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SALMON and SALMONIDAE .I The Salmonidae are an important See also:family of fishes belonging to the Malacopterygian Teleosteans, characterized as follows: Margin of the upper See also:jaw formed by the premaxillaries and the maxillaries—supraoccipital in contact with the frontals, but frequently overlapped by the parietals, which may meet in a sagittal suture; opercular bones all well See also:developed. Ribs sessile, parapophyses very See also:short or absent; epineurals, sometimes also epipleurals, See also:present. See also:Post-temporal forked, the upper See also:branch attached to the epiotic, the See also:lower to the opisthotic; postclavicle, as usual, applied to the inner See also:side of the clavicle. A small adipose dorsal fin. See also:Air-See also:bladder usually present, large. Oviducts rudimentary or absent, the ova falling into the cavity of the See also:abdomen before extrusion. The Salmonidae are very closely related to the Clupeidae, or See also:herring family, from which they are principally distinguished by the position of the postclavicle and by the presence of a rayless fin on the back, at a considerable distance from the true or rayed dorsal fin; this so-called adipose fin is an easy recognition-See also:mark of this family, so far as See also:British See also:waters are concerned, for, if it is present in several other families, these have no representatives in the See also:area occupied by the fresh-See also:water salmonids, with the exception of the See also:North See also:American Siluridae and Percopsidae, which are readily distinguished by the pungent spine or spines which precede the rays of the first dorsal fin. The imperfect See also:condition of the oviducts, quite exceptional among fishes, owing to which the large ripe eggs may be easily squeezed out of the abdomen, is a feature of See also:great See also:practical importance, since it renders artificial impregnation particularly easy, and to it is due the fact that the See also:species of Salmo have always occupied the first See also:place in the See also:annals of See also:fish-culture. The Salmonidae inhabit mostly the temperate and See also:arctic zones of the See also:northern hemisphere, and this is the See also:case with all fresh-water forms, with one exception, Retropinna, a See also:smelt-like fish from the coasts and See also:rivers of New See also:Zealand. A few deep-See also:sea forms (See also:Argentina, Microstoma, Nansenia, Bathylagus) are known from the Arctic ocean, the Mediterranean and the See also:Antarctic ocean, down to 2000 fathoms. The question has been discussed whether the salmonids, so many of which live in the sea, but resort to rivers for breeding purposes, were originally marine or fresh-water. The See also:balance of See also:opinion is in favour of the former See also:hypothesis, which is supported by the fact that the overwhelming See also:majority of the members of the suborder of which the salmonids See also:form See also:part permanently inhabit the sea. The clupeids, ' The Latin name salmo possibly means literally " the leaper," from salire, to leap, jump.for instance, which are their nearest See also:allies, are certainly of marine origin, as proved by their abundance in Cretaceous seas, yet a few, like the shads, ascend-rivers to spawn, in the same way as the salmon does, without this ever having been adduced as See also:evidence in favour of a fresh-water origin of the genus Clupea to which they belong. No remains older than See also:Miocene (Osmerus, Prothymallus, Thaumaturus) are certainly referable to this family, the various Cretaceous forms originally referred to it, such as Osmeroides and Pachyrhizodus, being now placed with the Elopidae. There is probably no other See also:group of fishes to which so much See also:attention has been paid as to the Salmonidae, and the species have been unduly multiplied by some writers. Perhaps not more than 8o should be regarded as valid, but some of them fall into a number of See also:local forms which are distinguished as varieties or subspecies by some authors, whilst others would assign them full specific See also:rank. These See also:differences of opinion prevail whether we See also:deal with Salmo proper or with Coregonus. See also:Classification.