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A historical review of the shad of

Item Type monograph

Authors Mansueti, Romeo; Kolb, Haven

Publisher Chesapeake Biological Laboratory

Download date 02/10/2021 23:42:48

Link to Item http://hdl.handle.net/1834/27445 State of BOARD OF NATURAL REsOURCEhS DEPART-MElNT OF RESEAWClH AND EDUCATION

A HISTORICAL REVIEW OF THE SHAD FISHERIES OF NORTH AMERICA

BY ROMEO hlAN,SIUETI md HAVEN KQLB

UBRART Unlvenity of Maviand, CES Chesapeake Biological Laboratog Solomom, Maryland 20688 US

Publication No. 97

CHESAPEAKE BIOLOGICAL LABOlRATORY Solomcms, Maryland State of Maryland DEPARTMbENT OF RESEARCH AND EDUCATION

Commissioners

FRANKLIND. DAY, Chairman...... Cenkrville EARLET. NAWKINS...... Towson ERNESTN. CORY...... College Park B. H. WILLIER- .....-...... "-...... 1...... B,altimore FREDERICKTRESSELT ...... "...... Thurmont

Director

R. V. TRUITT...... olomons Island

State Weather Service YP8C)*i .9 G. N.f~~~c~~o,.Met~rglogjstin Charge ...... Baltimore d $? - 9

A"" .*.-s : b "$; ,-, ) ' 2 Chesapeake Biological Laboratory

BYRONL. ASHBAUGH,M-ED., Education Assistant G. F. BEAYEN,M.A., Biologist I, Oyster Investigations DAVIDG. CARGO,M.S., Biologist 11, Crab Investigations HAROLDJ. ELSER, PH.IM., Biologist 111, Aquatic Investigations ADALINE GALL, Secretary GEORGEB. GRAY,B.S., Administrative Assistant II HARRYA. HEHSEL, JR., Investigator LEAH BELLEJONES, Senior Stenographer JOHN R. LONGWELL,M.S., BiologiSt III, Game Investigations JOSEPH H. MANNING,M.S., Biologist 11, Oyster Investigations ROMM)MANSUETI, M.S., Biologist ZI, Investigations HARVEYMISTER, Captain, Fish Culturist, in Charge of Commercial Fish Hatcheries RUDOLF' 8. SCHELTEMA,B.S., Biologist IZZ F. Pi. SIELING,B.S., Biologist II, in Charge of Chincoteague Project RICHARDE. TILLER, PH.D., Biologist I, Fishery Investigations R. V. T~UITT,PH.D., Biologist R. D. VAN DEUSEN,[M.S., Biologist I, in Charge of Natural Resource Inventory EARLT. WALKER,B.S., Biologist 11, Fishery Investigations CRAIGD. WHITESELL,B.S., Biologist 111, Forestry Investigation BETTYANN WOODBURN,Senior Typist ii ABSTRACT During the past 50 years the relative productivity and value of the shad fisheries of North America have decreased as reflected in recorded commercial catches. In 1897, the United States catch aggregated almost 50,000,000 pounds, and, at the time, was a growing, valuable industry. In 1949, the fisheries ranked 38th in voIume (about 11,000,000 pounds), and 26th in value ($1,636,000). The reasons for the decIine are due to many conditions that are biological and socio- economic in nature. These differ from one area to another in varying degrees. The factors which have been held responsible are: (a) pollution; (b) destruction or impairment of spawning and nursery areas by siltation; (c) ; (d) hydroelectric and canal dams; and, perhaps, (e) natural fluctuations in abund- ance. Natural catastrophes, parasites, and predators are not considered important in causing a decrease in commercial production. Attempts to rehabilitate the fisheries, by means of stocking artificially-reared fry and pond-reared fingerling shad, appear to have faiIed in every instance. The introduction of shad fry on the Pacific"Coast, however, has resulted in a major fishery. Early in the history of the decline, the first attempts at managing shad were in the form of week-end interdictions, limiting of seasons, and of declaring illegal, certain gear that were highly efficient. The most recent and significant program is a controlled catch man- agement plan, operating at this time only in Maryland, which attempts to control the rate for shad by restricting the number of fishermen and gear through a licensing program in order to assure a greater escapement of brood stock and thus, in time, a greater sustained yield. An analysis of the conditions existing at three fishways at the following locations, (a) Bonneville Dam on the , (b) Lawrence Dam on the Merrimack River, and (c) Holyoke Dam on the Con- necticut River, indicates that such passes are not fully successful despite the transportation of numbers of shad over dams. One of the reasons for difficulty in evaluating passes is that the percentage of the total run that ascends fishways cannot be determined. The total run is unknown for all but a few rivers in North America. In the final analysis, the prognosis of the shad fisheries does not appear promising. It appears doubtful that shad can be restored to the status of the late 19th century even with management programs and other devices.

