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Normal Stages of the Early Development of the Flying Fish, <I BULLETIN OF MARINE SCIENCE OF THE GULF AND CARIBBEAN VOLUME 11 1961 NUMBER 4 NORMAL STAGES OF THE EARLY DEVELOPMENT OF THE FLYING FISH, HIRUNDICHTHYS AFFINIS (GUNTHER) JOHN W. EVANS Bel/airs Research Institute of McGill. University, St. James, Barbados, W.I. ABSTRACT The embryology and early larval stages of the flying fish Hirundichthys affinis (Gunther) are described. Eggs and larvae show developmental features of a demersal marine teleost and are only secondarily pelagic. INTRODUCTION The flying fish, H irundichthys aDinis (GUnther), is one of the main sources of animal protein available to the population of the island of Barbados. There is no detailed account of the embryology of this species of the Exocoetidae available. The only papers concerned with the embryology of this group are a study by Miller (1952) which deals only with development up to the closure of the blastopore imd a des- cription by Nayudu (1923) of the embryo of a Cypselurus from the Indian Ocean at a stage equivalent to the 36-hour embryo of H. aDinis. References to the larvae are very scattered and are discussed to some extent by Hubbs and Kampa (1946) and Breder (1938). The only figures of young Hirundichthys aDinis were found in Breder ( 1938) in which young fish measuring 27 mm, 37 mm, 44 mm, 67 mm, 85 mm and 137 mm are discussed. The eggs of the Exocoetidae have attracted some attention. They are surprisingly variable in size and strUGture. Breder (1938) has stressed the taxonomic importance of a study of these eggs and Hubbs and Kampa (1943) have prepared a key for the identification of the eggs of flying fish and other Synentognathidae. The sargassum-weed fish "nest" is discussed by Gudger (1937). This nest was originally thought to have been made by an anten- nariid that lives in the weed, but is now known to be produced by several species of flying fish, including H. affinis. The following report deals with the development of the flying fish 484 Bulletin of Marine Science of the Gulf and Caribbean [11(4) from the freshly fertilized egg to the young fish about 10 days old and about 15 mm long. The author is indebted to Dr. N. J. Berrill of the Department of Zoology, McGill University, for encouragement and critical advice; to Dr. J. B. Lewis of the Bellairs Research Institute, Barbados, for advice and help in field work and to Dr. Anton F. Brunn, Copenhagen, for checking the identification of the parent fish. The work was sup- ported by a Colonial Development and Welfare Organization grant in aid of research. METHODS The methods of catching flying fish have been covered in works by Hall (1955) and Hornell (1923). In this study all fish were caught on a nylon line baited with either crab or fresh flying fish. The freshly caught fish were stripped by squeezing the abdomen with a downward motion. If the fish were in running condition, the gametes flow.edfreely from the vent. When a ripe male and female were caught at the same time, the gametes of each were added to a bottle of sea water and fertilization ensued. The parent fish were labelled and fixed in forma- lin. These fish were sent to Dr. Anton Brunn who very kindly checked the author's identification of the species. It was found, as Hall (1955) has noted before, that males were in running condition earlier than females. Females with nearly mature eggs would release upon stripping, but these nonviable eggs could easily be distinguished from mature eggs because they did not form the clump typical of the latter. This is due to the fact that the filaments which usually hold the mature eggs together in a mass were still tightly wrapped around the immature egg, making it look like a small ball of twine and preventing it from attaching to other eggs. Four sets of eggs were obtained on three different dates. Two clusters of eggs were collected on May 20th, 1958, fertilized im- mediately, and from this culture, eggs and embryos were fixed every six hours, starting with the six-hour stage, and preserved for later study. On June 9th and July 12, 1958, two other clusters of eggs were collected, fertilized and used for live observations, rough drawings and photographs. The eggs were incubated in 4 x 4-inch glass cylinders open at the upper end and sealed at the bottom by a piece of plankton silk held in place with a rubber band. The containers were kept three-quarters submerged in a table of running sea water. With this handling the 1961] Evans: Development of Flying Fish 485 eggs seemed to develop to hatching quite normally with a mortality rate of ten to fifteen per cent. The temperature at which the eggs developed