A Guide to the Study of Fishes, Volume 2 (of 2)
CHAPTER IV
SALMONIDÆ
=THE Salmon Family.=—The series or suborder _Salmonoidea_, or allies of the salmon and trout, are characterized as a whole by the presence of the adipose fin, a structure also retained in Characins and catfishes, which have no evident affinity with the trout, and in the lantern-fishes, lizard-fishes, and trout-perches, in which the affinity is very remote. Probably these groups all have a common descent from some primitive fish having an adipose fin, or at least a fleshy fold on the back.
Of all the families of fishes, the one most interesting from almost every point of view is that of the _Salmonidæ_, the salmon family. As now restricted, it is not one of the largest families, as it comprises less than a hundred species; but in beauty, activity, gaminess, quality as food, and even in size of individuals, different members of the group stand easily with the first among fishes. The following are the chief external characteristics which are common to the members of the family:
Body oblong or moderately elongate, covered with cycloid, in scales of varying size. Head naked. Mouth terminal or somewhat inferior, varying considerably among the different species, those having the mouth largest usually having also the strongest teeth. Maxillary provided with a supplemental bone, and forming the lateral margin of the upper jaw. Pseudobranchiæ present. Gill-rakers varying with the species. Opercula complete. No barbels. Dorsal fin of moderate length, placed near the middle of the length of the body. Adipose fin well developed. Caudal fin forked. Anal fin moderate or rather long. Ventral fins nearly median in position. Pectoral fins inserted low. Lateral line present. Outline of belly rounded. Vertebræ in large number, usually about sixty.
The stomach in all the _Salmonidæ_ is siphonal, and at the pylorus are many (15 to 200) comparatively large pyloric cœca. The air-bladder is large. The eggs are usually much larger than in fishes generally, and the ovaries are without special duct, the ova falling into the cavity of the abdomen before exclusion. The large size of the eggs, their lack of adhesiveness, and the readiness with which they may be impregnated, render the _Salmonidæ_ peculiarly adapted for artificial culture.
The _Salmonidæ_ are peculiar to the north temperate and Arctic regions, and within this range they are almost equally abundant wherever suitable waters occur. Some of the species, especially the larger ones, are marine and anadromous, living and growing in the sea, and ascending fresh waters to spawn. Still others live in running brooks, entering lakes or the sea when occasion serves, but not habitually doing so. Still others are lake fishes, approaching the shore or entering brooks in the spawning season, at other times retiring to waters of considerable depth. Some of them are active, voracious, and gamy, while others are comparatively defenseless and will not take the hook. They are divisible into ten easily recognized genera: _Coregonus_, _Argyrosomus_, _Brachymystax_, _Stenodus_, _Oncorhynchus_, _Salmo_, _Hucho_, _Cristivomer_, _Salvelinus_, and _Plecoglossus_.
Fragments of fossil trout, very imperfectly known, are recorded chiefly from Pleistocene deposits of Idaho, under the name of _Rhabdofario lacustris_. We have also received from Dr. John C. Merriam, from ferruginous sands of the same region, several fragments of jaws of salmon, in the hook-nosed condition, with enlarged teeth, showing that the present salmon-runs have been in operation for many thousands of years. Most other fragments hitherto referred to _Salmonidæ_ belong to some other kind of fish.
=Coregonus, the Whitefish.=—The genus _Coregonus_, which includes the various species known in America as lake whitefish, is distinguishable in general by the small size of its mouth, the weakness of its teeth, and the large size of its scales. The teeth, especially, are either reduced to slight asperities, or else are altogether wanting. The species reach a length of one to three feet. With scarcely an exception they inhabit clear lakes, and rarely enter streams except to spawn. In far northern regions they often descend to the sea; but in the latitude of the United States this is never possible for them, as they are unable to endure warm or impure water. They seldom take the hook, and rarely feed on other fishes. Numerous local varieties characterize the lakes of Scandinavia, Scotland, and Arctic Asia and America. Largest and most desirable of all these as a food-fish is the common whitefish of the Great Lakes (_Coregonus clupeiformis_), with its allies or variants in the Mackenzie and Yukon.
The species of _Coregonus_ differ from each other in the form and size of the mouth, in the form of the body, and in the development of the gill-rakers.
_Coregonus oxyrhynchus_—the _Schnäbel_ of Holland, Germany, and Scandinavia—has the mouth very small, the sharp snout projecting far beyond it. No species similar to this is found in America.
The Rocky Mountain whitefish (_Coregonus williamsoni_) has also a small mouth and projecting snout, but the latter is blunter and much shorter than in _C. oxyrhynchus_. This is a small species abounding everywhere in the clear lakes and streams of the Rocky Mountains and the Sierra Nevada, from Colorado to Vancouver Island. It is a handsome fish and excellent as food.
Closely allied to _Coregonus williamsoni_ is the pilot-fish, shad-waiter, roundfish, or Menomonee whitefish (_Coregonus quadrilateralis_). This species is found in the Great Lakes, the Adirondack region, the lakes of New Hampshire, and thence northwestward to the Yukon, abounding in cold deep waters, its range apparently nowhere coinciding with that of _Coregonus williamsoni_.
