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Fossil shells from the Eocene marl-beds at Jackson. THE Agricultural and Geological Survey of the State originated in an act of the Legislature, approved the 5th of March, , to take effect on the first of June following, entitled "an Act further to endow the University of Mississippi;" and its execution was committed to that Institution.

No assistant was obtained until the latter part of , and the gentleman then appointed relinquished the situation shortly after; having merely commenced a reconnoissance of the State, of which no report was made. The situation was tendered to the present incumbent on the 14th of January, Since that time he has been occupied in the performance of the duties committed to him, which were somewhat augmented by an amendatory act of the Legislature, passed after his appointment, by which a room in the State House was set apart and placed under his charge, for the better preservation of the collections in Natural History, which, as the State Geologist, he was required to make.

In the prosecution of this work, a, considerable portion of the State has been traversed, with a view of gaining such general knowledge of its character as would best guide and direct the subsequent, more detailed, and minute examination to be made. More than seven thousand three hundred miles have been travelled, collections amounting to several thousand specimens have been made, and the character, peculiarities, and productions of the different sections visited, have been observed and noted.

It was doubtless with a general knowledge of the geological features of the State that the Survey was authorized by the Legislature. Consisting chiefly of the more recent formations, the absence of the primitive and metalliferous rocks, in place, gave no reason to expect the existence of those ores and minerals which belong properly to an earlier period, and which constitute the chief resources of less favored and fertile districts than ours. The discovery of mines of copper, lead, or of the more precious metals, or, even of the true coal-fields, was obviously not to be expected.

It was, therefore, mainly in reference to its influence and bearing upon the agricultural prosperity of the State, that it was undertaken. XV calcareous fertilizers are as abundant, as varied ill character, and excellent in quality, as any other State can boast.

To ascertain and point out the chief deposits of these marls, and to determine their relative value and chemical constituents, becomes now an object of much importance. Exact analyses of the different varieties, characterizing them by the prevailing fossils, when such exist, so as at once to be identified by the planter, should be made; numerous experiments in their application should be encouraged; and the effects upon the growth of our different agricultural productions should be diligently observed, and accurately detailed.

The attention of planters has been pointed to these fertilizers on all suitable occasions, and in a few instances experiments on a limited scale have been commenced. Specimens have also been collected, with their-associated and characteristic fossils, and have been deposited for general inspection, in the State Cabinet at the Capitol, and in the Cabinet of the University at Oxford. Analyses of many varieties of our marls and soils should have been given in this Report.

Few of these, however, have been procured, owing, in part, to a defect in the law authorizing the Survey, and to the illness and subsequent resignation of Dr. The latter event occurring at a period so nearly approaching to that at which a report of the progress of the Survey was required, devolved unexpectedly upon the assistant that duty, which, under the existing circumstances, must otherwise have been unperformed.

Whilst captious and ill-natured criticism is ever to be deprecated, a fair and proper correction of error is as much to be desired; and in an essay of this character, in which the object is to impart useful knowledge, is rather to be invited. Errors have doubtless occurred in treating of the multifarious topics which are embraced in this Report; and to the end that the greatest accuracy may be attained, the writer will be gratified to have them pointed out, in order that they may be corrected hnd avoided in future.

Of the plan of this Report, it will be seen that, with the sanction of approved precedents, it has been considered that a short preliminary sketch of the discovery and early history of the country, not hitherto separately written, would not be out of place. In compiling and abridging this from other writers, it has been a somewhat difficult task to condense it within the required limits, except at the expense of much of the interest that would attach to a more detailed account.

To these highly respectable and authentic sources of information with respect to our early history, it gives me pleasure to acknowledge my indebtedness. From the manuscript correspondence of the late Mr. William Dunbar " of the Forest," I have also been enabled to glean some interesting facts, and to the representatives of his family I have to express my thanks for the opportunity afforded me of consulting it.

The Spanish archives preserved in the State, have also, to a limited extent, been consulted, and, had time permitted, might have been more profitably explored. I have to regret, notwithstanding, that this sketch is not more complete; the more so, as there is reason to believe that some authentic and interesting documents are preserved in the State, not as yet made public, which, if accessible, would no doubt serve to fill up some of the chasms and otherwise explain and illustrate our early history.

These I should have been pleased to avail myself of. As a subject of interest to the landed proprietors of the State, the chapter on Land Titles was considered as germain to the subject, and entitled to the short space which it occupies. In the details given of the different agricultural productions, the mode of cultivation, and the machinery for preparing these, I have been similarly aided. In all that has been said in this connection, universal concurrence is not expected.

If it be maintained that any of these details are erroneous, I can only say that any such will be most willingly corrected, when it can be done on better autlhority than that on which any specific fact or statement has been given. The tables of agricultural and other statistics have been prepared from the best sources, and will form matter for convenient and useful reference.

At this stage of the Survey, and in the first, and as it may be termed preliminary report, the notice of the Geology and other departments of natural history, will necessarily present a mere outline, and cannot assume that form and shape which will properly be given them in a final report.

Such an arrangement has been adopted, however, as far as these subjects are embraced, as will, it is believed, give a reasonably comprehensive and familiar view of those departments of the Report. Of the Fauna and Flora of the State, in the notice that has been taken of them, my own observations have been directed by the best available authorities; and in the former department, among others the works of DeKay, and of Audubon, and Bachman, among the most recent published, and by inference, the most complete and correct, have been consulted.

The aid of distinguished naturalists, also, has been liberally afforded; and I have to acknowledge my indebtedness, and express my thanks, to Professors Agassiz and Baird, and to Mr. Conrad, for their contributions to this department of the Report. The catalogues furnished by them, although not so complete or perfect as they will hereafter be made, have the stamp of authenticity and accuracy to recommend them. XiX to acknowledge the obligations I am also under to Dr.

Leidy and Mr. Cassin, of the Academy of Natural Sciences, of Philadelphia. As to the illustrations which accompany this Report, the limited means appropriated to the Survey, and the dearth of artistic skill available in this quarter, have made me dependent upon the early, imperfect, and selftaught attainment of drawing; and which, having been almost wholly unpractised for nearly thirty years, makes an apology necessary for their rude and unsatisfactory execution.

In making the collections required, the cases in the State Cabinet attest that a reasonable progress has been made with the means appropriated to this object, and upwards of a thousand duplicates have been deposited in the University at Oxford, for its cabinet. In my travels through the State, on this not very generally understood or properly appreciated mission, it was to be expected that occasionally little either of information or assistance would be afforded.

To those gentlemen whose hospitality and assistance have been kindly and liberally extended to me, on my various excursions in different quarters, I can only offer my sincere thanks, and express the hope that they may derive some gratification, if not profit, from the final issue of the Survey. Were a precedent required to sanction the very abridged historical outline here introduced, a distinguished one may be found in the able and elaborate memoir that forms the introduction to the Reports on the Natural History of New York, embracing a much wider scope than is here proposed, and comprehending the political history and social progress of the State.

To keep this sketch within the prescribed limits, and to exclude all matter not intimately connected with the subject, it will be restricted to occurrences strictly within the present boundaries of the State, except so far as may be necessary to preserve the natural sequence of events.

It will embrace little more, therefore, than a chronological outline, which, if desirable, may, as far as necessary, be enlarged in the final report of the Survey. On the west it is bounded by the Mississippi and Pearl Rivers, and on the east by a line dividing it from the State of Alabama, which is drawn from the mouth of Bear Creek on the Tennessee River to the northwestern corner of Washington County, Alabama, and thence south to a point on Grand Bay on the Gulf of Mexico, about seventeen miles due west from the Bay of Mobile.

The State also embraces the islands in the Gulf within six leagues of the northern shore, the principal of which are Horn, Ship, and Cat Islands. The width of the State along the northern boundary is one hundred and twenty miles; on the sea-shore seventy-eight miles; and along the of north latitude one hundred and eighty-six miles.

The greatest length from north to south is three hundred and thirty miles. It embraces an area of 55, square miles or 35,, acres. About the close of the year , Fernando de Soto, in his adventurous and romantic expedition, commenced the preceding year at the Bay of Espiritu Santo, and designed for the conquest of Florida, penetrated to the country of the Chickasaws in the northeastern part of the State. Here he continued, notwithstanding his disasters and the persevering ahd galling attacks of the Indians, until the 1st of April, The position of this winter encampment is conjectured to have been near the northeastern part of Pontitoc County, where, it is said, remains of ancient fortifications are still to be seen, and relics of European origin probably pertaining to this expedition have also been found.

Thus, far in the interior, distant from the sea-shore, and remote from the Mississippi, was the territory of the State first entered upon by Europeans. It is needless to trace his subsequent wanderings if it were practicable, even with approximate accuracy, to do so. There is little doubt, however, that De Soto traversed the country comprising the county now bearing his name, and in May of the same year discovered the Mississippi River, called by the natives "c Cicuaga," at a point near the extreme northwestern corner of the State.

Some writers assign the mouth of the Arkansas, and others that of Red River, as the place of De Soto's death; and the town of "Guachoa" is laid down on an English map published in 17 64 at the latter place. Still a year later, his followers, now led by Louis de MIuscoso, failing to reach Mexico by land, returned to the same village on the Mississippi, on which the small remnant of the expedition, reduced to about three hundred and fifty survivors, little more than one-fourth of the number of which it was first composed, embarked on the 2d of July, , for a final departure from the country, pursued and sorely harassed by the Indians, and arrived at the sea-shore after a voyage of twenty days.

From this period, for an interval of nearly one hundred and thirty-eight years, the native tribes were left in undisturbed possession of the country; and it was not until February, , when La Salle, accompanied by the Chevalier de Tonti, descended the Mississippi from Canada, that the country was revisited by European adventurers.

In April of this year, La Salle, having reached the ocean, on his return touched at the settlement of the Natchez, from which the hostile bearing of that people hastened his departure. Failing in his subsequent expedition, fitted out in France with a view to the establishment of a colony, to reach the mouth of the Mississippi by sea, having passed to the west of it, La Salle perished miserably in Texas by the hands of his despairing and mutinous followers; and another interval of eighteen years elapsed before the country was again visited by Europeans.

The authority of the map is not to be relied upon, since, among other inaccuracies, it places New Orleans above the Lafourche. In February, , an expedition led by Iberville arrived upon the coast, and occupied Ship Island. Iberville had offered to prosecute the plan of La Salle to colonize Louisiana, and under the patronage of Count de Pontchartrain, the French minister of marine, was put in command of an expedition fitted out at La Rochelle, consisting of two frigates and two smaller vessels, to be employed in this service.

After exploring the shores and inlets in that quarter, it was resolved to establish the proposed colony on the main land in the vicinity, and accordingly a landing was effected on the eastern extremity of the Bay of Baluxi. A fort of four bastions, with twelve pieces of cannon, was commenced on the first of May following, the colonists brought over by the expedition numbering about two hundred, including women and children, were settled around the fort, and the first European settlement was established in Mississippi.

Iberville, leaving his brothers in charge of the settlement, the elder, Sauvolle, as governor, and the younger, Bienville, as his lieutenant, set sail on his return to France for the purpose of reinforcing the infant colony he had founded, and procuring for it the necessary supplies.

In July, soon after the departure of Iberville, the colony was visited by two missionaries, Montegay and Davion, who had wandered from Canada, and had been residing among the Indian tribes. Bienville, who engaged actively in exploring the passes and outlets of the Mississippi, encountered an English ship in the river, commanded by Captain Bar, one of two vessels sent out by Daniel Cox of New Jersey, to take possession of a grant of land of which he was then the proprietor, made by Charles the First of England, in , to Sir Robert Heath.