—The See also:recent genera may be arranged in five See also:groups: The first, which includes Salmo, Brachymystax, Stenodus, Coregonus, Phylogephyra and Thymallus, has 8 to 20 branchiostegal rays, 9 to 13 rays in the ventral fin, the pyloric appendages more or less numerous (17 to 200) and breeding takes place in fresh water. The second group, with the single genus Argentina, is, like the following, marine, and is characterized by 6 branchiostegal rays, 11 to 14 ventral rays, the See also:stomach caecal, with pyloric appendages in moderate See also:numbers (12 to 20). The third group, genera Osmerus, Thaleichthys, Mallotus, Plecoglossus, Hypomesus, has 6 to to branchiostegal rays, 6 to 8 ventral rays, the stomach caecal, with pyloric appendages few (2 to 11) or rather numerous. The See also:fourth group, genera Microstoma, Nansenia, Bathylagus, deep-sea forms with the branchiostegal rays reduced to 3 or 4, ventral rays 8 to to, the stomach caecal and pyloric appendages absent; whilst the fifth group, with the genera Retropinna and Salanx, is distinguished from the preceding in having no air-bladder, branchiostegal rays 3 to 6, ventral rays 6 or 7, stomach siphonal and pyloric appendages absent. The genus Salmo, the most important from the economical and sporting points of view, is characterized by small smooth scales, which at certain seasons may become embedded in the slimy skin, a moderately high dorsal fin with to to 12 well-developed rays, and a large mouth provided with strong See also:teeth, which are present not only in the jaws and on the See also:palate, but also on the See also:tongue; the maxillary or posterior See also:bone-of the upper jaw extends to below or beyond the See also:eye. See also:Young specimens (see See also:PARR) are marked with dark See also:vertical bars on the sides (parr-marks), which in some -See also:trout are retained throughout See also:life, and have the caudal fin more or less deeply forked or marginate, the form of the fin changing with the See also:age and sexual development of the fish. Adult See also:males have the jaws more produced in front than See also:females, and both snout and See also:chin may become curved and hooked. As pointed out by A. See also:Gunther, who was the first to make a profound study of the members of this genus, and especially of the British forms, there is probably no other group of fishes which offers so many difficulties to the ichthyologist with regard to the distinction of species, as well as to certain points in their life-See also:history, the almost See also:infinite See also:variations which they undergo being dependent on age, See also:sex and sexual development, See also:food and the properties of the water. The difficulties in their study have rather been increased by the excessive multiplication of so-called specific forms. Opinions also vary as to the importance to be attached to the characters which serve to group the See also:principal species into natural divisions. Whilst A. Gunther admitted two genera, Salmo and Oncorhynchus, D. S. See also:Jordan and B. W. Evermann go so far as to recognize five, Oncorhynchus, Salmo, Hucho, Cristivomer and Salvelinus. The latter arrangement is certainly the more logical, the difference between the first genus and the second being of rather less importance than that between the second and the third. However, considering the slightness of the distinctive characters on which these divisions are based, and the See also:complete passage which obtains between them, the writer of this See also:article thinks it best to maintain the genus Salmo in the wide sense, whilst retaining the divisions as subordinate divisions or sub-genera, with the following See also:definitions : Oncorhynchus (Pacific salmon).—Vomer See also:flat, toothed along the See also:shaft, at least in the young; anal fin with 12 to 17 well-developed rays. Salmo (true salmon and trout).—Vomer flat, toothed along the shaft, at least in the young; anal fin with 8 to 12 well-developed rays. Salvelinus (See also:char).—Vomer See also:boat-shaped, the shaft strongly de-pressed behind the See also:head, which alone is toothed, the teeth forming an isolated fascicle; anal fin with 8 to to well-developed rays. Hucho (huchens).