The common whitefish (_Coregonus clupeiformis_) is the largest in size of the species of _Coregonus_, and is unquestionably the finest as an article of food. It varies considerably in appearance with age and condition, but in general it is proportionately much deeper than any of the other small-mouthed _Coregoni_. The adult fishes develop a considerable fleshy hump at the shoulders, which causes the head, which is very small, to appear disproportionately so. The whitefish spawns in November and December, on rocky shoals in the Great Lakes. Its food was ascertained by Dr. P. R. Hoy to consist chiefly of deep-water crustaceans, with a few mollusks, and larvæ of water insects. "The whitefish," writes Mr. James W. Milner, "has been known since the time of the earliest explorers as preeminently a fine-flavored fish. In fact there are few table-fishes its equal. To be appreciated in its fullest excellence it should be taken fresh from the lake and broiled. Father Marquette, Charlevoix, Sir John Richardson—explorers who for months at a time had to depend upon the whitefish for their staple article of food— bore testimony to the fact that they never lost their relish for it, and deemed it a special excellence that the appetite never became cloyed with it." The range of the whitefish extends from the lakes of New York and New England northward to the Arctic Circle. The "Otsego bass" of Otsego Lake in New York, celebrated by De Witt Clinton, is a local form of the ordinary whitefish.
Allied to the American whitefish, but smaller in size, is the Lavaret, Weissfisch, Adelfisch, or Weissfelchen (_Coregonus lavaretus_), of the mountain lakes of Switzerland, Germany, and Sweden. _Coregonus kennicotti_, the muksun, and _Coregonus nelsoni_, the humpback whitefish, are found in northern Alaska and in the Yukon. Several other related species occur in northern Europe and Siberia.
Another American species is the Sault whitefish, Lake Whiting or Musquaw River whitefish (_Coregonus labradoricus_). Its teeth are stronger, especially on the tongue, than in any of our other species, and its body is slenderer than that of the whitefish. It is found in the upper Great Lakes, in the Adirondack region, in Lake Winnipeseogee, and in the lakes of Maine and New Brunswick. It is said to rise to the fly in the Canadian lakes. This species runs up the St. Mary's River, from Lake Huron to Lake Superior, in July and August. Great numbers are snared or speared by the Indians at this season at the Sault Ste. Marie.
In the breeding season the scales are sometimes thickened or covered with small warts, as in the male _Cyprinidæ_.
=Argyrosomus, the Lake Herring.=—In the genus _Argyrosomus_ the mouth is larger, the premaxillary not set vertical, but extending forward on its lower edge, and the body is more elongate and more evenly elliptical. The species are more active and predaceous than those of _Coregonus_ and are, on the whole, inferior as food.
The smallest and handsomest of the American whitefish is the cisco of Lake Michigan (_Argyrosomus hoyi_). It is a slender fish, rarely exceeding ten inches in length, and its scales have the brilliant silvery luster of the mooneye and the ladyfish.
The lake herring, or cisco (_Argyrosomus artedi_), is, next to the whitefish, the most important of the American species. It is more elongate than the others, and has a comparatively large mouth, with projecting under-jaw. It is correspondingly more voracious, and often takes the hook. During the spawning season of the whitefish the lake herring feeds on the ova of the latter, thereby doing a great amount of mischief. As food this species is fair, but much inferior to the whitefish. Its geographical distribution is essentially the same, but to a greater degree it frequents shoal waters. In the small lakes around Lake Michigan, in Indiana and Wisconsin (Tippecanoe, Geneva, Oconomowoc, etc.), the cisco has long been established; and in these waters its habits have undergone some change, as has also its external appearance. It has been recorded as a distinct species, _Argyrosomus sisco_, and its excellence as a game-fish has been long appreciated by the angler. These lake ciscoes remain for most of the year in the depths of the lake, coming to the surface only in search of certain insects, and to shallow water only in the spawning season. This periodical disappearance of the cisco has led to much foolish discussion as to the probability of their returning by an underground passage to Lake Michigan during the periods of their absence. One author, confounding "cisco" with "siscowet," has assumed that this underground passage leads to Lake Superior, and that the cisco is identical with the fat lake trout which bears the latter name. The name "lake herring" alludes to the superficial resemblance which this species possesses to the marine herring, a fish of quite a different family.
Closely allied to the lake herring is the bluefin of Lake Michigan and of certain lakes in New York (_Argyrosomus nigripinnis_), a fine large species inhabiting deep waters, and recognizable by the blue-black color of its lower fins. In the lakes of central New York are found two other species, the so-called lake smelt (_Argyrosomus osmeriformis_) and the long-jaw (_Argyrosomus_ _prognathus_). _Argyrosomus lucidus_ is abundant in Great Bear Lake. In Alaska and Siberia are still other species of the cisco type (_Argyrosomus laurettæ_, _A. pusillus_, _A. alascanus_); and in Europe very similar species are the Scotch vendace (_Argyrosomus vandesius_) and the Scandinavian Lok-Sild (lake herring), as well as others less perfectly known.
The Tullibee, or "mongrel whitefish" (_Argyrosomus tullibee_), has a deep body, like the shad, with the large mouth of the ciscoes. It is found in the Great Lake region and northward, and very little is known of its habits. A similar species (_Argyrosomus cyprinoides_) is recorded from Siberia—a region which is peculiarly suited for the growth of the _Coregoni_, but in which the species have never received much study.
=Brachymystax and Stenodus, the Inconnus.=—Another little-known form, intermediate between the whitefish and the salmon, is _Brachymystax lenock_, a large fish of the mountain streams of Siberia. Only the skins brought home by Pallas a century ago are yet known. According to Pallas, it sometimes reaches a weight of eighty pounds.