It comprised a tract of truly royal dimensions, embracing not only the present State of Mississippi, but included several other adjoining States. Captain Bar, doubting whether the stream he had entered was the Mississippi, was easily induced by Bienville to retrace his steps; and the great bend in the river, at which his progress was terminated, has ever since been known from this circumstance as the "English Turn.

He brought out Leseur, a Geologist, who was sent by the French government to examine a greenish earth or ochre which had been noticed on the banks of the Mississippi. Peter's, which he ascended a considerable distance.

A greenish ochre was found covering the ground near a copper-mine, thirteen thousand pounds of which were gathered, brought to Baluxi and shipped to France; but no further notice appears to have been taken of it. This was the third voyage of this energetic and enterprising man down the Mississippi, first in company with La Salle, when he explored the river to its mouth, and again for the purpose of meeting his old associate and friend, who, he was apprised, was attempting to enter the Mississippi by sea, in which he was grievously disappointed.

De Tonti had distinguished himself in the European wars, and had lost a hand, which he had had supplied by an artificial substitute of iron, of which at times he was wont to make a formidable use, and which procured for him the sobriquet of the " Iron I-land.

There they met with St. Come, a missionary from Canada, who had fixed his residence among this people. The Natchez, greatly advanced beyond the other Indian tribes in civilization, had been reduced from a once powerful nation, and now numbered only about twelve hundred warriors. The Great Sun, as their king was termed, welcomed the French at the head of a large retinue, borne in state on the shoulders of some of his attendants. They were worshippers of the sun, and maintained a perpetual fire in their temples.

One of these, during the visit of the French, was set on fire by lightning, when the frenzied and superstitious women, at the call of the vociferating and demoniac priests, cast their infant children into the flames to appease their irritated divinity. Sauvolle died in July, , after the departure of Iberville, and was succeeded by Bienville as governor. The colonists suffered greatly from the want of provisions; and in the fall, disease following in the track of famine, many died, the number of survivors being reduced to one hundred and fifty.

The return of Iberville from France, late in December, afforded a timely relief. Besides the supplies, he brought with him also a reinforcement of troops. Under instructions from the king, Bienville moved his head-quarters to the western bank of Mobile River, leaving a detachment of twenty men in charge of the fort at Baluxi. A fort, with barracks and stores, was also erected on Dauphin Island, which possessed a better port and more convenient landing than Ship Island afforded. The seat of government of the Province being transferred beyond the present limits of the State, and there remaining within it but the small settlement at Baluxi, it will suffice to state, in reference to the progress of the colony for many years, that it was characterized by an entire neglect of agricultural pursuits, and that it was subjected to great hardships from famine and disease, the occasional supplies derived from France, St.

Domingo, and Vera Cruz, being so inadequate as to render it necessary occasionally to quarter the troops upon the adjacent Indian tribes to gain a precarious subsistence by hunting and fishing. Bienville, being appointed to the command of the establishments on the Mississippi, learning that the Natchez had plundered and killed some Frenchmen, led a detachment of troops against them, in , and, having decoyed some of their chiefs into his camp, compelled the restoration of the plundered goods and the punishment of the offenders; after which he accompanied the Natchez to their village, and with their assent conmmenced a fort on the spot Iberville had before chosen.

It was called Rosalie, and in June a small garrison was established in it under the command of an officer named Pailloux. The earthen mound or embankment which tradition points out as the site of this fort, is still to be seen crowning the bluff of the river, immediately below and in the suburbs of the city of Natchez. When the country camue under the dominion of Great Britain, it was called Fort Panmure, after a barony of that name in Scotland, a name it retained during the subsequent rule of the Spaniards, being so designated in all the grants of land made by that government.

Three of Crozat's ships arrived in March, , with three companies of infantry and fifty new colonists. Bienville was superseded as governor; and although the order of knighthood was conferred upon him in reward for his services, yet the arrival of L'Epinay, his successor, occasioned him much mortification, which the decoration of the cross of St.

Failing to establish a commerce with the Spaniards in Mexico, and disappointed in all his expectations, Crozat, in August, , surrendered his grant to the king. Marbois, however, attributes to him more statesmanship than was possessed by the ministers, and adds that his plans were wisely conceived, and as far as depended upon him he sent to the colony only robust and industrious people, and families recommended by their morals, who were the only settlers that succeeded.

In September, , a charter was granted to a new corporation, styled the'" Western Company," which originated with the celebrated Scotch adventurer and financier Law, a pr'ote'g of the Regent Duke of Orleans. It was also known as the "Mississippi Scheme. The company was authorized to nominate the governor and other officers, to grant lands, to levy troops, make treaties, and wage war with the Indians, and generally to exercise the most unlimited and extraordinary powers.

Of this powerful and privileged company, John Law was appointed Director-General. One of the first acts of the company, in February, , was to recall L'Epinay, and to reinstate Bienville as governor-a measure which gave great satisfaction to the troops, and to the inhabitants generally.

As the most effectual mode of encouraging agricultural enterprise, it was deemed expedient to make considerable concessions of land to wealthy and powerful personages: among these were grants of large extent, on the Yazoo River, to a company consisting of Le Blanc, Count de Belleville, Leblond, and others; and on St.

The Bay of St. The condition of all such grants was, the introduction of a certain number of emigrants upon them within a stated time. The experiment seems not to have been wholly successful; a few destitute peasants were first sent out to improve these lands, many of whom were prematurely swept away by the diseases attending the improvement of a new country and a change of climate.

It was a fatal error that the plantations had not been established nearer together for mutual protection. As Marbois remarks, the colonists feeling free from restraint settled wherever fancy or hope conducted them, indifferent even to the sanction of a grant to secure their possessions; they scattered themselves among the natives, and taking the Indian women for wives, were cordially received, and by right incorporated into the tribe.

In June, , De la Housaye and Scouvion, with their followers, eighty-two in number, settled upon the Yazoo. Of the emigrants of , three hundred were destined for Natchez, and three hundred and ninety for the Yazoo. Three hundred colonists arrived in , for the lands of Madame de Chaumont, at Pascagoula. War having broken out between France and Spain the same year, the attention of the colonists was mainly directed to attacks upon the Spanish possessions.

Pensacola was taken without resistance, but was surrendered again in August to as force sent from Havana to retake it. Three ships of the line arriving on the 1st of September, the place was again taken by the French with the Spanish shipping and eighteen hundred prisoners.

In the summer and fall of , Beaumanoir brought over sixty settlers to the grant on the St. A change in the seat of government being again determined upon by the directors of the company's concerns, in opposition to the views of Bienville and Hubert, the Bay of Baluxi was chosen for that purpose; a detachment of troops was sent to the western shore of the bay to erect houses and barracks, and the place thenceforth became known as New Baluxi.

The privileges and powers of the Mississippi Company had been greatly enlarged by the acquisition of the possessions and effects of the China and India Companies, which were dissolved; and from that time it assumed the style and became known as the Company of the Indies. Although a peremptory order had been given for the removal to Baluxi, both Bienville and Hubert were opposed to it; the former thought New Orleans was the most eligible site, and the latter went to France to induce the directors of the company to decide in favor of Natchez, near which, on the St.

Catharine's, he had an immense grant with a large plantation and considerable improvements. He was unsuccessful in his mission, and died a few days after his arrival. Finally, after considerable delay and opposition in the summer of , the order of council was executed, and Bienville and his staff removed to Baluxi.

Large additions to the colony had been made the previous year, chiefly of Germans, and negroes from the coast of Africa. On the 4th of June, , a company's ship, commanded by the Chevalier d'Arensbourg, brought over two hundred and fifty Germans. This intelligence was received with great dismay, and an apprehension was felt that the affairs of the colony, if not wholly neglected, would be prosecuted with less vigor; an apprehension soon realized to some extent in the failure of supplies.

To provide against impending famine, the troops were distributed in small detachments on Pearl River and Pascagoula, among the Indian tribes, to procure subsistence by hunting and fishing. Exasperated by hunger and distress, some of these mutinied, and attempted to reach the English settlements in Carolina.

The Indians were sent in pursuit, and all of them were captured or slain. The arrival of a ship in September afforded some relief, and it was learned that the Regent, after the failure and flight of Law, had placed the affairs of the company under the direction of three Commissioners. At that time, according to his account, the company had a warehouse, in charge of Seur Le Noir, at the latter place; the appearance of the country he describes as very agreeable, extensive meadows and handsome clumps of trees presenting themselves on every side, after surmounting the hill at the landing-place.

Fort Rosalie is spoken of by him as a 7kind of redoubt inclosed with a single palisade. The great village of the Natchez was situated near the St. Catharine, a few miles from the river, and about midway between the two French grants which formed a triangle with the fort, being distant from the latter and each other about one league; the St. Maloes grant being the lowermost on the creek, which discharged itself into the Mississippi about three leagues below.

The Tioux, allies of the Natchez, had a village in the neiglhborhood. Charlevoix regarded the country about Natchez the finest and most fertile in all Louisiana. In January, , Laharpe, on his way to the Arkansas, touched at Natchez, and found Fort Rosalie in a state of ruinous decay. Maneval, who commanded it, having only eighteen soldiers. Ascending the Yazoo River at the distance of nine miles from the mouth, he reached the settlement called Fort St.

Peter, commanded by De Grave. According to his statement, there was not more than thirty acres of arable land surrounding the fort, which was hemmed in by stony hills. The site of this fort was at the place now known as Hayne's Bluff, where the limestone is seen cropping out of the base of the hills. A group of mounds, one of them of considerable size, and about thirty feet high, is situated near the spot.

At that period, the Mississippi still flowed through what is now known as Old River; the cut-off, or present channel of the river, according to Charlevoix, having been recently formed, was not passable for boats, except at a high stage of water. In May, the copper coinage provided for the colony arrived at Baluxi. It was ordered to be used in payment of the troops, and was made a lawful tender in the company's stores. Specimens of this coin have been found at St.

State, formerly occupied by the French. An earthen vessel of Indian fabric containing several pieces of it was dug, some years since, from an Indian mound near the mouth of Pearl River. One of these coins, found on the reputed site of the governor's quarters in New Baluxi, is preserved in the State cabinet; and a similar one, from the mound, in the cabinet of the State university.

These coins bear date in and They bear on the face the cipher of Louis, the French monarch, surmounted by a crown, and surrounded by the legend, " Sit nomen domini benedictum. In September, a destructive tornado desolated the Province, prostrating many houses in New Orleans, and extending to Baluxi and Natchez; the crops were destroyed, and the inhabitants were menaced with impending dearth. An unexpected crop of rice, however, springing from the seed scattered by the hurricane, promised some relief.

The Indian tribes were becoming more open in their hostilities. In , a predatory band of Chickasaws killed a sergeant belonging to the garrison at St. Peter's, and his wife. Guenot, the director of the grant on St.

Catharine, was fired upon and wounded; some negroes were shot; two planters were taken and their heads cut off, and a number of cattle and horses were stolen. When Dustine, an officer of the garrison, arrived at New Orleans, bringing this intelligence, two suns of the Natchez were on a visit to Bienville. No punishment was inflicted upon the offending Indians; but these chiefs were dismissed with presents, under a pledge to put a stop to these outrages.

In consideration of the spiritual wants of the province, a number of Jesuits, and monks of other orders, as well as nuns, were introduced into the colony, and liberally provided for by the company, and curates were also provided for the missions. For several years, great distress was felt in the colony, growing out of the failure of Law's Scheme, and the attempts of the French government to regulate the currency, and to palliate the consequent embarrassment by the alteration of the value of money.