—Vomer as in the preceding, but teeth forming a single arched transverse See also:series continuous with the See also:palatine teeth; anal fin with 8 to to well-developed rays. The salmon itself (Salmo salar), the type of the family, is a large fish, attaining a length of 4 or 5 ft., and living partly in the sea, partly in fresh water, breeding in the latter. Fish which thus ascend rivers to spawn are called " anadromous." It may be briefly defined as of silvery coloration, with small See also:black spots usually confined to the side above the lateral See also:line, with the teeth on the shaft of the vomer disappearing in the adult, with 18 to 22 gill-rakers on the first branchial See also:arch, with It or 12 well-developed rays in the dorsal fin, See also:Ito to 125 scales in the lateral line, and II or 12 (exceptionally 13) between the latter and the posterior border of the adipose fin. The young, called "parr" or "samlet," characterized by a smaller mouth, the maxillary bone not extending much beyond the vertical of the centre of the eye, the presence of an alternating See also:double or zigzag series of teeth on the shaft of the vomer, the presence of dark vertical bars on the sides of the See also:body, together with more or less numerous small red spots, is hatched in the See also:spring; and usually remains for about two years in the rivers, descending at the third spring to the sea, where it is known as "smolt." In the sea it soon assumes a more See also:uniform silvery coloration and from this See also:state, or " grilse," develops its sexual See also:organs and re-enters rivers to breed, after which operation, much emaciated and unwholesome as food, it is known as " kelt," and returns to the sea to recuperate. It has now been ascertained by the investigations instituted in See also:Norway by K. See also:Dahl that the smolts, immediately after leaving the rivers, make for the open sea, and do not return to the See also:coast until they have reached the grilse See also:stage. Thus specimens measuring between 8 and 18 in. hardly ever fall into the hands of the See also:angler. The salmon inhabits the North See also:Atlantic and its tributary waters. It is known to extend as far north as Scandinavia, See also:Lapland, See also:Iceland, See also:Greenland and Labrador, and as far See also:south as the north-See also:west of See also:Spain and the state of See also:Connecticut. It ascends the See also:Rhine as far as See also:Basel. There are See also:land-locked forms in Scandinavia and in See also:Canada and See also:Maine, which are regarded by some authors as distinct species (S. hardinii from See also:Lake Wener, S. sebago from Sebago Lake in Maine, S. ouananiche from Lake St See also: It is usually estimated that a female salmon produces about 900 eggs for each See also:pound of her own weight ; but this See also:average is often exceeded.
The See also:time between fertilization and hatching, or the See also:escape of the young fish from the See also:egg-membrane, varies considerably with the temperature to which the eggs are exposed. It has been found that at a See also:constant temperature of 41° F. the period is 97 days; but the period may be as short as 70 days and as See also:long as 15o days without injury to the See also:health of the embryo. It follows therefore that in the natural conditions eggs deposited in the autumn are hatched in the See also:early spring. The newly hatched fish, or " alevin," is provided with a very large yolk-See also:sac, and by the absorption of the yolk is nourished for some time; although its mouth is fully formed and open, it takes no food. The alevin stage lasts for about six See also:weeks, and at the end of it the young fish is about t} in. long.
The grilse, after spawning in autumn, return again to the sea in the See also:winter or following spring, and reascend the rivers as mature spawning salmon in the following year. Both salmon and grilse after spawning are called " kelts.' The following recorded experiment illustrates the growth of grilse into salmon: a grilse-kelt of 2 lb was marked on See also: The Scottish rivers flowing into the See also:German Ocean and Pentland See also:Firth are almost all early, while those of the Atlantic slope are late. The See also:Thurso in See also:Caithness and the Naver in See also:Sutherlandshire contain fresh-run salmon in See also:December and January; the same is the case with the See also:Tay. In See also:Yorkshire salmon commence their ascent in See also:July, August or September if the season is wet, but if it is dry their See also:migration is delayed till the autumn rains set in. In all rivers more salmon ascend immediately after a spate or See also:flood than when the See also:river is See also:low, and more with the flood See also:tide than during the ebb. In their ascent salmon are able to pass obstructions, such as waterfalls and weirs of considerable height, and the leaps they make in surmounting such impediments and the persistence of their efforts are very remarkable.