Still another genus, intermediate between the whitefish and the salmon, is _Stenodus_, distinguished by its elongate body, feeble teeth, and projecting lower jaw. The Inconnu, or Mackenzie River salmon, known on the Yukon as "charr" (_Stenodus mackenziei_), belongs to this genus. It reaches a weight of twenty pounds or more, and in the far north is a food-fish of good quality. It runs in the Yukon as far as White Horse Rapids. Not much is recorded of its habits, and few specimens exist in museums. A species of _Stenodus_ called _Stenodus leucichthys_ inhabits the Volga, Obi, Lena, and other northern rivers; but as yet little is definitely known of the species.
=Oncorhynchus, the Quinnat Salmon.=—The genus _Oncorhynchus_ contains the salmon of the Pacific. They are in fact, as well as in name, the king salmon. The genus is closely related to _Salmo_, with which it agrees in general as to the structure of its vomer, and from which it differs in the increased number of anal rays, branchiostegals, pyloric cœca, and gill-rakers. The character most convenient for distinguishing _Oncorhynchus_, young or old, from all the species of _Salmo_, is the number of developed rays in the anal fin. These in _Oncorhynchus_ are thirteen to twenty, in _Salmo_ nine to twelve.
The species of _Oncorhynchus_ have long been known as anadromous salmon, confined to the North Pacific. The species were first made known nearly one hundred and fifty years ago by that most exact of early observers, Steller, who, almost simultaneously with Krascheninnikov, another early investigator, described and distinguished them with perfect accuracy under their Russian vernacular names. These Russian names were, in 1792, adopted by Walbaum as specific names in giving to these animals a scientific nomenclature. Five species of _Oncorhynchus_ are well known on both shores of the North Pacific, besides one other in Japan. These have been greatly misunderstood by early observers on account of the extraordinary changes due to differences in surroundings, in sex, and in age, and in conditions connected with the process of reproduction.
There are five species of salmon (_Oncorhynchus_) in the waters of the North Pacific, all found on both sides, besides one other which is known only from the waters of Japan. These species may be called: (1) the quinnat, or king-salmon, (2) the blue-back salmon, or redfish, (3) the silver salmon, (4) the dog-salmon, (5) the humpback salmon, and (6) the masu; or (1) _Oncorhynchus tschawytscha_, (2) _Oncorhynchus nerka_, (3) _Oncorhynchus milktschitsch_, (4) _Oncorhynchus keta_, (5) _Oncorhynchus gorbuscha_, (6) _Oncorhynchus masou_. All these species save the last are now known to occur in the waters of Kamchatka, as well as in those of Alaska and Oregon. These species, in all their varied conditions, may usually be distinguished by the characters given below. Other differences of form, color, and appearance are absolutely valueless for distinction, unless specimens of the same age, sex, and condition are compared.
The quinnat salmon (_Oncorhynchus tschawytscha_),[7] called quinnat, tyee, chinook, or king-salmon, has an average weight of 22 pounds, but individuals weighing 70 to 100 pounds are occasionally taken. It has about 16 anal rays, 15 to 19 branchiostegals, 23 (9 + 14) gill-rakers on the anterior gill-arch, and 140 to 185 pyloric cœca. The scales are comparatively large, there being from 130 to 155 in a longitudinal series. In the spring the body is silvery, the back, dorsal fin, and caudal fin having more or less of round black spots, and the sides of the head having a peculiar tin-colored metallic luster. In the fall the color is often black or dirty red, and the species can then be distinguished from the dog-salmon by its larger size and by its technical characters. The flesh is rich and salmon-red, becoming suddenly pale as the spawning season draws near.
Footnote 7:
For valuable accounts of the habits of this species the reader is referred to papers by the late Cloudsley Rutter, ichthyologist of the _Albatross_, in the publications of the United States Fish Commission, the _Popular Science Monthly_, and the _Overland Monthly_.
The blue-back salmon (_Oncorhynchus nerka_),[8] also called red salmon, sukkegh, or sockeye, usually weighs from 5 to 8 pounds. It has about 14 developed anal rays, 14 branchiostegals, and 75 to 95 pyloric cœca. The gill-rakers are more numerous than in any other salmon, the number being usually about 39 (16 + 23). The scales are larger, there being 130 to 140 in the lateral line. In the spring the form is plumply rounded, and the color is a clear bright blue above, silvery below, and everywhere immaculate. Young fishes often show a few round black spots, which disappear when they enter the sea. Fall specimens in the lakes are bright crimson in color, the head clear olive-green, and they become in a high degree hook-nosed and slab-sided, and bear little resemblance to the spring run. Young spawning male grilse follow the changes which take place in the adult, although often not more than half a pound in weight. These little fishes often appear in mountain lakes, but whether they are landlocked or have come up from the sea is still unsettled. These dwarf forms, called kokos by the Indians and benimasu in Japan, form the subspecies _Oncorhynchus nerka kennerlyi_. The flesh in this species is firmer than that of any other and very red, of good flavor, though drier and less rich than the king-salmon.
Footnote 8:
For valuable records of the natural history of this species the reader is referred to various papers by Dr. Barton Warren Evermann in the Bulletins of the United States Fish Commission and elsewhere.
The silver salmon, or coho (_Oncorhynchus milktschitsch_, or _kisutch_), reaches a weight of 5 to 8 pounds. It has 13 developed rays in the anal, 13 branchiostegals, 23 (10 + 13) gill-rakers, and 45 to 80 pyloric cœca. There are about 127 scales in the lateral line. The scales are thin and all except those of the lateral line readily fall off. This feature distinguishes the species readily from the red salmon. In color it is silvery in spring, greenish above, and with a few faint black spots on the upper parts only. In the fall the males are mostly of a dirty red. The flesh in this species is of excellent flavor, but pale in color, and hence less valued than that of the quinnat and the red salmon.