The colony, notwithstanding, had made rapid strides since it passed under the charge of the company. The' military force had increased to eight hundred men. In , Bienville was called to France to answer to charges preferred against him. Notwithstanding his able defence, he was removed, and Perrier was appointed in his stead on the 9th of August, Of the Indian tribes occupying the country at the period of its settlement by the French, the Choctaws, Chickasaws, and the Natchez were the most numerous.

There were many others, however, though too feeble and insignificant to merit more than a passing notice. Among these were the Baluxis and Pascagoulas to the south. Of the larger tribes, the Choctaws were by far the most numerous and powerful. They owned fifty iunportant villages, and could assemble twenty thousand warriors. They were first attached to the French, who managed, by their diplomacy and presents, to retain, throughout, a large majority of them in their interest.

The Chickasaws are described as a turbulent, warlike, and ferocious race; from their intercourse and trade with the English of Carolina, they espoused their interest, and were readily engaged in hostilities towards the French, and were consequently embroiled in continual warfare with the Choctaws. The Natchez, by far the most enlightened and furthest removed from barbarism, were rapidly declining from the condition of a numerous and once powerful tribe.

The institution of human sacrifices engrafted into their theology was the most efficient cause of their rapid course towards extinction. They were pacific in their disposition; but the French, by their harshness and encroachments upon their rights, forfeited their friendship and provoked their deadly hostility. It was the policy of Bienville, and most of the other governors, for the security of the colony from the united hostilities of the Indians, against which it could not have existed, to encourage the feuds among themselves.

This was done chiefly by keeping the traffic with them in their own hands, to the exclusion of the English traders of the Carolinas, and by supplying them with goods suited to their wants. Perrier, as the successor of Bienville, proved more harsh and less politic in his intercourse with his Indian neighbors; and when, from the failure of the necessary remittances from France, it became impossible to supply all the wants of their red allies, and to make them the customary presents, a considerable faction of the Choc.

The commandant at Natchez under Perrier, an officer named Chepar, was a man of intemperate habits, and of overweening vanity and self-importance. Professing an utter contempt for the Natchez, his conduct towards them was severe and exacting. On a beautiful and elevated plain on the western margin of Second Creek, about ten miles from Fort Rosalie, was situated the "Whiteapple Village. The land embracing this favorite village of the Natchez was coveted by Chepar.

The Natchez could not bring themselves to submit to this new act of aggression. But their remonstrances were unheard, nor was the offer of other lands as an equivalent embraced. They were constrained, therefore, to feign a reluctant acquiescence in the demand.

The suns and chieftains of the different villages held a secret council, and, resolving against submission, determined themselves to become the principals, instead of auxiliaries in the conspiracy against the French. Accordingly, they set to work to secure the co-operation of other tribes hostile to the French, and to destroy the whole settlement. The necessary messengers were dispatched, each provided with a bundle of sticks of equal numbers, one of which was to be withdrawn daily, to insure a concert of action between the allies, the attack to be made on the day that the last stick was removed.

This conspiracy was designed to be kept a profound secret among the chiefs, and especially from the women, some of whom were known to be too well affected to their French neighbors. That some secret and momentous measure was on foot was soon divined by one of the most shrewd and observant of the female suns, who, severely upbraiding her son in private for his want of confidence in her, artfully drew from him the details of the plot, which she lost no time in imparting to an officer of the garrison; but her warning was unheeded.

Not satisfied with this, the female sun, having in consideration of her rank access to the fane in which the bundle of sticks for her village was kept, secretly withdrew one or two of them at a time, trusting thus, by precipitating the attack by the Natchez before the arrival of the confederates, to afford the French a further chance of escape. Deceived by this artifice, and tempted also by the arrival of some boats laden with merchandise just landed from New Orleans, on the morning of the 29th of November, , before the arrival of the day first appointed, a simultaneous attack upon the garrison, town, and different plantations was made, a shot fired upon the boats, by a party who had secretly descended the hill for that purpose, being the concerted signal.

So well was the attack planned that, in less than three hours, upwards of two hundred Frenchmen were massacred, two only, a carpenter and tailor, being spared. Ninetytwo women and one hundred and fifty-five children, and all the negroes, were captured. The usual atrocities practised by savages ensued; the fort, houses, and boats were pillaged; and the liquor obtained furnished the means of a long-continued scene of carousal and debauchery.

A few only escaped, and succeeded in reaching New Orleans, bearing the first intelligence of this sad disaster. The first of these who arrived was Richard, followed shortly after by Couillard, and a few others.

Among the principal persons who fell were the Kollys, father and son, who had just arrived to take possession of the grant of Hubert on the St. Catharine's, which they had purchased. One house only, that of Laloire, the principal agent of the company at the post, made any defence. The two survivors escaped under cover of the night.

Laloire himself, who chanced to be on horseback when the attack commenced, defended himself bravely, and killed four Indians before he fell; these, with eight others killed from his house, twelve in all, constituted the entire loss of the Natchez. As to Chepar, he was held in such contempt and abhorrence, that death by the hands of a warrior was deemed too honorable for him, and at the conclusion of the massacre he was dragged from the garden to which he had fled, and beat to death with clubs by the most degraded of the Natchez race.

The destruction of the fort at Natchez being complete, and the habitations of the French reduced to ashes, some of the Yazoo tribe who were present at the massacre, accompanied by a party of Natchez, proceeded to the settlement on the Yazoo.

The fort was garrisoned by only twenty men, and the commander, Du Codier, having already perished at Natchez, where he chanced to be on a visit at the time of the massacre, was easily surprised, and the soldiers and the few families settled near it were put to death. Thus the French settlement on the Yazoo was entirely destroyed about the 1st of January, It has been charged that the Choctaws were to have aided in this massacre, and to have made a simultaneous attack upon New Orleans; and that, in consequence of the derangement of all their plans, and their disappointment in not sharing the plunder, by the premature attack made by the Natchez, they determined to avenge themselves by the destruction of that people.

No sooner had they learned that the Natchez threatened to put to death the women and children that had been captured, than they assembled a considerable force, headed by Leseur, a Frenchman, and attacked the Natchez on the 27th of January, whilst revelling on the banks of the St. Catharine, killed many of them, rescued the carpenter and tailor, and upwards of fifty French women and children, recovering at the same time about one hundred of the negroes.

In this attack fell the chief who had instigated the Natchez massacre. Perrier, the governor, who was assembling a force at Tunica to march against the Natchez, was less prompt in his movements. The Choctaws had marched a great distance by land, and were compelled to wait for many days for the arrival of the French, with whom they were to co-operate; and it was not until the fourteenth that Loubois, the French commander, after fruitless parleyings, had posted his artillery, and made his arrangements for an attack upon the forts in which the Natchez had entrenched themselves.

The guns of the French were mounted on the mound on which stood the great temple, and commanded the forts of the Indians; they were, however, only four pounders, hardly fit for service, and so badly managed that they made little impression. The Indians opposed three pieces, which were still more clumsily handled. More than ten days were consumed in this siege.

On the night of the 18th, eluding the vigilance of the besiegers, or, as some assert, with the connzivance of the French! Thus, with the escape of the Natchez, ended this expedition, so little creditable to the French arms, in which the rescue of the captured women and children, and whatever else of success attended it, were owing mainly to their Choctaw confederates. The women and children thus rescued were sent down the river to New Orleans, and most of them were eventually settled on concessions of land made to them at Point Coupie.

The country being thus abandoned, the French commenced the erection of a brick fort, the command of which, with a garrison of one hundred men, was given to the Baron de Cresnay. Another expedition was set on foot, at the head of which Perrier placed himself, and in January, , having discovered the place of retreat, and the fortified camp of the Natchez, near the junction of the Washita and Tensas, eventually succeeded in capturing forty-five men and four hundred and fifty women and children; the others escaped.

The prisoners captured by Perrier, including two suns and a princess, were taken to New Orleans, transported to San Domingo, and sold into slavery. Those who escaped from Perrier were headed by the Chief of the Flour, who led such of his tribe as he could collect against St.

Deyns at the Post of Natchitoches, whom he attacked with a force of about two hundred warriors; but he was repulsed. Pursuing his advantage, St. Denys, at the head of his small force, a few Spaniards and an inconsiderable number of Natchitoches Indians, sallied out, forced the entrenched camp of the Natchez, killed ninety-two of them, including all of their chiefs, and put the rest to flight. Thus St. Denys, with a very inconsiderable force, inflicted upon the Natchez the most fatal blow they had yet received.

The survivors of this fated race were now scattered among the Washitas and other small tribes; but most of them sought an asylum among the Chickasaws, with whom they incorporated themselves. They continued for several years, in conjunction with the latter tribe, to attack and harass the French on all favorable occasions, and still numbered two hundred warriors. When informed of these disasters, the company of the Indies decided that it was impracticable to sustain any longer so profitless and expensive a colony, and the directors proposed to surrender to the king the charter, the obligations of which it was thought would involve it in ruin.

After much negotiation, the retrocession was accepted, the French government resumed the administration of the colony, and on the 15th of November, these Indian families at Cape Francois, he replied that he was not aware of any other course to adopt than to order their sale or to send them back to Louisiana. They were thereupon ordered to be sold. The hostile disposition of the Indians, which had been so disastrous, and which seemed to be extending to all the tribes, was attributed in a great degree to the harshness of Perrier; and the return of Bienville was urged under the belief that his mildness and humanity would conciliate the Indians, with many of whom he had ever been a favorite, and possessed great influence.

Accordingly, under the new organization of the colony, Bienville was reappointed governor in , and on his arrival, which was hailed with much joy by the colonists, Perrier returned to France. From this period until near the close of the French rule, th6 country embraced in the limits of the State was little more than the theatre of Indian hostilities and warfare.

The Natchez and the Yazoos, who had taken refuge among the Chickasaws, resumed their predatory war upon the remote settlements of the colony, in which the Chickasaws frequently united with them, and intercepted or obstructed all communication by the way of the Mississippi.

Bienville, therefore, sent an officer to the Chickasaws to demand that the Natchez should be given up. This being refused, he commenced the preparation of an expedition against them. The party of Leblanc, although attacked by the enemy near the Yazoo River, reached its destination.

Another officer was sent among the Choctaws, and by the aid of liberal presents engaged the chiefs to unite their warriors with the force Bienville proposed to lead from New Orleans. The Chevalier d'Artaguette had distinguished himself in the war with the Natchez, and had subsequently been placed in command of the Fort at Natchez.

In obedience to his orders, with such forces as he could assemble, he repaired to the place of rendezvous on the 9th of May, the day previous to that on which he was directed to arrive. He encamped in sight of the enemy until the 20th, when he was no longer able to restrain his auxiliaries, who determined to fight or withdraw. Thus situated, he embraced the first alternative, and with an impetuous charge drove the enemy from the fort before which he was encamped, and the village it protected; the second fort was carried with equal gallantry; and he was in full pursuit of the foe, retreating to their third and last entrenchment when, unfortunately, he fell under repeated wounds.

His Indian confederates now basely deserted him, and fled in all directions. Forty-eight soldiers, all he could bring with him, and Father Senac, his chaplain, stood bravely by in defence of their prostrate leader; but they were too few to resist the overwhelming force by which they were assailed.

Overpowered by numbers, many of them were captured and led prisoners, with their wounded. And where was Bienville and his army in the mean time? John with thirty boats on the 4th of March, and did not arrive at the fort on' the Tombigbee until the 20th of April. On the 4th of May he reached his landing-place within twenty-seven miles only of the nearest Chickasaw village; here the last detachment of his Choctaw auxiliaries joined him, amounting in the whole now to twelve hundred warriors.