CONTENTS

1. Introduction ....." ...... -...... -..... I1. Life history ...... - ...-"...-. A . Name ...... - ...... -...... -.-.- " ... B . Description ...... ".- ...... -.... -..... - ...... C. Distribution ...... "...... -...... " .... - .... -...... D . Migration and tagging studies.. ...,...... w...... -...... -.-...... E. Factors affecting migrations...... -..-...... -...... F . Shad in the ocean..... "...... *...... -....*.....-...... -...... - ..-...... G. Shad in rivers.- ...... " ...... a .....-. .- ...... H . Sexual maturity...... "...."...... -...... -...... -" ...... I . Courtship and spawning grounds...... J . .Ovulation ...... ".- ...... K . Number of eggs...... L . "Down-runners'' ...... M . Young shad...... - .... "...... - ...- N . Movements of young shad...... '0. Food of young shad...... P . Food of adult shad...... "...... Q. Predators and parasites of shad...... R . Ag-ing and growth analysis of shad...... -..... I11 . Shad fisheries of .....l...... m...... -...... -...... - ..... IV. Shad fisheries of ...... -.... -....- ..... V . Shad fisheries of ...... -.... " ..... VI. Shad fisheries of ...... -...... VII. Shad fisheries of ...... ;...... "...... -...... " .... - ...... VIII. S,had fisheries of Maryland ...... " ..-.-..... -...... -..... IX. )Shad fisheries of State...... -...... X . Shad fisheries of ...... XI . Shad fisheries of ...... XI1. Shad fisheries of State...... "....-... "..... XI11. Shad fisheries of ....... "...... - "...... XIV . Shad fisheries of Rhode Island...... -...... XV . S'had fisheries of ...... XVI . Former shad fisheries of ...... XVII . Former shad fisheries of Xe~v.Hampshire ...... XVIII. Shad fisheries of ...... XIX . Shad fisheries of Eastern Canada...... "...... XX . Shad fishe~iesof the Pacific Coast...... -" ...... XXI. Shad fisheries of State...... XXII. Shad fisheries of ...... -...... q...... d,...... -.... "..... XXIII . Shad fisheries of C'alifornia ...... XXIV . Decline of shad production ...... A . Introduction .-...... B . Overfishing ...... IC . Dams and cessation of spawning...... D . Pollution and survival...... "....-...... E . Deforestation and freshets.- ...... *...... -...... F . Siltation and spawning...... "....-...... -...... G . Destruction of juveniles ...... H . Effects of gear, trap-nets, and other artificial conditions...... I . Natural catastrophes...... J . Natural mortality ...... "...... K . Predators and parasites...... "-..... L . Natural fluctuations in abundance...... M . Summary of factors causing the decline ...... XXV . lklanagernent of the shad fisheries...... , ...... A . Introduction ...-...... w...... w...... w..... B . ,Shad research as a managenlent tool ...... 1C . Laws as management measures...... "-...-.-...... -...... --..... D . Theory of controlled-catch management plan for shad fisheries...... E . Summary of the management of the shad fisheries ...... ,...... V