The dog-salmon, calico salmon, or chum, called saké in Japan (_Oncorhynchus keta_), reaches an average weight of about 7 to 10 pounds. It has about 14 anal rays, 14 branchiostegals, 24 (9 + 15) gill-rakers, and 140 to 185 pyloric cœca. There are about 150 scales in the lateral line. In spring it is dirty silvery, immaculate, or sprinkled with small black specks, the fins dusky, the sides with faint traces of gridiron-like bars. In the fall the male is brick-red or blackish, and its jaws are greatly distorted. The pale flesh is well flavored when fresh, but pale and mushy in texture and muddy in taste when canned. It is said to take salt well, and great numbers of salt dog-salmon are consumed in Japan.
The humpback salmon, or pink salmon (_Oncorhynchus gorbuscha_), is the smallest of the American species, weighing from 3 to 5 pounds. It has usually 15 anal rays, 12 branchiostegals, 28 (13 + 15) gill-rakers, and about 180 pyloric cœca. Its scales are much smaller than in any other salmon, there being 180 to 240 in the lateral line. In color it is bluish above, silvery below, the posterior and upper parts with many round black spots, the caudal fin always having a few large black spots oblong in form. The males in fall are dirty red, and are more extravagantly distorted than in any other of the _Salmonidæ_. The flesh is softer than in the other species; it is pale in color, and, while of fair flavor when fresh, is distinctly inferior when canned.
The masu, or yezomasu (_Oncorhynchus masou_), is very similar to the humpback, the scales a little larger, the caudal without black spots, the back usually immaculate. It is one of the smaller salmon, and is fairly abundant in the streams of Hokkaido, the island formerly known as Yezo.
Of these species the blue-back or red salmon predominates in Frazer River and in most of the small rivers of Alaska, including all those which flow from lakes. The greatest salmon rivers of the world are the Nushegak and Karluk in Alaska, with the Columbia River, Frazer River, and Sacramento River farther south. The red and the silver salmon predominate in Puget Sound, the quinnat in the Columbia and the Sacramento, and the silver salmon in most of the smaller streams along the coast. All the species occur, however, from the Columbia northward; but the blue-back is not found in the Sacramento. Only the quinnat and the dog-salmon have been noticed south of San Francisco. In Japan _keta_ is by far the most abundant species of salmon. It is known as saké, and largely salted and sold in the markets. _Nerka_ is known in Japan only as landlocked in Lake Akan in northern Hokkaido. _Milktschitsch_ is generally common, and with _masou_ is known as masu, or small salmon, as distinguished from the large salmon, or saké. _Tschawytscha_ and _gorbuscha_ are unknown in Japan. _Masou_ has not been found elsewhere.
The quinnat and blue-back salmon, the "noble salmon," habitually "run" in the spring, the others in the fall. The usual order of running in the rivers is as follows: _tschawytscha_, _nerka_, _milktschitsch_, _gorbuscha_, _keta_. Those which run first go farthest. In the Yukon the quinnat runs as far as Caribou Crossing and Lake Bennett, 2250 miles. The red salmon runs to "Forty-Mile," which is nearly 1800 miles. Both ascend to the head of the Columbia, Fraser, Nass, Skeena, Stikeen, and Taku rivers. The quinnat runs practically only in the streams of large size, fed with melting snows; the red salmon only in streams which pass through lakes. It spawns only in small streams at the head of a lake. The other species spawn in almost any fresh water and only close to the sea.
The economic value of the spring-running salmon is far greater than that of the other species, because they can be captured in numbers when at their best, while the others are usually taken only after deterioration.
The habits of the salmon in the ocean are not easily studied. Quinnat and silver salmon of all sizes are taken with the seine at almost any season in Puget Sound and among the islands of Alaska. This would indicate that these species do not go far from the shore. The silver salmon certainly does not. The quinnat pursues the schools of herring. It takes the hook freely in Monterey Bay, both near the shore and at a distance of six to eight miles out. We have reason to believe that these two species do not necessarily seek great depths, but probably remain not very far from the mouth of the rivers in which they were spawned. The blue-back or red salmon certainly seeks deeper water, as it is seldom or never taken with the seine along shore, and it is known to enter the Strait of Fuca in July, just before the running season, therefore coming in from the open sea. The great majority of the quinnat salmon, and probably all the blue-back salmon, enter the rivers in the spring. The run of the quinnat begins generally at the last of March; it lasts, with various modifications and interruptions, until the actual spawning season in November, the greatest run being in early June in Alaska, in July in the Columbia. The run begins earliest in the northernmost rivers, and in the longest streams, the time of running and the proportionate amount in each of the subordinate runs varying with each different river. In general the runs are slack in the summer and increase with the first high water of autumn. By the last of August only straggling blue-backs can be found in the lower course of any stream; but both in the Columbia and in the Sacramento the quinnat runs in considerable numbers at least till October. In the Sacramento the run is greatest in the fall, and more run in the summer than in spring. In the Sacramento and the smaller rivers southward there is a winter run, beginning in December. The spring quinnat salmon ascends only those rivers which are fed by the melting snows from the mountains and which have sufficient volume to send their waters well out to sea. Those salmon which run in the spring are chiefly adults (supposed to be at least three years old). Their milt and spawn are no more developed than at the same time in others of the same species which have not yet entered the rivers. It would appear that the contact with cold fresh water, when in the ocean, in some way causes them to run towards it, and to run before there is any special influence to that end exerted by the development of the organs of generation. High water on any of these rivers in the spring is always followed by an increased run of salmon. The salmon-canners think—and this is probably true—that salmon which would not have run till later are brought up by the contact with the cold water. The cause of this effect of cold fresh water is not understood. We may call it an instinct of the salmon, which is another way of expressing our ignorance. In general it seems to be true that in those rivers and during those years when the spring run is greatest the fall run is least to be depended on.