in the laboratory (26° C plus or minus 0.5°) differed slightly from the temperature at which they normally develop in the sea (28 ° to 29° C). The difference is probably due to the fact that under normal conditions the eggs develop at the surface of the sea which is almost always brilliantly sun- lit during the day, whereas in the laboratory the water and eggs were always shaded. The somewhat lower temperature probably increased the time required for hat.ching and may possibly have resulted in less pigmentation than is usual under natural conditions. Considerable difficulty was encountered in raising the larval fish and this was probably due to improper feeding. As the yolk sac dis- appears at about 24 hours after hatching, active feeding must begin very soon. On the first two occasions the newly hatched larvae were kept in sea water to which was added a suspension of sea urchin eggs. All the young fish died within four days. On the third attempt a large number of brine shrimp larvae were kept with the young fish from the time of hatching. On this occasion growth was rapid and one fish was kept alive for thirteen days. The eggs and larvae were preserved in five fixatives; 5 per cent saline formalin, Stockard's solution, Stockard's solution plus 0.5 per cent Tergitol, Bouin's fixative and Bouin's solution plus 0:5 per cent Tergitol. The Tergitol was very kindly supplied by the Carbide and Carbon Chemicals Company and as it is a strong detergent it was hoped that it would increase the permeability of the zona radiata to fixatives. However, no difference was noted in the fixing properties of either pair of solutions. Formalin produced much distortion of both egg and embryo and was used only in the preparation of specimens for somite counts. Stockard's solution resulted in the most life-like preservation of whole mount material. The yolk remained crystal clear and the embryo be- came an opaque white. However, it caused the yolk to swell slightly so that the embryo became squeezed between the zona radiata and the expanding yolk and was eventually pushed down into the yolk mass. As a histological fixative, Stockard's solution was unsatisfac- tory. Bouin's solution was a very satisfactory fixative histologically but gave poor results on whole mounts. It caused the yolk mass to shrink 486 Bulletin of Marine Science of the Gulf and Caribbean [11(4) to about two-thirds its normal volume which resulted in a secondary distortion of the embryo. However whole embryos fixed in Bouin's solution were useful for observing certain features such as the degree of pectoral fin development and the extent to which the tail had lifted off the yolk. Whole mounts were drawn with the aid of a camera lucida and photographs of living eggs were used to correct for the distortions caused by fixation. Composite drawings of the whole embryo were usually made from eggs fixed in Stockard's solution. Little success was met with in at- tempting to remove the embryo from the yolk and flatten it under a a cover glass as the tissue was too delicate and brittle. Instead, four or five drawings were made with a camera lucida of different regions of the embryo on the yolk. These drawings were then pieced together to form a composite. In the study of larval and post-larval stages a careful search was made for cartilage. The metachromatic stain, Azure A, was used for this purpose on specimens preserved in Stockard's solution. The young fish were dissected and the parts thought to contain cartilage were stained. Transverse and longitudinal serial sections were made of Bouin- fixed embryos at the ages of 36 hours, 48 hours, hatching, and 7 days after hatching. Sections were cut in paraffin at 7p.. Difficulty was en- countered in removing the embryo from the yolk with the heart and digestive tract intact. Consequently in the first three stages these structures could not be adequately studied or described. Sections were stained in fast green and phloxine. DESCRIPTION OF THE NORMAL DEVELOPMENT OF THE EGGS OF Hirundichthys affinis The egg of H. affinis can be identified to species using the key provided by Hubbs and Kampa (1946). The mature egg, enclosed in its zona radiata, measures 1.6 mm in diameter plus or minus 0.1 mm. The yolk is a clear, light yellow fluid with no oil globules present in the early stages. The zona radiata is a perfectly transparent, very tough, elastic membrane. As in most species of flying fish and their relatives the zona radiata bears a number of filaments. There are two clumps of filaments restricted to opposite poles of the egg. At one pole the filaments are short and fine and resemble a piece of wool stuck to the egg.
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