INTRODUCTIION During the past 50 years the relative productivity and value of the shad fisheries of North -4merica have decreased as reflected in the catch records (see Figure 1). In 1897 the United States catch of the , sapidissima (Wilson), aggregated almost 50,000,000 pounds, and at the time, was a growing, valuable indus- try. By 1935, however, the catch had dwindled to 8,000,000 pounds, and the shad fisheries ranked 21st among the various Atlantic coast fisheries with respect to volume, though 11th in value. The shad fisheries ranked 38th in volume (about 11,000,000 pounds), and 26th in value (about $1,636,000). in 1949. Since the North American shad fisheries have steadlly declined in production, now constituting but 0.2 percent of the total catch (4,796,000,000 pounds) of the fisheries of the United States and Canada, it appears that a general survey of the biological and econon~icaspects of the entire industry is in order. The importance of investigating and solving the diverse problems of the shad fisheries has been realized by both Atlantic and Pacific coast fishery officials, most of whom are support- ing or actively pursuing research in the management of shad fisheries. The United States Fish and WiIdlife Service is also concerned about the decline of shad, and has conducted many investigations. At this time the Shad Investigations section of the United States Fish and Wildlife Service's (U. 5. F. W. 8.) Middle and Bouth Atlantic Fishery Investigations, is carrying on an intensive study of shad on the Atlantic coast. HARVEST OF SHAD

YEAR Figure 1 The following historical review was begun in 1951 in order to sumniarize and bring up-to-date the information to be found in the literature and available from various state and federal agencies. It is not exhaustive or as complete as the re- port of the shad fisheries of the United States by Stevenson (1899), which continues to be the most comprchensive study extant. Nor does it contain a biological sum- mary that is in any way comparable to the monograph by Leim (1924), to the general review of artificial propagation of shad by Leach (1925), or to the detailed summary of madieting of shad by Johnson (1938), all of which are references invaluable to analysis of the shad fisheries of North America or its geographical sections. The review has attacked, on the contrary, several aspects that are not synthes- ized in the current literature, namely: (1) the role of management and regulations in the restoration of the shad fisheries; (2) the role of fishways in shad migrations; (3) the evaluation of hatcheries in the rehabilitation of shad runs; (4) a general study of the socio-economics of the fisheries; and (5) a review of the over-all and sectional trends and the annual recorded commercial production of shad. In addition, the study attempted to summarize the essential points in: (1) a review of the early history of shad in each state and geographic area; (2) the current fisheries for shad in these areas; (3) factors contributing to the decline in production of shad in each state; and (4) recent information on life history, with emphasis on the phenomenon of migrations. A complete bibliography of American and foreign shads has been corn- piled. At the same time, many scattered and significant studies dealing with basic research on the biology of shad, and effective means of managing the fisheries have been abstracted and cited in appropriate places in the report.