The blue-back salmon runs chiefly in July and early August, beginning in late June in Chilcoot River, where some were found actually spawning July 15; beginning after the middle of July in Frazer River.
As the season advances, smaller and younger salmon of these species (quinnat and blue-back) enter the rivers to spawn, and in the fall these young specimens are very numerous. We have thus far failed to notice any gradations in size or appearance of these young fish by which their ages could be ascertained. It is, however, probable that some of both sexes reproduce at the age of one year. In Frazer River, in the fall, quinnat male grilse of every size, from eight inches upwards, were running, the milt fully developed, but usually not showing the hooked jaws and dark colors of the older males. Females less than eighteen inches in length were not seen. All of either sex, large and small, then in the river had the ovaries or milt developed. Little blue-backs of every size, down to six inches, are also found in the upper Columbia in the fall, with their organs of generation fully developed. Nineteen-twentieths of these young fish are males, and some of them have the hooked jaws and red color of the old males. Apparently all these young fishes, like the old ones, die after spawning.
The average weight of the adult quinnat in the Columbia, in the spring, is twenty-two pounds; in the Sacramento, about sixteen. Individuals weighing from forty to sixty pounds are frequently found in both rivers, and some as high as eighty or even one hundred pounds are recorded, especially in Alaska, where the species tends to run larger. It is questionable whether these large fishes are those which, of the same age, have grown more rapidly; those which are older, but have for some reason failed to spawn; or those which have survived one or more spawning seasons. All these origins may be possible in individual cases. There is, however, no positive evidence that any salmon of the Pacific survives the spawning season.
Those fish which enter the rivers in the spring continue their ascent till death or the spawning season overtakes them. Doubtless not one of them ever returns to the ocean, and a large proportion fail to spawn. They are known to ascend the Sacramento to its extreme head-waters, about four hundred miles. In the Columbia they ascend as far as the Bitter Root and Sawtooth mountains of Idaho, and their extreme limit is not known. This is a distance of nearly a thousand miles. In the Yukon a few ascend to Caribou Crossing and Lake Bennett, 2250 miles. At these great distances, when the fish have reached the spawning grounds, besides the usual changes of the breeding season their bodies are covered with bruises, on which patches of white fungus (_Saprolegnia_) develop. The fins become mutilated, their eyes are often injured or destroyed, parasitic worms gather in their gills, they become extremely emaciated, their flesh becomes white from the loss of oil; and as soon as the spawning act is accomplished, and sometimes before, _all_ of them die. The ascent of the Cascades and the Dalles of the Columbia causes the injury or death of a great many salmon.
When the salmon enter the river they refuse to take bait, and their stomachs are always found empty and contracted. In the rivers they do not feed; and when they reach the spawning grounds their stomachs, pyloric cœca and all, are said to be no larger than one's finger. They will sometimes take the fly, or a hook baited with salmon-roe, in the clear waters of the upper tributaries, but this is apparently solely out of annoyance, snapping at the meddling line. Only the quinnat and blue-back (there called redfish) have been found at any great distance from the sea, and these (as adult fishes) only in late summer and fall.
The spawning season is probably about the same for all the species. It varies for each of the different rivers, and for different parts of the same river. It doubtless extends from July to December, and takes place usually as soon as the temperature of the water falls to 54°. The manner of spawning is probably similar for all the species. In the quinnat the fishes pair off; the male, with tail and snout, excavates a broad, shallow "nest" in the gravelly bed of the stream, in rapid water, at a depth of one to four feet and the female deposits her eggs in it. They then float down the stream tail foremost, the only fashion in which salmon descend to the sea. As already stated, in the head-waters of the large streams, unquestionably, all die; it is the belief of the writer that none ever survive. The young hatch in sixty days, and most of them return to the ocean during the high water of the spring. They enter the river as adults at the age of about four years.
The salmon of all kinds in the spring are silvery, spotted or not according to the species, and with the mouth about equally symmetrical in both sexes. As the spawning season approaches the female loses her silvery color, becomes more slimy, the scales on the back partly sink into the skin, and the flesh changes from salmon-red and becomes variously paler, from the loss of oil; the degree of paleness varying much with individuals and with inhabitants of different rivers. In the Sacramento the flesh of the quinnat, in either spring or fall, is rarely pale. In the Columbia a few with pale flesh are sometimes taken in spring, and an increasing number from July on. In Frazer River the fall run of the quinnat is nearly worthless for canning purposes, because so many are "white-meated." In the spring very few are "white-meated"; but the number increases towards fall, when there is every variation, some having red streaks running through them, others being red toward the head and pale toward the tail. The red and pale ones cannot be distinguished externally, and the color is dependent on neither age nor sex. There is said to be no difference in the taste, but there is little market for canned salmon not of the conventional orange-color.