We reproduce here, with additions, See also:Professor See also:Noel See also:Paton's See also:summary (published first in the loth edition of this Encyclopcedia) of observations on the life-history of the salmon. Important advances in our knowledge of the life-history of the salmon have been made through the investigations of Professor F. Miescher on the Rhine at Basel, of Professor P. P. C. Hoek in See also: In a few months a smolt will increase from a few ounces to 4 or 5 lb; while Archer's weighings of 16 salmon which had been marked and recaptured in the following year showed an average gain of 36 %, reckoned on from kelt stage to kelt stage. During the season of 1895 Tosh, at See also:Berwick-on-See also:Tweed, opened between March and August 514 fish, and found food in the stomachs of 76, or over 14% of the whole. As to the nature of the food, it was found to be as follows: Herring . . 36 or 47% See also:Crustacea, amphipods, &c. . 14 ,, 18% See also:Sand eels . 11 „ 14 % See also:Haddock and See also:whiting . . 8 „ 10% Feathers and See also:vegetable See also:matter . 7 ,, 9 % Excluding the feathers and vegetable matter, which are not really of the nature of food, all the material found in the stomach was of marine origin. Hoek, out of 2000 fish examined by him, found 7 with food in the stomach, and, curiously enough, 4 of these were taken on the same See also:day. In each case marine fish constituted the food. As to where salmon go to feed in the sea, our See also:information is still very deficient, but the prevalence of herring in the stomach would seem to indicate that they must follow the shoals of these fish which approach the coast during the summer months. While there can be no doubt that salmon feed in the sea, the question of whether they feed in fresh water has been much debated. It is difficult for the popular mind to conceive of an active fish like the salmon subsisting for several months without food, and the fact that the fish so frequently not only takes into its mouth but actually swallows See also:worms and various lures has still further tended to confirm many See also:people in the conviction that salmon do feed in fresh water. In discussing the question it is well clearly to understand what is meant by feeding. It is the taking, digesting and absorbing of material of use in the See also:economy in such quantities as to be of benefit to the individual. Accepting this See also:definition, it may at once be said that all the evidence we possess is entirely opposed to the view that salmon feed when in fresh water. Miescher examined the stomachs of about 2000 salmon captured at Basel, about 50o m. from the mouth of the Rhine, and in only two did he find any indication of feeding. These two fish were male kelts. One contained the remains of a cyprinoid fish, and the other had a dilated stomach with an See also:acid secretion, but no food remains. Hoek, who, as already stated, examined about 2000 fish, found food of marine origin in 7, but in none food derived from fresh water. Of the 132 stomachs of salmon from the estuaries and upper waters of Scottish rivers examined in the laboratory of the College of Physicians not one contained any food remains. The stomach of salmon captured in fresh water is collapsed and shrunken. Its mucous membrane is thrown into folds, and it contains a small amount of mucus of a neutral reaction. The See also:intestine, which usually contains numerous tape-worms, is full of a greenish-yellow viscous material which, when examined under the See also:microscope, is found to consist of mucus with See also:shed See also:epithelial and other cells and with masses of crystals of carbonate of See also:lime. In no case does the microscope reveal any food remains such as fish-scales, plates of crustacea or bristles of worms or annelids. In the fish taken in the estuaries up to the month of August the See also:gall-bladder is distended; in those taken later in the year it is empty. In all the fish from the upper waters the gall-bladder is empty and collapsed. According to the investigations of Hoek and of Gulland, the lining membrane of the stomach and intestine degenerates while the fish is in the river, but the correctness of these observations has been denied by F. B. See also: Briefly stated, these investigations show that the supply of albuminous material and fats stored in the muscles and used while the fish is in the river is amply sufficient for the greatest requirements of the fish. The amount of energy liberated from the fats and albuminous material is 570 times more than is required to raise the fish from the level of the See also:estuary to that of the upper waters! These analyses further show that all the materials required for the construction of the ovaries and the testes are found in sufficient quantity in the muscles, with the exception of See also:iron, which is, however, abundantly present in the See also:blood.