SCOPE ANGDPROCEDURE There are slightly over 1200 references in the bibliography that deal directly or indirectly with shad; of these, about 200 concern foreign shad or anadromous clupeoids closely related to the American shad. Obviously, it was inlpossible to con- sult, digest, and integrate all of this material in the present report. The sources from which much of the report was drawn are cited in the text. Of greater concern during the survey was the dearth of material in the literature on the present condi- tion of the fishery and recently-acquired knowledge about shad. Despite the tre- mendous amount of written material on the American shad, a large portion is sub- jective in nature, or repetitious of the writings of some of the pioneer shad biologists and fish-culturists of the old United States Fish Commission (U.S.F.C.). An attempt was made to select the literature that contributed most objectively to an accurate and adequate treatment of the various subjects discussed in the text. Thus, by reviewing many of the older papers listed in the bibliography, a basic framework or outline of information was constructed upon which much of the recent published or unpublished material was integrated. A major portion of the present survey was prepared from material gathered by means of (1) personal interviews, (2) questionnaires, (3) personal letters, and (4) unpublished manuscripts and observations (for which permission was obtained to abstract pertinent portions from the responsible authorities). A list of the individ- uals who have contributed information, time, or their efforts, and the states for which they supplied information, is given in the acknowledgments. Various fishery agencies of different states, particularly Pennsylvania and the southeastern United States, were visited, and authorities or their libraries or files were consulted. Most of the questionnaires, which contained 10 general questions regarding the subjects listed in the fourth paragraph of the introduction, were answered and returned by officials or authorities of various Atlantic coastal states who were familiar with local conditions and the developments of their respective shad fisheries. Much information was obtained and many points were clarified by correspondence. As the tentative report for each state was prepared, carbon copies were sent to the various collabora- tors in the states or geographic sections with which they were most familiar. 4 copy of each section on the shad fisheries was also sent to the Shad Investigations group of the U. S. F. W. S.' Middle and South Atlantic Fishery Investigations for criticisms and advice. Almost all changes, suggestions, and corrections that were received from the collaborators were included in the final report. The senior author is responsible for most of the compilation and writing of the report, while the junior author was mostly concerned with the editorial aspects. ACKNOWLED~GMENTS Sincere appreciation is extended to many individuals and agencies for their aid and whole-hearted support of the shad project. Special consideration must be accorded to Dr. R. V. Truitt, Director, Maryland Department of Research and Edu- cation, for suggesting, outlining, and supporting the study. He contributed a great many ideas and facts, and fol!owed progress with great interest and enthusiasm. Dr. Clinton E. Atkinson, formerly Chief, and Dr. Gerald Talbot, Chief, Middle and South Atlantic Fishery Investigations, allowed the writers to refer to, peruse, and abstract much illaterial on shad in the files of the shad investigations group at Beaufort. In particular, they deserve credit for giving many valuable suggestions and for pointing ont where certain passages might be improved. The many people who supplied information for the reports for the various states are listed geographi- cally as follows: (1) FLORIDA:J. F. Dequine, B. 0. Freeman, G. Vathis, and R. W. Yerger; (2) GEORGIA:E. P. Creaser, F. J. Diclison, J. Stearns; (3) SOUTHCAROLINA: D. H. Gadsden, G. It. Lunz, A. Seabrook; (4) NORTH CAROLINA:B. B. Brandt, W. E. Fuller, J. Nelson, E. Roelofs; (5). VIRGINIA: E. Hollis, J. L. McHugh, T.Y. Massmann, C. Quittmeyer, W. F. Yarrlngton; (6) MARYLAND:G. F. Beaven, C. Coker, H. Elser, R. C. Hammer, H. Rensel, E. Hollis, R. E. Tiller, R. V. Truitt, E. Walker, D. Wallace; (7) DELAWARE:L. E. Cronin, F. C. Daiber, J. L. Harmic, E. Hollis; (8) NEW JERSEY:J. H. Allen, L. E. Cronin, T. Nelson, J. H. Underhill, J. R. Westman; (9) PENN~YL~ANIA:It. L. Bortner, @. E. French, E. R. Hoffert, P. LeFever, M. V. Seitz, %I. E. Shoemaker, H. R. Stackhouse, R. E. Turner, G. K. Urban, R. N. Weaver; (10) NEW YORKSTATE: A. W. Bromley, P. B. Duryea, M. Fredin, 3. Greeley, B. Lehman, A. Perimutter, W. C. Senning, G. Talbot; (11) RIIODE ISLAND:L. D. Stringer, R. E. Tiller, C. Whaley; (12) CONNECTICUT: E. Lehman, D. D. Moss, G. Talbot, E. Walker; (13) MASSACHUSETTS:A. D. Mugnier, F. W. Sargent, W. C. Schroeder; (14) NEW HAMPSHIRE:C. F. Jackson; (15) MAINE: R. L. Dow, W. H. Everhart; (16) CANADA:E. D. Fraser, A. G. Huntsman, A. H. Leim; (17) WASHINGTONSTATE: I. Donaldson, D. R. Johnson, J. L. McHugh, J. W. Sloat; (18) OREGON:I. Donaldson, D. R. Johnson, J. L. McHugh, J. W. Sloat; (19) :D. H. Fry, Jr., J. L. MuHugh, P. M. Roedel; (20) GENERALASSIST- ANCE:J. Cating, M. Fredin, B. Lehman, J. Sykes, C. H. Walburg, A. W. Anderson, R. P. Silliman, and C. Quittmeyer. Dr. W. C. Senning, assistant director, Division of Fish and Game of the New York Conservation Department, has provided the colored frontispiece for reproduc- tion. we express our sincere appreciation for the loan of the plates and for permis- sion to reproduce the painting. LIFE HISTOIRY OF THE SHAD Name The purpose of this short account of the life history of the shad is to review a considerable amount of scattered information recorded in published and unpublished sources. The shad, Alosa sapidissima (Wilson), mas named by Alexander Wilson, known as the father of American ornithology, in the American edition of Rees' Encyclopedia, in 1811. The type locality is believed to be Philadelphia. The name Alosa is Latin for "shad," and sapidissima is the superlative in Latin for sapidzts, meaning "good to eat." The shad appears to have been confused bp early American writers on fish with the shads found in England (these include, the Allis shad, Alosa alosa, the Twaite shad, Alosa finta j%zta, and the Killarney shad, Alosa finta kill- arnensis, according to Regan (1932). The American shad is unique because it possesses so few vernacular names that are widely used even though it is a com- mercially-important and abundant species. Sonie of the names are: "white shad," "Atlantic shad," "American shad," "Atlantic coast shad," '-1 acific coast shad," and "poplar-back shad." Consult the index for local common names. Two other species of shad are recognized in American waters; namely, the Alabama shad, Alosa alabarnae, found on the Gulf Coast, from which it ascends rivers to an unknown distance, and the Ohio or Gulf shad, Alosa ohiensis, found in the Mississippi River system where it ascends streams to Louisville, Kentucky, and to Montgomery, West Virginia (see Evermann, 1902A). These two species are of little commercial import- ance. They have not been artificially propagated. 3 Description , The shad is the largest, best known, and one of the more valuable members of the family in the United States. Its sides are silvery, and it is white below. When fresh from the ocean, it is dark bluish or greenish above, but this coloration largely is lost after the fish enters fresh water at spawning time. There is a dark blotch behind the gill openings, frequently succeeded by a series of similar spots in one or two longitudinal rows, which are seen more plainly after the scales have been removed. The ventral fins often have black or dusky edges. The body is covered with large, silvery scales, easily detached, 52 to 64 of which occupy the lateral line. The ventral edge of the body is provided with bony scutes or modified scales giving