As the season advances the difference between the males and females becomes more and more marked, and keeps pace with the development of the milt, as is shown by dissection. The males have (1) the premaxillaries and the tip of the lower jaw more and more prolonged, both of the jaws becoming finally strongly and often extravagantly hooked, so that either they shut by the side of each other like shears, or else the mouth cannot be closed. (2) The front teeth become very long and canine-like, their growth proceeding very rapidly, until they are often half an inch long. (3) The teeth on the vomer and tongue often disappear. (4) The body grows more compressed and deeper at the shoulders, so that a very distinct hump is formed; this is more developed in the humpback salmon, but is found in all. (5) The scales disappear, especially on the back, by the growth of spongy skin. (6) The color changes from silvery to various shades of black and red, or blotchy, according to the species. The blue-back turns rosy-red, the head bright olive; the dog-salmon a dull red with blackish bars, and the quinnat generally blackish. The distorted males are commonly considered worthless, rejected by the canners and salmon-salters, but preserved by the Indians. These changes are due solely to influences connected with the growth of the reproductive organs. They are not in any way due to the action of fresh water. They take place at about the same time in the adult males of all species, whether in the ocean or in the rivers. At the time of the spring runs all are symmetrical. In the fall all males, of whatever species, are more or less distorted. Among the dog-salmon, which run only in the fall, the males are hook-jawed and red-blotched when they first enter the Strait of Fuca from the outside. The humpback, taken in salt water about Seattle, have the same peculiarities. The male is slab-sided, hook-billed, and distorted, and is rejected by the canners. No hook-jawed females of any species have been seen.
On first entering a stream the salmon swim about as if playing. They always head towards the current, and this appearance of playing may be simply due to facing the moving tide. Afterwards they enter the deepest parts of the stream and swim straight up, with few interruptions. Their rate of travel at Sacramento is estimated by Stone at about two miles per day; on the Columbia at about three miles per day. Those which enter the Columbia in the spring and ascend to the mountain rivers of Idaho must go at a more rapid rate than this, as they must make an average of nearly four miles per day.
As already stated, the economic value of any species depends in great part on its being a "spring salmon." It is not generally possible to capture salmon of any species in large numbers until they have entered the estuaries or rivers, and the spring salmon enter the large rivers long before the growth of the organs of reproduction has reduced the richness of the flesh. The fall salmon cannot be taken in quantity until their flesh has deteriorated; hence the dog-salmon is practically almost worthless except to the Indians, and the humpback salmon was regarded as little better until comparatively recently, when it has been placed on the market in cans as "Pink Salmon." It sells for about half the price of the red salmon and one-third that of the quinnat. The red salmon is smaller than the quinnat but, outside the Sacramento and the Columbia, far more abundant, and at present it exceeds the quinnat in economic value. The pack of red salmon in Alaska amounted in 1902 to over two million cases (48 pounds each), worth wholesale about $4.00 per case, or about $8,000,000. The other species in Alaska yield about one million cases, the total wholesale value of the pack for 1902 being $8,667,673. The aggregate value of the quinnat is considerably less, but either species far exceed in value all other fishes of the Pacific taken together. The silver salmon is found in the inland waters of Puget Sound for a considerable time before the fall rains cause the fall runs, and it may be taken in large numbers with seines before the season for entering the rivers.
The fall salmon of all species, but especially of the dog-salmon, ascend streams but a short distance before spawning. They seem to be in great anxiety to find fresh water, and many of them work their way up little brooks only a few inches deep, where they perish miserably, floundering about on the stones. Every stream of whatever kind, from San Francisco to Bering Sea, has more or less of these fall salmon.
The absence of the fine spring salmon in the streams of Japan is the cause of the relative unimportance of the river fisheries of the northern island of Japan, Hokkaido. It is not likely that either the quinnat or the red salmon can be introduced into these rivers, as they have no snow-fed streams, and few of them pass through lakes which are not shut off by waterfalls. For the same reason neither of these species is likely to become naturalized in the waters of our Eastern States, though it is worth while to bring the red salmon to the St. Lawrence. The silver salmon, already abundant in Japan, should thrive in the rivers and bays of New England.
=The Parent-stream Theory.=—It has been generally accepted as unquestioned by packers and fishermen that salmon return to spawn to the very stream in which they were hatched. As early as 1880 the present writer placed on record his opinion that this theory was unsound. In a general way most salmon return to the parent stream, because when in the sea the parent stream is the one most easily reached. The channels and runways which directed their course to the sea may influence their return trip in the same fashion. When the salmon is mature it seeks fresh water. Other things being equal, about the same number will run each year in the same channel. With all this, we find some curious facts. Certain streams will have a run of exceptionally large or exceptionally small red salmon. The time of the run bears some relation to the length of the stream: those who have farthest to go start earliest. The time of running bears also a relation to the temperature of the spawning grounds: where the waters cool off earliest the fish run soonest.
The supposed evidence in favor of the parent-stream theory may be considered under three heads:[9] (1) Distinctive runs in various streams. (2) Return of marked salmon. (3) Introduction of salmon into new streams followed by their return.
Footnote 9:
See an excellent article by H. S. Davis in the _Pacific Fisherman_ for July, 1903.
