masses and larval development of Gastropoda from the Iranian Gulf, Danish Sci. Invest. Iran, 2, 159
Thorson G., 1946, Reproduction and larval development of Danish marine bottom invertebrates, Medd. Komm. Damn. Fisk.- og Havunders., ser. Plankton, 4, 523
Vaney C., 1925, L'incubation chez les Holothuries, Trav. Stat. zool. Wimereux, 8, 254
Wesenberg‐Lund Elise, 1949, Polychaeta, Danish Sci. Invest. Iran, 4, 247
Wiborg K.R., 1946, Unders0kelser over oskjellet, Rep. Norw. Fish. Mar. Invest., 8, 1
Williamson H. C.(1900).Contributions to the life‐history of the Edible Crab (Cancer pagurus). 18th Ann. Rep. Fish. Bd. Scotland for 1899 pp.77–143.
Wilson B. CH., 1900, The habits and early development of Cerebratulus lacteus, Quart. J. micr. Sci. ser. 2, 43, 97
Wingfield C. A., 1939, The activity and metabolism of poikilothermal animals in different latitudes. IV, Proc. zool. Soc. Lond., A, 109, 103, 10.1111/j.1096-3642.1939.tb00051.x
Yatsu N., 1902, On the development of Lingula anatina, J. Coll. Sci. Tokyo, 17, 1
Zeuthen E., 1947, Body size and metabolic rate in the animal kingdom, C.R. Lab. Carlsberg, sér. chim., 26, 20