(1) Alabama shad, Alosa alabamae Jordan & Evermann; female. Drawing from the type. (After Evermann, 1902A).

(2) Alabama shad, Alosa alabamae Jordan & Evermann; male. Drawing from the type. (After Evermann, 1902A). Flau~m2. The Alabama shad, Alosa alabamae Jordan and Evermann, is a little known species found ascending streams of the Gulf of Mexico coast of the United States. It was described in 1895 from the Black Warrior River, near Tuscaloos'a, Alabama. It is believed that shad harvested from the west coast of Florida during 1961 (see page 22) are of this species. The Alabama shad has never been of any great importance commercially. the abdominal outline a strongIy serrated appearance. The fins are small and weak; the dorsal is much nearer to the snout than the base of the caudal and is composed of 17 to 19 rays; the anal has 19 to 21 rays. The body is comparatively deep and com- pressed, the depth varying with sex and spawning condition. The head is also deep, with the free portion of the cheek deeper than long. The jaws are about equal in length, the lower one fitting into a deep notch on the tip of the upper in adult fish. Teeth are present in the young, but are not found in the jaws of the adult. The gill rakers are long, slender, and numerous, numbering from 93 to 120 on the first arch of the adult fish. The numerous pin-like bones that are encountered when shad is eaten are chiefly several series of intermuscular bones, which support the muscle segments above the ribs. The peritoneum is white, and on the side of each body wall lies a narrow strip of dark muscle. Shad are distinguished from the other clupeoids with which they are frequently associated in rivers as follows: In all of these, except in the case of the ' or hickory jack, Pornolobus mediocris, the cheek is longer than deep. In the latter 4 species it is about as long as deep, but the projecting jaw closely resembles that of the shad in shape. The river are much smaller than the shad, have fewer or shorter gill rakers and larger eyes. In the branch herring or alewife, Pomolobus pseudoharengus, the peritoneum is pale, while in the glut herring, Pornolobus aesti- valis, it is black. More detailed differences between shad and herrings are to be found in Hildebrand and Schroeder (1928). Great difficulties are often experienced by workers with young clupeoids in dis- criminating between the various species. Leim (19e4), Leach (1925) and Hildebrand and Schroeder (1938) give as the most prominent and constant distinguishing marks, particularly among juveniles where other differences are frequently less pronounced, the differences in the shape of the cheek and the structure of the lower jaw. In the herrings the lower jaw is bent sharply upward from its tip, forming a prominent angle midway of its length, while in the shad, this character is much less pronounced. Leim presented detailed data on the characters of the young shad at various stages which allow their separation from alewives of similar sizes. It was found that the

(1) Ohio shad, Alosa ohiensis Evermann; female. Drawing from the type. (After Evermann, 1802A).