Here they loitered, erecting houses and stores, within a day's march of the enemy, for more than twenty days. The British flag was flying, and several Englishmen were observed in the fort, which was surrounded by thick palisades pierced with loopholes for firing through; and within, the Indians were further protected by trenches, from which they could securely fire without exposing themselves to the shot of their assailants.

Since the settlement of the country by the present inhabitants, two small field-pieces and a box of bullets have been here recovered from the river, into which they were probably thrown on the retreat of the French army. Against this stronghold the force of Bienville was therefore led, powerless to inflict damage on an enemy thus protected, whilst the assailants were exposed and galled by an incessant shower of balls, which, however, was sustained for several hours, when, several of the best officers being killed or disabled, a retreat was ordered, after a loss of thirty-two killed and sixty-one wounded.

The French did not renew the attack. The following day the Choctaws had some desultory skirmishing with the enemy, and on the 29th the army commenced its retreat for the landing-place, where it arrived on the third day with the wounded. After distributing the remainder of his goods to his Choctaw allies, Bienville re-embarked his troops, floated down the river, and returned to New Orleans; thus terminating a most disastrous expedition, reflecting the deepest disgrace upon the French arms, the prowess of which was lowered immeasurably in the estimation of the savage foe.

In the mean time, the gallant but unfortunate D'Artaguette, suffering with his wounds, had been kept a prisoner, with his captured companions, under the belief that their ransom would secure favorable terms from Bienville, of whose imposing force, on its approach, they were in great dread. When it was known that he had been repulsed, and had ingloriously withdrawn, these gallant men were brought out into the plain, and D'Artaguette, Father Senac, and fifteen others were burned alive.

A sergeant of D'Artaguette's party succeeded in obtaining his liberty, reached New Orleans on the 1st of July, and made known the fate of his gallant commander. Gayerre gives a somewhat different version. According to his account, the force of D'Artaguette was much greater than represented; and his attack, although sufficiently gallant, was less successful.

He had also been apprised by a messenger of the delay of Bienville, and the cause. The battle of Ackia was so called from the town of that name, on which the attack was made. It was situated, among several other villages, in a beautiful prairie of about six miles in extent, probably near the site of De Soto's encampment of The French officers were also deserted by the larger portion of their men, whom it was impossible to force into battle. Feats of daring heroism were performed by the officers and a few of the men, many of the former being killed or wounded.

It was not until after the British sent McIntosh among them as agent, that they were induced to leave their towns and form separate settlements. To effect this dispersion, considered essential to the welfare of the nation, the agent established himself at a place near Tocshish, in the same county, represented in the early maps of the country as McIntoshville.

It is due to the reputation of Bienville to say that he alleged, in defence of his retreat, that he had reason to apprehend the desertion of the Choctawrs, and could place no reliance upon the cowardly vagabonds who had been sent him as soldiers, very few of whom were five feet in height, and many of them under that stature. A second expedition against the unsubdued Chickasaws was recommended to the French government by Bienville, to proceed up the Mississippi, instead of by the more direct and truly less objectionable route up the Tombigbee, formerly pursued, to be undertaken when the proper force, and an armament suited to the object, could be furnished.

The plan was approved, and, after considerable delay, Bienville was supplied with artillery, arms, ammunition, and provisions, and seven hundred men. With these was M. With few exceptions, preferred spawning habitat consists of gravel-bedded stream reaches with moderate depth and current cm deep and cm per second, respectively; Quinn , pg.

Females excavate a nest redd in the gravel to receive the eggs, which are fertilized by one or more competing males as they are released and subsequently buried by the female. The seasonality of spawning and incubation is roughly the same for all species of Pacific salmon, although the timing can vary somewhat by species, population, and region.

In general, salmon spawn during summer or early fall and the fry emerge from the spawning gravel the following spring. While in the gravel, the embryos develop within their eggs and then hatch into fry that continue to subsist on yolk sacs. After emerging from the gravel, basic life history patterns of the five species differ in notable ways.

Species-specific life history and ecology Sockeye salmon Sockeye salmon originate from river systems along the North American and Asian shores of the North Pacific and Bering Sea, roughly from the latitude of the Sacramento River to that of Kotzebue Sound. The largest North American populations occur between the Columbia and Kuskokwim rivers Burgner , pg. Spawning sockeye are readily identified by their striking red bodies with green heads and tails; males additionally develop a large hump in front of the dorsal fin.

Sockeye are unique among salmon in that most stocks rely on lakes as the primary freshwater rearing habitat. Some sockeye spawn within the nursery lake where their young will rear. Others spawn in nearby stream reaches, and their fry migrate to the nursery lake after emerging from spawning redds. Sockeye are by far the most abundant salmon species in the Bristol Bay region Salomone et al.

Riverine sockeye populations spawn and rear throughout the Nushagak River basin Figure 2. Juveniles in Bristol Bay systems rear for one or two years in their nursery lakes West et al. Fish then typically spend two or three years at sea West et al. At sea, sockeye salmon feed on a range of invertebrates, small fish, and squid Burgner , pg Sockeye salmon distribution in the Nushagak and Kvichak watersheds. There are more than a thousand North American spawning populations and a much smaller number in Asia.

These populations tend to be relatively small, however, making Chinook the rarest of North America's Pacific salmon species Healey , pg. They are also the largest of the Pacific salmon; at least one specimen over 60 kg has been reported, but most weigh less than 23 kg Mecklenburg etal. Chinook salmon have two different behavioral forms.

The "stream type" form is predominant in Bristol Bay, as well as other areas of northern North America, Asia, and the headwaters of Pacific Northwest rivers Healey , pg. These fish spend one or more years as juveniles in fresh water, range widely at sea, and return to spawning streams during spring or summer.

In fresh water, juvenile Chinook tend to occupy flowing water and feed on aquatic insects. At sea, Chinook are generally pisciverous Brodeur and feed higher on the food chain than other salmon species Satterfield and Finney Chinook spawn and rear throughout the Nushagak River basin and in many tributaries of the Kvichak River Figure 3.

Some life history data are available from adults returning to the Nushagak River, Bristol Bay's largest Chinook salmon run. Chinook salmon individuals in recent Bristol Bay commercial catches have averaged Chinook salmon distribution in the Nushagak and Kvichak watersheds. Bristol Bay's rainbow trout are of the coastal variety sensu Behnke , pg.

While classified in the same genus as the Pacific salmon, there are some key differences. Foremost, rainbow trout are not genetically programmed to die after spawning, making iteroparity i. Also, most coastal drainages support populations of both resident and anadromous i. Finally, rainbow trout spawn in the spring, as opposed to summer or early fall, although their spawning habitat and behavior is otherwise generally similar to that of salmon.

Bristol Bay rainbow trout tend to mature slowly and grow to relatively large size. Bristol Bay trout utilize complex and varying migratory patterns that allow them to capitalize on different stream and lake habitats for feeding, spawning, and wintering. From there, they appear to utilize a variety of habitats, as some tagged individuals have been recovered in other Iliamna Lake tributaries and in the Newhalen and Kvichak Rivers Russell , pg.

In the Alagnak River watershed, a number of rainbow trout life history types have been identified, each with their own habitat use and seasonal migratory patterns Meka et al. These consist of lake, lake-river, and river residents, the latter of which range from non-migratory to highly migratory Meka et al. Individuals comprising each of these life history types migrate in order to spend the summer in areas with abundant spawning salmon Meka et al.

Eggs from spawning salmon are a major food item for Bristol Bay trout and are likely responsible for much of the growth attained by these fish. Upon the arrival of spawning salmon in the Wood River basin, rainbow trout shifted from consuming aquatic insects to primarily salmon eggs for a 5-fold increase in ration and energy intake Scheuerell et al.

With this rate of intake, a bioenergetics model predicts a g trout to gain 83 g in 76 days; without the salmon-derived subsidy, the same fish was predicted to lose five g Scheuerell et al. Rainbow trout in Lower Talarik Creek were significantly fatter i. Coho salmon Coho salmon are native to coastal drainages in western North America and eastern Asia, approximately from the latitude of the Sacramento River to that of Point Hope Sandercock , pg.

Coho salmon occur in relatively small populations, and are second only to Chinook salmon in rarity. Most Alaskan coho salmon populations tend to spend two years in fresh water and one year at sea Sandercock , pg. Coho salmon individuals in recent Bristol Bay commercial catches have averaged 6.

At sea, coho salmon consume a mix of fish and invertebrates Brodeur , pg. Their trophic position is intermediate for Pacific salmon; Chinook salmon consume more fish while sockeye, pink, and chum salmon eat more zooplankton and squid Satterfield and Finney In fresh water, coho salmon feed primarily on aquatic insects, although salmon eggs and flesh can be important nutritional subsidies Heintz et al.

In press. They utilize a wide range of lotic and lentic freshwater habitats, including stream channels, off- channel sloughs and alcoves, beaver ponds, and lakes. Coho distribute widely into headwater streams, where they are often the only salmon species present Woody and O'Neal , King et al. Production of juvenile coho is often limited by the extent and quality of available wintering habitats Nickelson et al. Rinella, unpublished data. Pink salmon Pink salmon spawning populations occur on both sides of the North Pacific and Bering Sea, as far south as the Sacramento River and northern Japan.

Northward, small spawning populations are scattered along the North American and Asian shores of the Arctic Ocean. The most abundant Pacific salmon overall Irvine et al. They typically spawn in shallow, rocky stream reaches relatively low in the watershed and their young migrate to sea soon after emerging Heard , pg. Essentially all pink salmon breed at two years of age, and this strict two-year life cycle results in genetic isolation of odd- and even-year spawning runs, even within the same river system.

For reasons not entirely clear, large disparities between odd- and even-year run sizes occur across geographic regions and extend over many generations. An extreme example is the Fraser River, in southern British Columbia, where millions of pink salmon return during odd- numbered years, yet no fish return during even-numbered years Riddell and Beamish , pg. In Bristol Bay rivers, even-year runs dominate the returns Salomone et al. Pink salmon are the smallest of the Pacific salmon species; individuals in recent Bristol Bay commercial catches have averaged 3.

Sexually mature males become highly laterally compressed and develop a massive dorsal hump, hence the common name "humpy. Scattered spawning populations also occur on the Asian and North American shores of the Arctic Ocean. Populations tend to be relatively large, and chum salmon are the third most abundant species, behind pink and sockeye salmon. Chum salmon, like pink salmon, migrate to sea soon after emerging from spawning gravel.

Across their range, the vast majority spends two to four years at sea Salo , pg. At sea, chum salmon consume a range of invertebrates and fishes, and gelatinous material is commonly found in stomachs leading to speculation that jellyfish may be a common prey item Brodeur , pg. Individuals in recent Bristol Bay commercial catches have averaged 6. Commercial fishing interests were among the original supporters of the purchase of Alaska from Russia in King , pg. The first canneries were established eleven years later, and by the s salmon surpassed mining as Alaska's major industry as Alaska became the world's principal salmon producer Ringsmuth , pg.

In the early years, fish packing companies essentially had a monopoly on the harvest of salmon. Packers in Bristol Bay and elsewhere built industrial fish traps, constructed of wood pilings and wire fencing with long arms that guided schools of migrating salmon into holding pens King , pg. In Bristol Bay, packing interests also upheld a federal ban on fishing with power boats until Ostensibly a conservation measure, this law served to protect obsolete cannery-owned sailboat fleets by excluding independent Alaska-based fishermen who largely used power boats by this time Troll , pg.