Under the first head it is often asserted of fishermen that they can distinguish the salmon of different streams. Thus the Lynn Canal red salmon are larger than those in most waters, and it is claimed that those of Chilcoot Inlet are larger than those of the sister stream at Chilcat. The red salmon of Red Fish Bay on Baranof Island are said to be much smaller than usual, and those of the neighboring Necker Bay are not more than one-third the ordinary size. Those of a small rapid stream near Nass River are more wiry than those of the neighboring large stream. The same claim is made for the different streams of Puget Sound, each one having its characteristic run. In all this there is some truth and perhaps some exaggeration. I have noticed that the Chilcoot fish seem deeper in body than those at Chilcat. The red salmon becomes compressed before spawning, and the Chilcoot fishes having a short run spawn earlier than the Chilcat fishes, which have many miles to go, the water being perhaps warmer at the mouth of the river. Perhaps some localities may meet the nervous reactions of small fishes, while not attracting the large ones. Mr. H. S. Davis well observes that "until a constant difference has been demonstrated by a careful examination of large numbers of fish from each stream taken _at the same time_, but little weight can be attached to arguments of this nature."
It is doubtless true as a general proposition that nearly all salmon return to the region in which they were spawned. Most of them apparently never go far away from the mouth of the stream or the bay into which it flows. It is true that salmon are occasionally taken well out at sea, and it is certain that the red salmon runs of Puget Sound come from outside the Straits of Fuca. There is, however, evidence that they rarely go so far as that. When seeking shore they do not reach the original channels.
In 1880 the writer, studying the salmon of the Columbia, used the following words, which he has not had occasion to change:
"It is the prevailing impression that the salmon have some special instinct which leads them to return to spawn in the same spawning grounds where they were originally hatched. We fail to find any evidence of this in the case of the Pacific-coast salmon, and we do not believe it to be true. It seems more probable that the young salmon hatched in any river mostly remain in the ocean within a radius of twenty, thirty, or forty miles of its mouth. These, in their movements about in the ocean, may come into contact with the cold waters of their parent rivers, or perhaps of any other river, at a considerable distance from the shore. In the case of the quinnat and the blue-back their 'instinct' seems to lead them to ascend these fresh waters, and in a majority of cases these waters will be those in which the fishes in question were originally spawned. Later in the season the growth of the reproductive organs leads them to approach the shore and search for fresh waters, and still the chances are that they may find the original stream. But undoubtedly many fall salmon ascend, or try to ascend, streams in which no salmon was ever hatched. In little brooks about Puget Sound, where the water is not three inches deep, are often found dead or dying salmon which have entered them for the purpose of spawning. It is said of the Russian River and other California rivers that their mouths, in the time of low water in summer, generally become entirely closed by sand-bars, and that the salmon, in their eagerness to ascend them, frequently fling themselves entirely out of water on the beach. But this does not prove that the salmon are guided by a marvelous geographical instinct which leads them to their parent river in spite of the fact that the river cannot be found. The waters of Russian River soak through these sand-bars, and the salmon instinct, we think, leads them merely to search for fresh waters. This matter is much in need of further investigation; at present, however, we find no reason to believe that the salmon enter the Rogue River simply because they were spawned there, or that a salmon hatched in the Clackamas River is more likely, on that account, to return to the Clackamas than to go up the Cowlitz or the Des Chûtes."
Attempts have been made to settle this question by marking the fry. But this is a very difficult matter indeed. Almost the only structure which can be safely mutilated is the adipose fin, and this is often nipped off by sticklebacks and other meddling fish. The following experiments have been tried, according to Mr. Davis:
In March, 1896, 5000 king-salmon fry were marked by cutting off the adipose fin, then set free in the Clackamas River. Nearly 400 of these marked fish are said to have been taken in the Columbia in 1898, and a few more in 1899. In addition a few were taken in 1898, 1899, and 1900 in the Sacramento River, but in much less numbers than in the Columbia. In the Columbia most were taken at the mouth of the river, where nearly all of the fishing was done, but a few were in the original stream, the Clackamas. It is stated that the fry thus set free in the Clackamas came from eggs obtained in the Sacramento—a matter which has, however, no bearing on the present case.
In the Kalama hatchery on the Columbia River, Washington, 2000 fry of the quinnat or king-salmon were marked in 1899 by a V-shaped notch in the caudal fin. Numerous fishes thus marked were taken in the lower Columbia in 1901 and 1902. A few were taken at the Kalama hatchery, but some also at the hatcheries on Wind River and Clackamas River. At the hatchery on Chehalis River six or seven were taken, the stream not being a tributary of the Columbia, but flowing into Shoalwater Bay. None were noticed in the Sacramento. The evidence shows that the most who are hatched in a large stream tend to return to it, and that in general most salmon return to the parent region. There is no evidence that a salmon hatched in one branch of a river tends to return there rather than to any other. Experiments of Messrs. Rutter and Spaulding in marking adult fish at Karluk would indicate that they roam rather widely about the island before spawning. An adult spawning fish, marked and set free at Karluk, was taken soon after on the opposite side of the island of Kadiak.
The introduction of salmon into new streams may throw some light on this question. In 1897 and 1898 3,000,000 young quinnat-salmon fry were set free in Papermill Creek near Olema, California. This is a small stream flowing into the head of Tomales Bay, and it had never previously had a run of salmon. In 1900, and especially in 1901, large quinnat salmon appeared in considerable numbers in this stream. One specimen weighing about sixteen pounds was sent to the present writer for identification. These fishes certainly returned to the parent stream, although this stream was one not at all fitted for their purpose.
But this may be accounted for by the topography of the bay. Tomales Bay is a long and narrow channel, about twenty miles long and from one to five in width, isolated from other rivers and with but one tributary stream. Probably the salmon had not wandered far from it; some may not have left it at all. In any event, a large number certainly came back to the same place.