(2) Ohio shad, Alosa ohiensis Evermann ; male. Drawing from one of the cotypes. After Evermann, 1902A). FIGum 3. The Ohio shad, Alosa ohiensis Evermann, is an indigenous shad of the Mississippi River basin, where, at certain localities, it has been taken commercially, by sport fishermen, and by certain fishermen for use as bait for catfish. It has never been abundant, nor as widely distrihuted as Alosa sapidissima, hence its contribution to the total production of shad in North America is negligible. number of myotomes anterior to the anus was the best character to be used in this connection until the fish reached such a size that the vertebrae could be counted. Gill raker counts run higher in young shad than in hickory shad, although it is recognized that the number increases greatly with age in certain clupeoids. The female shad is larger than the male, the average difference in weight of adults being more than a pound. The mature males taken in the fisheries of the Atlantic coast weigh from 1% to 6 pounds, the average being about 3 pounds; the females usually weigh from 3 to 6 pounds, the average being about 4% pounds. The general average for both sexes is between 3% and 4 pounds. In this paper a con- version of 3.5 pounds per shad is used in estimating total poundage of shad from numbers. Dr. C. E. Atkinson, former Chief, Middle and South Atlantic Fishery Investigations, U. S. F. W. S., used a conversion factor of 3.61 pounds per shad in estimating the catch in pounds of shad in the for the Pennsylvania 5 Joint Legislative Committee in 1950. In the early history of the fisheries, shad weighing up to 11, 12, and even 14 pounds were reported, but 9-pound shad are now very rare on the Atlantic Coast; 10 pounds seem to be the maximum. Colonel H. E. Shoemaker, president of the Pennsylvania Folklore Society, stated that the largest shad ever caught in Pennsylvania waters was 30 inches long and 20 inches deep, and possessed the astounding weight of 20 pounds, more than 5 pounds above the maximum recorded weights for shad. This large shad was taken between 1890 and 1904 on the south side of Green's Dam at Clark's Ferry, and was mounted and dis- played by a taxidermist at a local tavern for many years. It has since been lost. At present, during some seasons an unusual number of large shad (7 to 9 pounds) rappear in certain streams. On the Pacific Coast, according to Leach (1925) shad iaverage a pound or more heavier than on the Atlantic, some occasionally attaining n weight of 14 pounds; many have been reported weighing 9 to 12 pounds.

(1) American shad, Alosa sapidissinza (Wilson) ; female. (After Evermann, 1902A).

(2) American shad, dlosn snpifissssima (Wilson): male. (After Evermann, 1902A). Figure 4. The eeriean shad, Alosa sapid