Salmon harvest peaked in then declined steadily for many years, leading to a federal disaster declaration in the s King , pg. A lack of scientific management, poor federal oversight, excessive harvest during World War II, and natural changes in ocean conditions contributed to the decline. Declining salmon runs, along with Alaskans' desire for more control over their fisheries, was a significant factor in the drive toward Statehood Augerot , King , pg. In , Alaskans began to develop a state constitution that included provisions intended to preserve Alaska's fisheries and, unique among state constitutions, to guarantee equal access to fish and game for all residents.

Alaska became a state in , the year that marked the lowest salmon harvest since King , pg. Statehood was a turning point for Alaska's salmon fisheries, with the end of federal management, fish traps, and undue control of the resource by the canning industry. With the mandate for equal access came decentralization of 10 the fishing industry, and thousands of individual fishermen began harvesting salmon for market to the canneries Ringsmuth , pg.

Inventorying fish stocks, understanding basic ecology, and improving run strength forecasting were central research goals. Of particular importance was the development and application of methods for counting salmon runs in spawning streams, which allowed the establishment of escapement goals and management based on scientific principles of sustained yield.

The latter, funded largely by the salmon processing industry, began researching factors controlling sockeye salmon production in While the scope of their investigations has expanded over the years, sockeye monitoring is still a focus and represents the world's longest-running program for monitoring salmon and their habitats. Over time, a number of state and federal policy changes have affected Bristol Bay salmon fisheries.

A constitutional amendment set the stage for a bill that limited participation in Alaska commercial salmon fisheries. This legislation, designed to curb the expanding commercial fishery, set an optimum number of permits for each fishery, which were then issued by the State based on an individual's fishing history.

Permits are owned by the individual fisherman and are transferable, making them a limited and valuable asset King , pg. The Fishery Conservation and Management Act of , commonly known as the Magnuson-Stevens Act, was introduced to Congress by the late senator Ted Stevens as a means to curtail high seas salmon fishing.

In response to intensive Japanese gill netting in the western Aleutians and Bering Sea since , this legislation extended America's jurisdiction from 12 to miles 19 to km offshore. This ensured that salmon produced in Alaskan rivers would be harvested and processed locally and gave Alaska's fishery managers much more control in deciding when and where salmon are harvested.

The former established a comprehensive policy for the regulation and management of sustainable fisheries and the latter defined procedures for establishing and updating salmon escapement, including a process for public review of allocation disputes associated with escapement goals The Alaska Department of Fish and Game is responsible for managing fisheries under the sustained yield principle. The Board of Fisheries consists of seven citizens, appointed by the governor and confirmed by the legislature, that serve three-year terms.

Eighty-one advisory committees, whose members are elected in local communities around the state, provide local input. While regulations and management plans provide the framework for fisheries regulation, local fisheries managers are ultimately responsible for their execution. They are delegated authority to make "emergency orders," in-season changes to fishing regulations, which allow rapid adjustments to changing conditions, often with very short notice.

Managers use them to provide additional protection to fish stocks when conservation concerns 11 arise and to liberalize harvest when surplus fish are available. Alaska's management of its salmon fishery has proven successful; it was the second fishery in the world to be certified as well managed by the Marine Stewardship Council Hilborn and is regarded as a model of sustainability Hilborn et al.

Current management of commercial salmon fisheries While all five species of Pacific Salmon are harvested in Bristol Bay, sockeye salmon dominate the runs and harvest by a huge margin Table 1. Salmon return predominately to nine major river systems, located on the eastern and northern sides of the Bay, and are harvested in five fishing districts in close proximity to the river mouths that allow managers to regulate harvest individually for the various river systems Figure 1.

The Naknek-Kvichak district includes those two rivers as well as the Alagnak. The Egegik, Ugashik, and Togiak districts include the rivers for which they are named. Table 1. Mean harvest by species and fishing district, Fishing is conducted with drift or set gillnets. Set gillnets have a maximum length of fathoms m and are fished from boats no longer than of 32 ft. Set gillnets are fished from beaches, often with the aid of an open skiff, and have a maximum length of 50 fathoms 91 m.

The management of the Bristol Bay sockeye salmon fishery is focused on allowing an adequate number of spawners to reach each river system while maximizing harvest in the commercial fishery Salomone et al. This balancing act is achieved through the establishment of escapement goals which represent the optimum range of spawners for a given river system. Escapement goals are established using a time series of spawner counts where a spawning run of a given size i.

Established stock-recruit models Ricker , Beverton and 12 Holt are then used to estimate the stock size that results in the largest number of recruits, or the maximum sustained yield Baker et al. In theory, spawning runs that are too small or large can result in reduced recruitment. With the former, too few eggs are deposited. With the latter, superimposition of spawning redds can diminish egg viability and competition in nursery lakes can reduce growth and survival.

Once escapement goals are set, the timing and duration of commercial fishery openings are then adjusted during the fishing season i. Escapement goals are periodically reviewed and updated based on regulatory policies, specifically, the Policy for the Management of Sustainable Salmon Fisheries and the Policy for Statewide Salmon Escapement Goals.

Each of Bristol Bay's nine major river systems has an escapement goal for sockeye salmon Table 2 , and in-season management of the commercial fishery is used to keep escapement in line with the goals. Management responsibility is divided among three managers: one for the Naknek, Kvichak, and Alagnak rivers; one for the Nushagak, Wood, Igushik, and Togiak rivers; and one for the Ugashik and Egegik rivers.

Fishery openings are based on information from a number of sources, including preseason forecasts, the test fishery at Port Moller, the early performance of the commercial fishery, and in-river escapement monitoring. Table 2. Bristol Bay escapement goal ranges for sockeye salmon. River Kvichak Alagnak Naknek Egegik Ugashik Wood River Igushik Nushagak-Mulchatna Togiak Escapement range thousands 2,, minimum , , , , Preseason forecasts are the expected returns of the dominant age classes in a given river system, and they are based on the number of spawning adults that produced each age class.

In the Port Moller test fishery, gill netting at standardized locations provides a daily index of the overall number of fish entering Bristol Bay Flynn and Hilborn , with approximately seven days' lead before they enter the commercial fishing districts. Genetic samples from the test fishery are analyzed within four days Dann et al. As salmon move upstream, escapement is monitored with counting towers on each of the major rivers, except the Nushagak where a sonar system is used.

Counting towers are elevated platforms along small to medium-sized m wide , clear rivers from which migrating salmon are visually counted Woody Since tower and sonar monitoring occurs well upstream of the 13 commercial fishery, all information regarding the performance of the fishery must be analyzed on a continual basis to ensure escapement levels will be met Clark , pg.

The fishery is typically opened on a schedule during the early part of the season, during which time the frequency and duration of openings are primarily based on preseason forecasts and management is conservative. As the fishing season progresses and more information becomes available, managers make constant adjustments to fishing time and area. If the escapement goal is exceeded at a given monitoring station, the fishery is opened longer and more frequently.

If the escapement goal is not reached, the fishery is closed. Fishing time is opened and closed using emergency orders, and fishermen often learn of changes only a few hours before they go into effect. Since the bulk of the sockeye salmon harvest occurs during a short timeframe - from the last week of June until the middle of July - this short warning system is needed to maximize fishing time while ensuring that escapement levels are met.

Migrating fish move quickly through the fishing districts, and delaying an opener by one day during the peak of the migration can forego the harvest of a million salmon. This is a significant loss of revenue to individual fishermen, and compounded by the missed revenue of workers, processors, and marketers Clark , pg.

The fishery will periodically close de facto during the peak of the season when catch rates exceed processing capacity and processors stop buying fish. This lack of buyers can also curtail salmon harvest early and late in the season when numbers offish do not warrant keeping processing facilities operational. In-season management is also used to help meet an escapement goal for Chinook salmon on the Nushagak River Table 3 , where escapement is monitored by sonar. There are also Chinook salmon goals for the Togiak, Alagnak, Naknek, and Egegik rivers and a chum salmon goal for the Nushagak River Table 3 , but in-season management is not used to help attain these goals.

Bristol Bay salmon fisheries are regarded as a management success Hilborn et al. Bristol Bay escapement goal ranges for Chinook and chum salmon. River Togiak Nushagak Nushagak Alagnak Naknek Egegik Species Chinook Chinook chum Chinook Chinook Chinook Escapement goal 9, minimum 40,, , minimum 2, minimum 5, minimum minimum 14 Description of sport fisheries The sport fisheries in Bristol Bay's river systems are regarded as world class. The area has been acclaimed for its sport fisheries since the s.

For example, Fly Rod and Reel Williams says "No place on earth is wilder or more beautiful or offers finer salmonid fishing. Large numbers of salmon and trout are caught in Bristol Bay's sport fisheries each year see below , but the area is best known for its rainbow trout fishing. Fish Alaska magazine calls the Iliamna system "One of the greatest trophy trout fisheries in the world Unlike commercial fisheries, whose salient features tend to be readily quantifiable e.

Despite the potential to catch high numbers of sizeable fish, Bristol Bay anglers rate aesthetic qualities as most important in selecting fishing locations. Of 11 attributes that capture different motivations and aesthetic preferences, including "catching and releasing large numbers offish" and "chance to catch large or trophy-sized fish," Alaska resident and nonresident anglers picked the same top five: "natural beauty of the area", "being in an area with few other anglers", "being in a wilderness setting", "chance to catch wild fish", and "opportunities to view wildlife" Duffield et al.

The same priorities apply for nonresident anglers across Alaska Romberg , pg. The Bristol Bay region is not linked to the State's highway system and roads connected to the major communities provide very limited access. Small aircraft with floats are the primary source of access followed by boats based out of communities and remote lodges Dye and Schwanke , pg.

A range of services are available for recreational anglers. Modest river camps, with cabins or wall tents, are a lower-budget option. Many self-guided expeditions center on multi-day raft trips that use chartered aircraft for transport to and from access points along a river. Site-specific data regarding participation, effort and harvest have been collected from sport fishing guides and businesses since Sigurdsson and Powers In addition, Table 4 shows figures for , the first year of data collection, and , a peak year.

Table 4. The number of businesses and guides operating in the Nushagak and Kvichak watersheds in , and Three local management plans guide sport fishing regulations in the Bristol Bay region in addition to several statewide plans. The Southwest Alaska Rainbow Trout Management Plan instated conservative trout management uniformly throughout the region, replacing the fragmentary restrictions that had been established over the previous decades.

The Division of Sport Fish uses the annual Statewide Harvest Survey, mailed to randomly-selected licensed anglers, to monitor effort, catch, and harvest. Total annual sport harvest for the same period ranged from 39, to 71, fish, of which sockeye, Chinook and coho salmon comprise the majority Dye and Schwanke , pg.

Harvest rates are lower for these species than for salmon, likely due to restrictive bag limits and the popularity of catch-and-release fishing Dye and Schwanke , pgs. Chinook salmon In the Nushagak drainage, the general season runs from May 1 to July 31 for Chinook salmon, although some areas close on July 24 in order to protect spawners. The daily limit is two per day, only one of which can be over 28 inches 71 cm.

The annual limit is four fish. The Nushagak-Mulchatna King Salmon Management Plan calls for an in-river return of 75, fish with a spawning escapement of 65, fish. The guideline harvest for the sport fishery is 5, fish, although restrictions are triggered if the in-river return falls below 55, fish. In other 16 Bristol Bay drainages, the daily limit for Chinook salmon is three and the annual limit is five, although there are additional restrictions in the Wood and Naknek river drainages.