It is a very See also:common opinion that kelts feed voraciously while still in fresh water, and this has been used as an See also:argument that they should be destroyed. It is not easy to bring forward such satisfactory evidence as has been adduced in the case of unspawned salmon, since it is illegal to kill kelts; but none of the 25 kelts procured by the Scottish See also:Fishery See also:Board, and examined in the College of Physicians' laboratory, contained any food, and Mr See also: By See also:November they have risen to 20 % in the Tweed and in Holland, and to 23 % in the upper reaches of the Rhine. According to Archer's observations, the development of the ovaries in grilse in the earlier months somewhat lags behind that in the salmon. The growth of the testes has been chiefly investigated by Archer and Tosh in the Tweed and by Miescher at Basel. From March to the middle of July in the Tweed these organs increase from about .19 to .35% of the weight of the fish. In July their rate of growth increases, and they reach their maximum development at the end of September, when they are about 6% of the body weight. In the Rhine in March they weigh about • 1 %, and they reach their maximum development of about 5 % in See also:October. What leads to the migration of salmon from sea to river and river to sea ? It is usually supposed that they come to the river to spawn; that it is the See also:nisus generativus that drives them from the sea, where their ova will not develop, to the fresh water where development is possible. But it is found that salmon are passing from sea to river at all seasons of the year, and with their genitalia in all stages of development—some fish, running in March with ovaries only 1% of the body weight, other fish not running till October with ovaries t5 or 16% of the body weight. It is difficult, then, to accept the theory that the sexual See also:act is the governing See also:factor. That it is a secondary factor seems to be indicated by the great run offish in See also:June, July and August, when the genitalia are most rapidly growing. There is one respect, however, in which all the fish leaving the sea for the river agree, and that is in the amount of stored material accumulated in their bodies. In the early running fish this material is largely confined to the muscles, but in the later coming fish it is more equally distributed between muscles and genitalia. The amount of stored material may be measured by the amount of solids, and if we See also:express the results of all the fish examined in terms of fish of uniform size—too cm. in length— the following results are obtained Nov.' Feb. See also:Mar. See also:April. May July Oct. Kelts. and and and June. Aug. Nov. Muscles 2481 2214 2355 2599 2210 2270 1750 946 Ovaries 23 24 24 33 47 72 545 9 See also:Total 2504 2238 2379 2632 2257 2342 2295 955 It would thus appear that, when the salmon has in the sea accumulated a certain definite amount of nourishment, it ceases to feed, and returns to the river irrespective of the state of its genital organs. See also:Nutrition, and not the nisus generativus, appears to be the See also:motive See also:power. That the fish after` spawning returns to the sea in See also:search of food is fully recognized by all. Course of Migration.—It is well known that while salmon run all the year through in greater or lesser numbers, the run of grilse takes place in the summer months, from May to August. But it is further possible to See also:divide the salmon into classes—the so-called winter salmon of the Rhine, large fish running from October to See also:February, with unripe ovaries and testes; and the summer salmon, running for the most part from March to October, with genitalia more or less ripe. These summer fish are small in the early months, but increase in size as the autumn advances. The winter salmon, along with the early summer or spring fish, appear to pass directly to the upper reaches of the river, and to spawn there, while the larger late-coming fish appear to populate the lower waters. This seems to be indicated by the comparison of upper-water and estuary fish throughout the year. The period at which male and female fish enter the rivers also appears to be somewhat different. The observations of Tosh, Miescher and Hoek show that throughout the year the female fish exceed the males in number, and, secondly, that during the earlier months of the year female fish run in much larger numbers than do male fish. It is only in September that anything like an equality between the two sexes is established. But in Great See also:Britain it is not until the end of August that the nets are removed, and one cannot but believe that the destruction of such a very large proportion of females as are captured during the early months of the season must have a most prejudicial effect upon the breeding stock. Rate of Migration.—By a comparison of the first appearance of winter salmon and of grilse in the markets of Holland and of Basel—50o m. up the river—Miescher gives some data for the determination of the average rate at which salmon ascend an unobstructed stream. It was found that winter salmon appeared at Basel about 54 days after their appearance in Holland, which would give a rate of passage of about to m. per diem. From a smaller number of observations on grilse, it appears that they travel at a somewhat slower rate. It is, however, doubtful how far these figures are of value in deciding the rate at which fish pass up the lower reaches of the river.
Great difficulties have been experienced in ascertaining the age and rate of growth of salmon. The practice has long ago been resorted to of " marking " salmon, the most satisfactory mark being a small oblong See also:silver See also:label, oxidized or blackened, bearing distinctive letters and numbers, to the dorsal fin. But of late the structure of the scales has been studied with the See also:object of obtaining indications of the age, growth and spawning See also:habit. H. W. See also:Johnston in 1905 contributed an interesting See also:paper on the subject. The scales See also:bear concentric lines, which vary in number and relative distance according to the growth of the fish, and during the feeding periods these lines are added with more rapidity and a greater degree of separation than at other times. Johnston has endeavoured to ascertain their meaning in Tay salmon, and he has shown that the number of lines external to their last See also:annual See also:ring gives some See also:clue to the time at which they See also:left the sea; he is thus able to distinguish among ascending salmon such as are on their first return from such as have made the See also:journey once or oftener before.