That the salmon rarely go far away is fairly attested. Schools of king-salmon play in Monterey Bay, and chase the herring about in the channels of southeastern Alaska. A few years since Captain J. F. Moser, in charge of the _Albatross_, set gill-nets for salmon at various places in the sea off the Oregon and Washington coast, catching none except in the bays.
Mr. Davis gives an account of the liberation of salmon in Chinook River, which flows into the Columbia at Baker's Bay:
"It is a small, sluggish stream and has never been frequented by Chinook salmon, although considerable numbers of silver and dog salmon enter it late in the fall. A few years ago the State established a hatchery on this stream, and since 1898 between 1,000,000 and 2,000,000 Chinook fry have been turned out here annually. The fish are taken from the pound-nets in Baker's Bay, towed into the river in crates and then liberated above the dike, which prevents their return to the Columbia. When ripe the salmon ascend to the hatchery, some two or three miles farther up the river, where they are spawned.
"The superintendent of the hatchery, Mr. Hansen, informs me that in 1902, during November and December, quite a number of Chinook salmon ascended the Chinook River. About 150 salmon of both sexes were taken in a trap located in the river about four miles from its mouth. At first thought it would appear that these were probably fish which, when fry, had been liberated in the river, but unfortunately there is no proof that this was the case. According to Mr. Hansen, the season of 1902 was remarkable in that the salmon ran inshore in large schools, a thing which they had not done before for years. It is possible that the fish, being forced in close to the shore, came in contact with the current from the Chinook River, which, since the stream is small and sluggish, would not be felt far from shore. Once brought under the influence of the current from the river, the salmon would naturally ascend that stream, whether they had been hatched there or not."
The general conclusion, apparently warranted by the facts at hand, is that salmon, for the most part, do not go to a great distance from the stream in which they are hatched, that most of them return to the streams of the same region, a majority to the parent stream, but that there is no evidence that they choose the parental spawning grounds in preference to any other, and none that they will prefer an undesirable stream to a favorable one for the reason that they happen to have been hatched in the former.
=The Jadgeska Hatchery.=—Mr. John C. Callbreath of Wrangel, Alaska, has long conducted a very interesting but very costly experiment in this line. About 1890 he established himself in a small stream called Jadgeska on the west coast of Etolin Island, tributary to McHenry Inlet, Clarence Straits. This stream led from a lake, and in it a few thousand red salmon spawned, besides multitudes of silver salmon, dog-salmon, and humpback salmon. Making a dam across the stream, he helped the red salmon over it, destroying all of the inferior kinds which entered the stream. He also established a hatchery for the red salmon, turning loose many fry yearly for ten or twelve years. This was done in the expectation that all the salmon hatched would return to Jadgeska in about four years. By destroying all individuals of other species attempting to run, it was expected that they would become extinct so far as the stream is concerned.
The result of this experiment has been disappointment. After twelve years or more there has been no increase of red salmon in the stream, and no decrease of humpbacks and other humbler forms of salmon. Mr. Callbreath draws the conclusion that salmon run at a much greater age than has been supposed—at the age of sixteen years, perhaps, instead of four. A far more probable conclusion is that his salmon have joined other bands bound for more suitable streams. It is indeed claimed that since the establishment of Callbreath's hatchery on Etolin Island there has been a notable increase of the salmon run in the various streams of Prince of Wales Island on the opposite side of Clarence Straits. But this statement, while largely current among the cannerymen, and not improbable, needs verification.
We shall await with much interest the return of the thousands of salmon hatched in 1902 in Naha stream. We may venture the prophecy that while a large percentage will return to Loring, many others will enter Yes Bay, Karta Bay, Moira Sound, and other red salmon waters along the line of their return from Dixon Entrance or the open sea.
=Salmon-packing.=—The canning of salmon, that is, the packing of the flesh in tin cases, hermetically sealed after boiling, was begun on the Columbia River by the Hume Brothers in 1866. In 1874 canneries were established on the Sacramento River, in 1876 on Puget Sound and on Frazer River, and in 1878 in Alaska. At first only the quinnat salmon was packed; afterwards the red salmon and the silver salmon, and finally the humpback, known commercially as pink salmon. In most cases the flesh is packed in one-pound tins, forty-eight of which constitute a case. The wholesale price in 1903 was for quinnat salmon $5.60 per case, red salmon $4.00, silver salmon $2.60, humpback salmon $2.00, and dog-salmon $1.50. It costs in round numbers $2.00 to pack a case of salmon. The very low price of the inferior brands is due to overproduction.
The output of the salmon fishery of the Pacific coast amounts to about fifteen millions per year, that of Alaska constituting seven to nine millions of this amount. Of this amount the red salmon constitutes somewhat more than half, the quinnat about four-fifths of the rest.
In almost all salmon streams there is evidence of considerable diminution in numbers, although the evidence is sometimes conflicting. In Alaska this has been due to the vicious custom, now done away with, of barricading the streams so that the fish could not reach the spawning grounds, but might be all taken with the net. In the Columbia River the reduction in numbers is mainly due to stationary traps and salmon-wheels, which leave the fish relatively little chance to reach the spawning grounds. In years of high water doubtless many salmon run in the spring which might otherwise have waited until fall.
The key to the situation lies in the artificial propagation of salmon by means of well-ordered hatcheries. By this means the fisheries of the Sacramento have been fully restored, those of the Columbia approximately maintained, and a hopeful beginning has been made in hatching red salmon in Alaska.