The major Chinook salmon sport fisheries in the BBMA include the Nushagak, Naknek, Togiak and Alagnak rivers and the average annual harvest is 11, fish for the period from to Sockeye salmon Sockeye salmon fishing is open year round with a daily limit of five fish.

Runs enter rivers starting in late June, peak in early July, and continue into late July or early August. The Kvichak River Drainage Sockeye Salmon Management Plan places restrictions on the sport fishery to avoid conflicts with subsistence users when the escapement falls below the minimum sustainable escapement goal of two million fish. Restrictions include actions such as reducing the daily limit for sockeye and closure of areas for sport fishing that are used by both subsistence and recreational anglers.

Sockeye are the most abundant salmon species in the BBMA. Recent annual sport harvest ranged from 8, to 23, fish Dye and Schwanke , pg. The two locations that support the largest sport harvest are the Kvichak River, near the outlet of Iliamna Lake, and the Newhalen River, just above Iliamna Lake Dye and Schwanke , pg. Other drainages that support moderate harvests of sockeye salmon include the Naknek and Alagnak rivers and the Wood River lake system Dye and Schwanke , pg.

Rainbow trout Due to their relatively small spawning populations and their popularity as a game fish, fishing regulations for rainbow trout are more restrictive than those for any other species. Special management areas were created to preserve a diversity of sport fishing opportunities: eight catch-and-release areas, six fly-fishing catch-and-release areas, and eleven areas where only single-hook artificial lures can be used Dye and Schwanke , pgs.

Only single-hook artificial lures can be used in the Kvichak River drainage, and all sport fishing is banned from April 10 through June 7 to provide protection for spawning rainbow trout. From June 8 through October 31 anglers are allowed to keep one trout per day, with the exception of a number of streams where no harvest is allowed.

From November 1 through April 9, when anglers are few, the daily limit increases to five fish although only one may be longer than 20 inches 51 cm. Rainbow trout fishing regulations are similarly restrictive in other drainages across the BBMA. The most popular rainbow trout fisheries are found in the Kvichak drainage, the Naknek drainage, portions of the Nushagak and Mulchatna drainages, and streams of the Wood River Lakes system Dye and Schwanke , pg.

Field surveys and the Statewide Harvest Survey show that harvest has decreased over the past decade but that total catch and effort have remained stable or increased Dye and Schwanke , pg. The annual BBMA-wide harvest 17 between and averaged fish, but the catch estimate over this period was nearly times greater , fish; Dye and Schwanke , pgs.

Although the fishery is widespread, approximately eighty percent of the total catch , fish was from the eastern portion of the BBMA, where the Naknek and Kvichak systems are located. The relative abundance of Pacific salmon species relates to their life histories, as those species that are not constrained by the availability of stream rearing habitat i. The highest Pacific-wide salmon harvest occurred in and totaled million fish, over million of which were pink salmon Irvine et al.

Approximately five billion juvenile salmon are released annually from hatcheries around the North Pacific Irvine et al. When combined with commercial, subsistence and sport harvest, data from escapement monitoring allows estimates of total run sizes. A recent synthesis of salmon returns for 12 regions around the North Pacific also extends back to the s, allowing comparisons of wild sockeye salmon returns between Bristol Bay and other regions for the period to Ruggerone et al.

The average global abundance of wild sockeye salmon over that period was Total returns to Bristol Bay ranged from a low of 3. Other regions that produce high abundances of wild sockeye salmon include the Kamchatka Peninsula, northern British Columbia, Cook Inlet and Kodiak Island Ruggerone et al. Relative abundance of wild sockeye salmon stocks in the North Pacific, See Bristol.

AK Other. Kamchatka W.. Kamchatka Russia N.. WA Appendix 1 for data and sources. Stocks are ordered from west to east across the North Pacific. Kamchatka E. AK ""' S. AK Pen. BC "" S. Wild sockeye salmon abundances by region in the North Pacific, See Appendix 1 for data and sources.

Each graph shows three regions organized from west to east across the North Pacific. No hatchery production has occurred in the Bristol Bay region. On average, the Kvichak River has the largest sockeye salmon run in Bristol Bay, with an average annual return of Runs exceeding 30 M fish have occurred three times in the Kvichak River: The Egegik River supports Bristol Bay's second largest run, with a mean annual return of 6.

The Nushagak and Wood rivers are smaller runs and average returns from to were 1. As noted above, returns to the Kvichak River have averaged The Fraser River system supports the world's second largest run, with an average of 8. The Kamchatka Peninsula in Russia also has rivers with large sockeye runs, but abundances for individual rivers were not readily available.

The combined runs for the western and eastern Kamchatka Peninsula averaged less than 5 M sockeye during the period from to Ruggerone et al. Sockeye salmon abundances for major rivers of the North Pacific, The top graph includes time series for the Nushagak-Wood and Kvichak-Alagnak systems from to , the Chignik River from to , and the Karluk River from to The bottom graph shows the Kenai River late run from to , the Copper River wild run from to , the Skeena River from to , and the Fraser River from to Rivers are listed in the graphs as they occur from west to east across the North Pacific.

A major driver is the Pacific decadal oscillation PDO , an inter-decadal pattern of correlated changes in sea-level pressures and sea- surface temperatures Mantua et al. The warm phase of the PDO is characterized by warmer than average winter sea surface temperatures along the western coastline of North America and increased stream flows around the Gulf of Alaska, both of which are linked to increased salmon survival Mantua et al.

There are three regime shifts documented in the recent climate record that correlate with salmon productivity: , and From to , the PDO was in a cool phase marked by low productivity for Alaskan and British Columbia sockeye salmon. For Bristol Bay stocks, this warm phase corresponded with increased marine growth and, in turn, increased abundances and numbers of recruits returning adults generated per spawner Ruggerone et al.

Bristol Bay stocks more than doubled during this warm phase and remained high until the mids, when declines in the Kvichak and other rivers reduced the overall abundance Figure 4, Ruggerone et al. Biological indicators suggest that decreased productivity associated with a cool phase began in , while climate indices point to a short-lived reversal from to , followed by a return to a warm phase Hare and Mantua Late marine growth and adult length-at-age of Bristol Bay sockeye decreased after the regime shift, potentially reducing stock productivity Ruggerone et al.

Another factor affecting sockeye salmon productivity is competition with increasing numbers of hatchery smolts released into the North Pacific. Alaska produces the most hatchery pink salmon in the world, averaging 42 M fish for the period to , followed next by Russia, with Japan dominates the production of hatchery chum salmon, with Coming in a distant second behind Japan, Southeast Alaska averaged 9.

Bristol Bay sockeye smolts that migrated to sea during even-numbered years and interacted with dominant odd-year Asian pink salmon experienced decreased growth, survival and adult abundance compared to the smolts that migrated during odd-numbered years Ruggerone et al. Additionally, Kvichak sockeye salmon productivity was negatively correlated with a running three-year mean of Kamchatka pink salmon abundances Ruggerone and Link In the freshwater environment, spawning and rearing habitats can limit sockeye salmon populations through negative density dependence.

The amount of suitable spawning habitat is fixed within a given system, so when spawning densities are high and suitable spawning sites are occupied, females will dig nests on top of existing nests, dislodging many of the previously laid eggs, or die without spawning Semenchenko , Essington et al. As such, the amount of available spawning habitat can impose an upper limit on potential fry production. In nursery lakes, juvenile growth rates decrease with rearing densities Kyle et al.

Together, these processes limit the number of recruits potentially produced by a large spawning run. Kvichak sockeye abundances follow five-year cycles that are unique amongst the nine major systems of Bristol Bay. Previous hypotheses for the cycle included natural depensatory mechanisms, such as predation, and fishing-related depensation. Since the first escapement goal was established for the Kvichak River in until the most recent change in , the escapement goals were managed to match the cycle year.

Most recently, off-cycle years had an escapement goal range of 2 to 10 M spawners, while pre-peak and peak cycle years were managed for escapement of 6 to 10 M spawners Baker et al. In , the escapement goal was changed to one goal for all years of 2 to 10 M spawners. Ruggerone and Link recently analyzed the population characteristics of Kvichak sockeye and found that the cycle is likely perpetuated by three factors: density dependence during pre-peak and peak cycle years reducing productivity in off-cycle years, higher percentage interceptions in off-cycle years biasing productivity low, and the dominance of age 2.

Kvichak salmon were shown to have high interception rates in the Egegik and Ugashik fisheries in years when the Egegik and Ugashik returns were more than double the Kvichak return, which biased the number of returning recruits during off-cycle years. They did not find any evidence of natural depensatory mechanisms, nor did they find reason to believe that the change in the escapement goal in could have had any effect on the decline in the s. Beginning in , with the spawning return of the brood year, Kvichak runs dropped to an average of 4.

Mean annual returns of sockeye salmon in Bristol Bay, , and percent of total by river system. Rivers are listed from east to west across Bristol Bay. The average number of smolts out- migrating from the Kvichak River during the years to was approximately M, which declined to an approximate average of 50 M from to Ruggerone and Link The declines were accompanied by a shift in the dominant age structure of Kvichak spawners from 2. Across the nine monitored Bristol Bay watersheds, the decrease in the percentage of 2.

The decrease in spawner length at age starting in and higher than normal sea surface temperatures in June from both may have contributed to lower reproductive potential, since smaller females produce fewer eggs. Competition with Asian pink salmon also may have played a role. Abundances of Asian pink salmon have been linked to decreased size at age of returning Bristol Bay sockeye salmon in addition to decreased abundance during even-year migrations when interactions are highest Ruggerone et al.

Abundances of Kamchatka pink salmon were high from to , the beginning of which correlates to age-1 smolts from the brood year. The three eastern Bristol Bay stocks that experienced the largest declines during the s Kvichak, Egegik and Ugashik rivers have greater overlap with Asian pink salmon stocks in their marine distribution than other stocks that did not decline significantly Ruggerone and Link , pg.

Ultimately, conditions outside of the freshwater environment likely led to the decline of Kvichak sockeye salmon. Warmer summer temperatures in both fresh water Schindler et al. Because ocean-age-two salmon interact with only one Asian pink salmon population at sea, the effects on growth and abundance are greater than for ocean-age-three salmon, which interact with both large even and small odd Asian pink salmon populations at sea and thus, have the opportunity for higher growth rates during odd years Ruggerone et al.

The decrease in spawner to smolt survival may also be related to marine conditions causing smaller length at age of returning adults and reduced reproductive success Ruggerone and Link , pg. Since , Bristol Bay returns have again totaled more than 40 million fish annually and in the Kvichak run increased to over 9. Chinook salmon The total commercial harvest of Chinook salmon in the North Pacific ranged between three and four million fish until the early 90s; recent total catches have decreased to one to two million fish Eggers et al.

Lacking escapement data for many runs, commercial harvest is a good surrogate for salmon abundance, and suggests a decline in Chinook salmon abundance in recent decades. The U. Recreational, subsistence, and aboriginal catch is significant for this salmon species and totaled approximately one million annually in Heard et al. Washington dominates hatchery production of Chinook salmon, with over one billion juveniles released annually from Heard et al.

The Columbia River historically produced the largest Chinook salmon run in the world, with peak runs spring, summer, and fall combined estimated at 3. Peak catches for the Columbia River summer-run Chinook salmon occurred at this time, until overfishing decimated the run. Fishing effort then shifted to the fall run, which suffered a similar demise in the early s. Currently, the largest runs of Chinook salmon in the world originate from three of the largest watersheds that drain to the North Pacific: the Yukon, Kuskokwim and Fraser rivers Table 6.