The group of Pacific salmon, or See also: They See also:die after the breeding season is over, and never return to the sea. For the important Salmonidae known as TROUT, CHAR,See also:WHITEFISH,SMELT,See also:GRAYLING, &C., see the See also:separate articles. The huchen (S. hucho) of the See also:Danube is an elongate, somewhat See also:pike-like form, growing to the same size ' Winter fish not due to spawn till following November. as the salmon, of silvery coloration, with numerous small black dots, extending on the dorsal fin. Allied to it are S. fluviatilis from See also:Siberia and S. perryi or blackistoni from the northern See also:island of See also:Japan. The genus Stenodus is intermediate between Salmo and Coregonus (whitefish). S. leucichthys is an anadromous species, inhabiting the See also:Caspian Sea and ascending the See also:Volga and the Ural; it is also found in the Arctic ocean, ascending the Ob, See also:Lena, &c. It grows to a length of 5 ft. A second species occurs in Arctic North America; this is the " Inconnu," S. mackenzii, from the See also:Mackenzie river and its tributaries. The capelin (Mallotus villosus, so called from the villous bands formed by the scales of mature males) is a salmonid of the coasts of Arctic America and north-eastern Asia; it deposits its eggs in the sand along the shores in incredible numbers, the See also:beach becoming a quivering See also:mass of eggs and sand. Plecoglossus, a salmonid from japan and See also:Formosa, is highly remarkable for its lamellar, See also:comb-like, lateral teeth. The siel-smelts, Argentina, are deep-sea salmonids, of which examples have occasionally been taken off the coasts of Scotland and See also:Ireland. Bathylagus, another salmonid discovered by the " Challenger " expedition, is still better adapted for life at great depths (down to 1700 fathoms), the eyes being of enormous size. AuTHORIIIES.—On the systematic and life histories: A. Gunther, See also:Catalogue of Fishes in the British Museum, vol. vi. (1866) ; F. Day, British and Irish Salmonidae (See also:London, 1887) ; F. A. Smitt, Kritisk Forteckning ofver de i Riksmuseum befintliga Salmonider (See also:Stockholm, 1886) ; V. Fatio, Faune See also:des vertebras de la Suisse, vol. v. (189o); D. S. Jordan and B. W. Evermann, Fishes o North America, vol. i. (1896), and American Food and See also:Game Fishes (London and New See also:York, 1902) ; F. F. Kavraisky, Die Lachse der Kaukasuslander (See also:Tiflis, 1896). On growth and migrations: Die histochemischen and physiologischen Arbeiten von See also:Friedrich Miescher, See also:Band ii., pp. 116, 192, 304, 325 (See also:Leipzig, 1897) ; P. P. C. Hoek, Statische and biologische Untersuchungen an in den Niederlandern gefangenen Lachsen (Chark,ctenburg, 1895) ; Annual Reports of the Fishery Board for Scotland, part ii., " See also:Report on Salmon Fisheries," Nos. TI, 12, 13, 14 (1893—1894–95–96) ; Report of Investigations on the Life-History of the Salmon to the Fishery Board for Scotland, edited by Noel Paton, presented to See also:parliament and published 1898; K. Dahl, Orret og unglahs samt lovgivningens forhold til dem (See also:Christiania, 1902); H. W. Johnston, " The Scales of Tay Salmon as indicative of Age, Growth and Spawning Habit," See also:Ann. See also:Rep. Fish. Board, Scotland, See also:xxiii., appendix ii. (1905). Introduction in See also:Tasmania and New Zealand: M. See also:Allport, Proc. Zool. See also:Soc. (1870), pp. 14 and 750; A. See also:Nichol, See also:Acclimatization of the Salmonidae at the See also:Antipodes (London, 1882) ; W. See also:Arthur, ` History of Fish Culture in New Zealand," Tr. N. Zeal. Inst. xiv. (1881) p. 18o; P. S. Seager, " Concise History of the Acclimatization of the Salmonids in Tasmania," Proc. R. Soc. Tasm. (1888) p. 1; also R. M. Johnston, l.c. p. 27. On the salmon disease: T. H. See also:Huxley, Quart. Jour. Micr. Sci. xxii. (1882) p. 311. (G. A. Additional information and CommentsThere are no comments yet for this article.
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