The shad is distributed from the Gulf of St. Lawrence to Florida, the area of its greatest abundance being from North Carolina to Connecticut. On the Pacific Coast, :from the original introductions in 1871, shad liave iiiultiplied and distributed them- selves along nearly 3,000 miles of the coast from southern California to southeastern ACllaslca. For a more detailed account of the distribution of shad in specific areas, see the various accounts of the shad fisheries of the Atlantic and Pacific coastal states. Mildebrand and Schroeder (1928) and Leach (1925) state that many experi- ments in transplanting the shad to waters in which it was not native were made by the U. S. F. C. when hatching operations were first undertaken. Fry were liberated in the various streams of the hlississippi Valley, and Gulf of Mexico, also in several lakes, including the Great Lakes, in Bear and Jordan Rivers (both tributary to Great Salt Lake, Utah), in Colorado River at Needles, Arizona, and, of course, on the Pacific Coast. They became established only on the Pacific Coast. Migration and Tagging Studies of Shad Despite the commercial importance of shad and the attention that has been devoted to its artificial propagation for so many years, very little explicit knowledge of its habits and life history has been obtained. Facts about the extent and direction of the migrations of anadromous fishes are factors of vital interest in the successful management of a fishery. For example, it" was widely held that it was impossible to transport adult shad, and yet Sykes (1950 and 1951) reported how he successfully moved adults up to distances of 95 miles; he also gave an account of conditions under which adults and fingerlings can be moved. Leach (1925) has posed a few interesting questions: "Bow is the run in one river affected by the fishing operations in another? To what degree does artificial reproduction at one point yield returns of value to another? What is the origin of "sea shad" which are a feature of the local markets on the northeastern coast." There are no complete answers to these questions, but the account of migrations may suggest some. The annual migration of adult shad from the ocean to the rivers is for the sole purpose of reproduction. The fish ascend waterways to suitable spawning grounds in fresh water above tidal effects, and take several weeks in depositing and fertilizing their eggs in any given stream. The young hatch and remain in fresh water until autumn, after which time they descend streams to the ocean. They do not return until they are either three (Greeley, 1937) or four or five (Leim, 1924) years old. Not much is known about the movements or whereabouts of young shad, but it is believed that they travel great distances. Efforts have been made to determine the distances travelled, the distribution, and the total population of shad in certain rivers by tagging experiments. More important, however, has been the efforts to test the parent-stream or homing tendency of shad. The parent-srream theory has resolved itself into three questions: (1) do shad return "intentionally" to the natal stream to after three or four years of life at sea or do they, by random movements, wander into the same stream in the instances that are recorded; (2) what percentage of adult shad return to streams in which they have already spawned; and (3) where do adult shad go after they have finished spawning in fresh water? I The only successful attempt made to investigate the homing instinct of juvenile I shad was the experiment conducted by Hollis (1948). He tagged 2,466 fingerlings (mostly shad; some were river herrings which were accidentally introduces in the sample) measuring about 10 em. long at Edenton, North Carolina in October, 1941. The shad were held in Ringer's solution after tagging until the incisions in the abdomen throngh which the celluloid tags were placed had healed. A total of 1,078 1 fish was released in late October in Pembroke Creek. Three recoveries from this 1 study are recorded from the Chowan River and Albemarle Sound within a radius of ' 10 miles of the place of release, three, four, and five years later, i. e., April, 1944, March, 1945, and April, 1946, respectively. This work is the only objective observa- tion on returns of young shad that is generally consistent with the theory that they return to the stream of their nativity. Although the comparison of the number of returns with that of the number tagged is not statistically significant, the records are strengthened by the lack of recoveries of tagged shad elsewhere. The tagging of juvenile shad was continued by the U. S. F. W. S. and the Chesapeake Biological Laboratory, Maryland Department of Research and Education LC. B. L., Md. D. R. E.] in 1948 and 1950, when several hundred pond-reared juveniles were tagged and released at Solomons, Maryland, but no results are available. One of the studies concerning the possible influence of environmellt upon juvenile shad was made from scale markings as revealed from a study of 11,399 scale samples and body measurements fr6m adult shad collected in 1940 and reported in an unpub- lished .thesis by Hammer (1942). He described two patterns of scale development, i. e., (I)an inner fresh-water mark; and (2) an outer salt-water periphery. Measure- ments of the fresh-water zone were used to compute the juvenile body lengths attained at the termination of the fresh-water period. The different calculated lengths indi- , cated that mature Chesapeake Bay shad return to the stream of their origin; that is, specific populations of shad visited definite spawning grounds year after