Total Chinook escapements to the Kuskokwim and Yukon rivers have not been quantified directly due to their large watershed area, but recent total run estimates based on mark-recapture studies put them at , and , fish, respectively Molyneaux and Brannian , pg. On the Fraser River, the average size of the spring, summer, and fall Chinook runs combined including the Harrison River for the most recent ten-year period was , fish PSC , pg.

Nushagak River Chinook average run sizes for , in comparison to other rivers across the North Pacific. Other rivers are sorted in order of decreasing run size. Watershed Nushagak R. Taku R. Copper R. Kenai R. Yukon R. Grays Harbor Chehalis R. Nehalem R. The Nushagak produces runs that are periodically at or near the world's largest Figure 8 , which is remarkable considering its relatively small watershed area Table 6.

Returns consistently number over , fish, while returns greater than , fish have occurred eleven times between and Figure 8. An especially productive six-year period from produced three returns greater than , fish Figure 8. The Harrison River is the dominant fall run stock for the Fraser River. Chinook salmon abundances by river system, The top graph shows total runs for the Yukon River Canadian stock from to , the Kuskokwim River from to , the Nushagak River from to , and the Kenai River from to The bottom graph shows total runs for the Copper River from to , the Taku River from to , the Skeena River from to , and the Fraser River from to Rivers are organized from west to east across the North Pacific.

The Nushagak Chinook stock is considered stable Heard et al. Both the Yukon and Kuskokwim Chinook were listed as stocks of yield concern in Estensen et al. The Yukon River stock is still listed but the Kuskokwim River Chinook stock was delisted as a stock of concern in , based on higher than normal returns starting in Estensen et al.

The decline in Yukon and Kuskokwim Chinook stocks that began in the late s may have resulted from the El Nino Kruse , Myers et al. The decline in Chinook stocks that persisted after the El Nino indicate that multiple ocean age classes were affected by this event Ruggerone et al. Chinook salmon hatchery production contributes to harvests in both southeast and southcentral Alaska.

There are no salmon hatcheries located in western Alaska and none of the total runs for the Alaskan rivers listed in Figure 8 or Table 6 include contributions from hatcheries Yukon, Kuskokwim, Nushagak, Kenai, Copper, and Taku rivers. Threatened and endangered salmon and conservation priorities Although it is difficult to quantify the true number of extinct salmon populations around the North Pacific, estimates for the Western United States California, Oregon, Washington and Idaho have ranged from to populations Nehlsen et al.

Chinook had the largest number of extinctions followed by coho and then either chum or sockeye Nehlsen et al. Many of the patterns of population extinction are related to time spent in fresh water: interior populations have been lost at a higher rate than coastal populations, stream-maturing Chinook and steelhead which may spend up to nine months in fresh water before spawning had higher losses than their ocean-maturing counterparts, and species that relied on fresh water for rearing Chinook, coho, and sockeye had higher rates of extinction than pink or chum salmon, which go to sea soon after emergence Gustafson et al.

No populations from Alaska are known to have gone extinct. Salmon populations in the southern extent of their range have suffered higher 32 extinction rates and are considered at higher risk than populations further to the north Brown et al. In addition to the large number of populations now extinct, there are many that are considered at risk due to declining population trends. The Columbia River basin dominated the list of at risk stocks identified by Nehlson et al.

Approximately half of the stocks evaluated were listed as high risk because they failed to replace themselves fewer than one recruit per spawner or had recent escapements below individuals. More recent analyses of the status of salmon populations in the North Pacific continue to highlight the declines in the Pacific Northwest.

A detailed assessment of salmon populations in the Columbia River basin from to showed that many are declining and this trend is heightened when hatchery fish are excluded McClure et al. A comparison between time periods reflecting both good and bad ocean productivity for Columbia River salmon populations further indicates that the declining trends are not due to the regime shift of McClure et al.

Washington, Oregon, California, and Idaho had the highest concentrations of high-risk stocks Augerot , pgs. A detailed assessment of sockeye salmon populations across the North Pacific highlights threats for this species in British Columbia Rand At the global population level, sockeye salmon are considered a species of least concern. Eighty subpopulations were identified for assessment, five of which are extinct and 26 did not have the necessary data with which to conduct a status assessment.

Of the remaining 49 subpopulations, 17 were identified as threatened and two as nearly threatened. Three key threats to sockeye salmon were identified: mixed stock fisheries that lead to high harvests of small, less productive populations; poor marine survival rates and high rates of disease in adults due to changing climatic conditions; and negative effects of enhancement activities such as hatcheries and spawning channels Rand Twenty-five subpopulations were assessed for Alaska: 10 were data deficient, 12 were of least concern including the one subpopulation identified for Bristol Bay , one subpopulation in the eastern Gulf of Alaska was listed as vulnerable four of six sites had declining trends: Bering, East Alsek, Italic, and Situk rivers , and two subpopulations in Southeast Alaska McDonald and Hugh Smith Lakes were listed as endangered.

Both were de-listed within four years after runs met escapement goals for several consecutive years following implementation of successful fishing restrictions Piston , pg. Government agencies in the United States and Canada are tasked with identifying and protecting salmon populations at risk. In the U. Salmon stocks considered for listing under ESA must meet the definition of an Evolutionary Significant Unit ESU : it must be substantially reproductively isolated from other nonspecific population units and it must represent an important component of the evolutionary legacy of the species 33 Federal Register , November 20, Current determinations for the U.

All listed ESUs occur in the western contiguous U. California, Oregon, Washington, and Idaho. On the Asian side of the Pacific, no information was found regarding listings of threatened or endangered salmon populations under a legal framework. Other assessments of Asian salmon distribution and status have relied on interviews with fishery biologists due to the scarcity of data and the dominance of hatcheries in Japanese fisheries Augerot , pg.

In California, both the building of dams that eliminated access to upstream spawning and rearing areas and destruction of coastal habitat from extensive logging were major contributors to the decline of coho salmon populations in the southern extent of their range Brown et al.

Heavy fishing pressure at the end of the 19th century followed by extensive impacts to river habitats 35 from agriculture, logging, mining, irrigation and hydroelectric dams all led to the extensive decline of Columbia River salmon by the mid 20th century Chapman , McConnaha et al. Restoration activities to help restore salmon habitat and populations in the Pacific Northwest require huge expenditures with results that are often difficult to measure due to annual variation, the time lapse between restoration action and effect on the population, and changing climate and ocean conditions GAO , pg.

Predicted outcomes from restoration rarely take into account climate change scenarios. Models developed to predict the outcome of restoration on Snohomish basin Chinook salmon habitat showed that increased temperatures resulting from climate change changed snow to rain in high elevation watersheds and affected three hydrologic parameters that decreased fish populations: higher flows during egg incubation, lower flows during spawning, and increased temperatures during pre-spawning Battin et al.

Often used as mitigation for lost habitat, salmon hatcheries have resulted in decreased survival of the wild populations they are intended to support NRC , pg. Impacts of hatchery fish include overfishing of wild populations in mixed-stock fisheries Hilborn and Eggers , competition with wild salmon in both fresh water and the ocean Ruggerone and Nielsen , and a reduction in life history diversity making populations more susceptible to climate variability Moore et al. Due to the high costs of restoration and the difficulty in predicting or measuring outcomes, some have argued that the best way to protect salmon for future generations is to create salmon sanctuaries that maintain intact and connected habitats throughout the watershed from headwaters to the ocean Rahr et al.

Protecting entire watersheds is especially important to sockeye, Chinook, and coho salmon, which spend years rearing in fresh water prior to entering the ocean. These sanctuaries would provide habitat for salmon populations with heightened resilience to factors outside of management control, such as climate change and changes in the ocean environment.

The salmon populations in Bristol Bay meet all the criteria for selecting sanctuaries across the North Pacific by having intact habitats, abundant populations, and a high diversity of life history patterns Schindler et al. In addition, several studies have targeted Bristol Bay as a high priority for salmon conservation. The Kvichak, Nushagak, and Wood watersheds were ranked third, 44th, and fourth, respectively, in an analysis of physical complexity of watersheds from California to the Kamchatka Peninsula Luck et al.

Pinsky et al. Bristol Bay, the Kamchatka Peninsula, and coastal British Columbia all had clusters of high conservation value catchments. No published materials specifically address the question "Why do Bristol Bay watersheds support so many salmon? Obviously, the simplest answer is "Habitat. Our inquiry led us to the conclusion that interplay between the quantity, quality, and diversity of habitats in these river systems accounts for their productivity. The major habitat attributes discussed here were identified in personal communications with Dr.

Tom Quinn University of Washington and Dr. Jack Stanford University of Montana. Habitat quantity An obvious feature of the Bristol Bay watershed is the abundance of large lakes Figure 9. The Wood River, a major tributary to the lower Nushagak River, drains an interconnected chain of four major lakes- lakes Kulik, Beverly, Nerka, and Aleknagik- and several smaller lakes. Lakes cover 7. Lakes cover Within the Nushagak River basin, lakes cover With the exception of Chikuminuk Lake, all of the major lakes named above are accessible to anadromous salmon.

Since watershed elevations in the Bristol Bay region are relatively low Table 8 , barriers to fish migration are few and a large proportion of the watershed can be accessed by salmon. Since fish use must be documented firsthand by field biologists, a large proportion of anadromous fish habitat undoubtedly remains undocumented.

For example, a recent survey targeted undocumented headwater i. Comparison of landscape features potentially important to sockeye salmon production for watersheds across the North Pacific top portion of table and across the Bristol Bay watershed bottom portion of table. All landscape data are from the Riverscape Analysis Project Luck et al. Wood Kvichak inc. Pat Shields, ; Copper is from sockeye brood tables for Copper River pers.

Tim Baker, Map of surveyed anadromous streams in the Nushagak and Kvichak watersheds. First of all, Bristol Bay streambeds tend to have abundant gravel, which is essential substrate for salmon spawning and egg incubation Bjornn and Reiser , pgs. Several Pleistocene glacial advances have left behind a complex landscape of gravel-rich moraines, melt-water deposits, and outwash plains Stilwell and Kaufman , Hamilton and Kleiforth As stream channels meander and cut through these deposits, gravel and other sediments are captured and formed into riffles, bars and other habitat features.

Groundwater inputs to streams and lakes are also an important feature of salmon habitat in the Kvichak and Nushagak watersheds. Rainwater and melting snow infiltrate the extensive glacial deposits and saturate pore spaces below the water table, thus recharging the groundwater aquifer Power et al. Ponds are common on the Bristol Bay landscape and contribute disproportionately to groundwater recharge Rains Once in the aquifer, groundwater flows slowly downhill and eventually surfaces in areas of relatively low elevation, like stream channels or lake basins.

Areas of groundwater upwelling are heavily used by spawning sockeye salmon because they provide circulation, stable flows, and stable temperatures Burgner , pgs. These habitats include lake beaches and spring-fed ponds, creeks, and side channels Burgner , pgs. Studies in the Wood River system of Bristol Bay demonstrate the importance of groundwater upwelling to spawning sockeye salmon. In lakes, densities of beach spawners were highest at sites with the strongest upwelling, while spawners were absent at beach sites with no upwelling Burgner , pg.

In a spring-fed tributary to Lake Nerka, the distribution of sockeye salmon spawners also corresponded with areas of groundwater upwelling Mathisen , pgs. Large numbers of sockeye salmon in the Kvichak River system also spawn in lake beaches, spring-fed ponds, and other groundwater-associated habitats Morstad , pgs. In addition to spawning sockeye, groundwater is an important habitat feature for other salmon species and life history stages.

Chum salmon have been shown to preferentially spawn in areas of groundwater upwelling Salo , pg. Groundwater also maintains ice-free habitats used extensively by wintering fishes, helps to maintain streamflow during dry weather, and provides thermal refuge during periods of warm water Reynolds , Power et al.

Salmon themselves are another important habitat feature of Bristol Bay watersheds. Each year, the region's spawning salmon populations convey massive amounts of energy and nutrients from the North Pacific to fresh waters. These marine-derived nutrients MDN , released as excreta, carcasses, and energy-rich eggs, greatly enhance the productivity of freshwater ecosystems, making Pacific salmon classic examples of keystone species that have 40 large effects on the ecosystems where they spawn Willson and Halupka , Power et al.

Salmon contain limiting nutrients i. Given the high densities of spawning salmon in some streams, MDN subsidies can be large. On average, spawning sockeye salmon import 50, kg of phosphorus and , kg of nitrogen to the Kvichak River system and 12, kg of phosphorus and , kg of nitrogen to the Wood River system each year Moore and Schindler In high latitudes, the importance of marine nutrients is magnified by the low ambient nutrient levels in freshwater systems Gross et al.

In Iliamna Lake, for example, nitrogen inputs from spawning salmon greatly exceed inputs from the watershed Kline et al. Resident fishes e. The increase in rainbow trout diet was attributable to salmon eggs, salmon flesh, and maggots that colonized salmon carcasses, while the increase in Arctic grayling diet was attributable to consumption of benthic invertebrates dislodged by spawning salmon Scheuerell et al.

A bioenergetics model suggested that these subsidies were responsible for a large majority of the annual growth of these fish populations Scheuerell et al. In a stream in the Kvichak River basin, Dolly Varden moved into ponds where sockeye salmon spawned and fed almost entirely on salmon eggs Denton et al.

The growth rate of these Dolly Varden increased three-fold while salmon eggs were available Denton et al. On the Kenai Peninsula, Alaska, recent work has shown that the number of salmon spawning in a given stream is an important predictor of the growth rate and energy storage among coho salmon and Dolly Varden rearing there Rinella et al.

These and other studies indicate that the availability of MDN enhances growth rates Bilby et al. MDN is also linked with bottom-up effects on aquatic food webs, but any resulting effects on fish are less clear. In streams, increased standing stocks of biofilm Wipfli et al. Stream-dwelling fishes may benefit indirectly through increased macroinvertebrate production, but this has yet to be established. However, it is not clear if these nitrogen inputs have measurable effects on sockeye salmon populations Schindler et al.

The prolonged depression of salmon stocks in the Columbia River basin is a prime example, where a chronic nutrient deficiency hinders the recovery of endangered and threatened Pacific salmon stocks Gresh et al. Density-dependent mortality has been documented among juvenile Chinook, despite the fact that populations have been reduced to a fraction of historic levels, suggesting that nutrient deficits have reduced the carrying capacity of spawning streams in the Columbia River basin Achord et al.

A population viability analysis has indicated that declines in MDN have very likely contributed to low productivity of juvenile salmon and that increasing the productivity could lead to large increases in the salmon population Zabel et al. Diminished salmon runs, thus, present a negative feedback loop where the decline in spawner abundance reduces the capacity of streams to produce new spawners Levy Fisheries managers recognize the importance of MDN in sustaining the productivity of salmon systems and are now attempting to supplement nutrient stores by planting hatchery salmon carcasses and analogous fertilizers in waters throughout the Pacific Northwest Stockner , Shaff and Compton In addition to their inherent natural productivity, Bristol Bay watersheds have not been subjected to anthropogenic watershed disturbances that have contributed to declining salmon populations elsewhere.

For example, Nehlsen et al. They found that stocks appeared to face a risk of extinction; of these, habitat loss or modification was a contributing factor for These cases were in addition to at least stocks that had already gone extinct Nehlsen et al. A National Research Council committee NRC , convened to review the population status of Pacific Northwest salmon, summarized that: The ecological fabric that once sustained enormous salmon populations has been dramatically modified through heavy human exploitation - trapping, fishing, grazing, logging, mining, damming of rivers, channelization of streams, ditching and draining of wetlands, withdrawals of water for irrigation, conversions of estuaries, modification of riparian systems and instream habitats, alterations to water quality and flow regimes, urbanization, and other effects.

Thus, it is generally agreed that a complex and poorly understood combination of factors - with direct and indirect effects of habitat degradation at the fore - are responsible for declining Pacific Northwest salmon stocks NRC , Gregory and Bisson , Lackey In watersheds of the Bristol Bay region, including the Nushagak and Kvichak rivers, human habitation is confined to a few small towns and villages, roads are few, and large-scale habitat modifications are absent.

Additionally, invasive fishes and riparian plants, which can negatively impact native fish populations, have not been introduced to Bristol Bay's watersheds. Habitat diversity A diverse assemblage of spawning and rearing habitats is an exceedingly important feature of Bristol Bay's riverine ecosystems.

Since salmon adapt in predictable ways to conditions within their specific environments, a high level of habitat diversity fosters a correspondingly high level of population and life history diversity. The utilization of different types of spawning habitat is an easily observable example. Suitable lotic habitats range from small gravel-bed creeks to large cobble-bed rivers Hilborn et al.

Spring-fed ponds are also used, as are areas of groundwater upwelling on mainland lake beaches, and rocky beaches of low-lying islands Hilborn et al. Sockeye salmon have adapted to each of these environments in predictable ways, optimizing behavioral and physiological traits like timing of spawning, egg size, and the size and shape of spawning adults Table 9; Hilborn et al. The result is a stock complex comprised of hundreds of distinct spawning populations, each adapted to its own spawning and rearing environment.

This complexity is compounded by variation within each spawning population, likely in ways that are not yet fully understood Hilborn et al. One clear example is variation in the amount of time spent rearing in fresh water and at sea Table Within a given cohort, most individuals rear for either one or two years in fresh water, although a small number may spend three years or go to sea shortly after hatching i.

The latter life history is relatively common among Nushagak River sockeye, many of which rear in rivers as opposed to lakes. Once at sea, most fish will rear for an additional two or three years, although a few will rear for as little as one year or as many as five years. This life history complexity superimposed on localized adaptations results in a high degree of biological complexity within the stock complex.

A summary of life history variation within the Bristol Bay stock complex of sockeye salmon from Hilborn et al. Element of biocomplexity Range of traits or options found Watershed location within Bristol Bay complex Seven different major watersheds, ranging from maritime-influenced systems on the Alaska Peninsula to more continental systems Time of adult return to fresh water June -September Time of spawning July- November Spawning habitat Major rivers, small streams, spring fed ponds, mainland beaches, island beaches Body size and shape of adults - mm body depth at mm male length: sleek, fusiform to very deep bodied, with exaggerated humps and jaws Egg size 88 - mg at mm female length Energetic allocation within spawning period Time spent rearing in fresh water Time between entry into spawning habitat and death ranges from days to several weeks years Time spent at sea years Table Variation in time spent rearing in fresh water and at sea for Bristol Bay sockeye salmon.

Numbers represent percentage of fish returning to the respective river systems after a given combination of freshwater and sea rearing periods. This is because differences in habitat and life history lead to different population responses despite exposure to the same prevailing environmental conditions. For example, a year with low stream flows might negatively impact populations that spawn in small streams but not those that spawn in lakes Hilborn et al.

Asynchrony in population dynamics of Bristol Bay sockeye has been demonstrated at both the local scale i. The latter is demonstrated nicely by the relative productivity of Bristol Bay's major rivers during different climatic regimes Hilborn et al. Population and life history diversity within Bristol Bay sockeye populations can be equated to spreading risk with a diversified portfolio of financial investments Schindler et al. Under any given set of conditions, some assets perform well while others perform poorly, but maintenance of a diversified portfolio stabilizes returns over time.

Within the sockeye stock complex, the portfolio of population and life history diversity greatly reduces year-to-year variability in run size, making the commercial salmon fishery much more reliable than it would be otherwise. If Bristol Bay sockeye lacked biocomplexity and the associated stabilizing effects, run sizes would fluctuate widely and complete fishery closures would happen every two to three years Schindler et al.

While the analyses described here apply to the Bristol Bay commercial sockeye fishery, portfolio effects certainly stabilize populations of other fish species and increase the reliability of sport and subsistence fisheries. In addition, portfolio effects stabilize and extend the availability of salmon to consumers in the watershed food webs. Poor runs in some habitats will be offset by large runs in others, allowing mobile predators and scavengers e.

Different populations vary in the timing of spawning, which substantially extends the period when salmon are occupying spawning habitats Schindler et al. Since a diversified salmon stock complex is contingent upon a complex suite of habitats, an important question becomes: How does habitat diversity in Bristol Bay watersheds compare to that in other salmon-producing regions? The Riverscape Analysis Project calculated remotely-sensed indices of physical habitat complexity, allowing comparisons among salmon producing watersheds at the North Pacific Rim scale Luck et al.

Rankings of overall physical complexity were based on 10 attributes: variation in elevation; floodplain elevation; density of floodplains and stream junctions; human footprint; the proportion of watershed covered by glaciers, floodplains, and lakes; and the elevation and density of lakes. While the characterization of habitat complexity at this broad spatial scale is necessarily imprecise and certainly fails to detect nuanced habitat features, it does seem to quantify attributes that are important to salmon as it explained general patterns in salmon abundance in validation watersheds Luck et al.

The studies reviewed here demonstrate how biocomplexity in salmon populations provides resilience to environmental change. This resilience can break down when habitats are degraded or when the genetic diversity that allows salmon to utilize the full complement of available habitats is diminished.

The loss of habitat diversity and associated loss of population diversity has contributed to declines of once prolific salmon fisheries, including those in the Sacramento Lindley et al. Lindley et al. The life history diversity of this historical assemblage would have buffered the overall abundance of Chinook salmon in the Central Valley under varying climate conditions.

Levin, and R. Density-dependent mortality in Pacific salmon: the ghost of impacts past? Ecology Letters Southwest Alaska rainbow trout management plan. Anchorage, AK. Augerot, X. Atlas of Pacific salmon : the first map-based status assessment of salmon in the North Pacific. Portland, OR. Azuma, T. Diel feeding habits of sockeye and chum salmon in the Bering Sea during the summer. Nippon Suisan Gakkaishi Baker, T. Fair, F. West, G.

Buck, X. Zhang, S. Fleischmann, and J. Review of salmon escapement goals in Bristol Bay, Alaska, Alaska Department of Fish and Game, Anchorage. Battin, J. Wiley, M. Ruckelshaus, R. Palmer, E. Korb, K. Bartz, and H.

Projected impacts of climate change on salmon habitat restoration. Proceedings of the National Academy of Sciences Behnke, R. Native trout of western North America. Beverton, R. On the dynamics of exploited fish populations. The Blackburn Press. Bilby, R. Fransen, and P. Incorporation of nitrogen and carbon from spawning coho salmon into the trophic system of small streams: Evidence from stable isotopes.

Canadian Journal of Fisheries and Aquatic Sciences Fransen, P. Bisson, and J. Response of juvenile coho salmon Oncorhynchus kisutch and steelhead Oncorhynchus mykiss to the addition of salmon carcasses to two streams in southwestern Washington, USA. Bjornn, T. Habitat requirements of salmonids in streams. Pages in W. Meehan, editor.

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