year. Although he did not find significant differences between the calculated lengths deter- mined from scales collected within the James, York and Pocomoke rivers, he found 7 other scale characteristics indicating the existence of populations peculiar to each river. Differences were found between lengths of fingerlings of the Rappahannock, Potomac, and Susquehanna River shad. Two self-perpetuating shad populations were thought to exist in the two tributaries of the York River, namely, the Mattaponi and the Pamunkey Rivers, on the basis of scale measurements. Hammer believed that is was possible that each spawning area ". . . may have unique chemical and physical properties directly influencing juvenile growth and behavior." He concluded that it is probable that the fry from each river have features peculiar only to shad of the river of their origin. Unfortunately, there is no additional information from the study areas over a long period of time upon which to determine without question whether the different areas possess subtle factors that are unchangeable from year to year under varying conditions, or otherwise. Whether adult shad tagged in a spawning stream will return to the same stream after a year at sea has been extensively investigated. In 1937 a limited amount of tagging was carried out in Chesapeake Bay by the Chesapeake biological Laboratory in order to discover if there were a general mixing of shad populations along the Atlantic Coast. A shad tagged at Spesutie Island, Maryland, was recaptured in June, 1937, at Race Point, Cape Cod, Massachusetts, after travelling a distance of about 900 miles. In 1938 the tagging program was expanded and it was found that about 28 per cent of 330 tagged shad were recaptured in the vicinity of the point of tagging during the same year. One specimen tagged at the head of Chesapeake Bay was taken later from New Jersey waters. Although no actual data is available, tagging returns from tk 1938 releases were announced by Truitt (1940), who stated that indications wer:, . . . that this form visits the same breeding areas during successive years . . . Cable and Hollis (1949) stated that a few adult shad tagged in Chesapeake Bay were later recaptured in the same season in North Carolina waters. A few were caught along the New Jersey coast, and along the south shore of Long Island, New a York. A few others were taken in the Bay of Fundy. A shad tagged by Vladykov (1948 A) on the south shore of the St. Lawrence River was later recaptured off Cape Cod, Massachusetts. A total of 20 out of 2316 shad tagged in the ocean off the coast of Maine by the C. B. L., Md. D. R. E. and the U. S. F. W. S. in 1947 was recovered by fishermen from Massachusetts to Georgia in the spring of 1948, the maximum distance travelled by a shad between Maine and Georgia being about 1,200 miles. 1 The shad investigations group of the U. S. F. W. S. conducted extensive tagging I operations in the Connecticut and Hudson Rivers in 1950 and 1'951, and in Chesa- I peake Bay in ,195t2, as a part of the mark-and-recapture method of estimating total population of shad in a stream and fishing mortality. A progress report of the group I by the U. S. F. W. 8. (1952) stated that from 1,000 shad tagged in the Windsor I Locks Canal, , between June 29 and July 5, 1951, only one tag I was recovered, December 13, 1951, off Long Branch, New Jersey. The results from tagging in the were more extensive. Shad were marked during 1950 / and 1951 at the mouth of the river, and after spawning, some shad migrated along the shores of Long Island and northward to Maine, according to tag recoveries. Individual tags were also recovered in 1950 off Portland, Maine, on August 8, and north of Gloucester, Massachusetts, on November 11. During March, April and May, 1951, all along the coast from Mann's Harbor, North Carolina, to Raritan Bay, New Jersey, marked shad were recovered. Some tags were returned from catches made in the mouths of Chesapeake and Delaware bays. The report indicated that "Though tags were recovered in mouths of several streams, or rather bays into which streams empty, not one tag was returned from a spawning ground in any other stream than the Hudson. This indicates that shad return to the same stream each year for spawning." Extensive experiments were carried out by Vladykov (1950), who tagged and released a total of 1,8155 shad in the St. Lawrence River, Quebec. A total of 44 recaptures, which was a part of the six percent retur;, was recorded. Of these, 18 fish, tagged principally at Green Island, demonstrated . . . clearly that Quebec shad, after spending the winter months in the Atlantic, ascend the St. Lawrence River in May and June." Four recaptures outside of Quebec were as follows: around Octo- ber 1, 1947, at Provincetown, Cape Cod, after being in the water for 444 days; on March 30th, 1947, at Browns Bank, Nova Scotia, after 258 days; on May 31, 1948, on the Atlantic side of the Canso Strait (Arichat, Nova Scotia), after 327 days; . and on June 27,1946, in the Cumberland basin (Minudie, Nova Scotia) after 322 days. The recapture made farthest inland was that of a shad tagged at Green Island and recovered about 300 miles to the west in the Ottawa River (near Pointe-Fortune). 8 The maximum ulstance travelled from the point of $elease to the place of recapture was over 1,200 miles. Vladykov concluded that the . . . recaptures clearly indicate that mature Quebec shad . . . return again to their spawning grounds located in the St. Lawrence River. . . ." It was noted that shad were recovered on their migra- tions to and from spawning grounds during the first (same year they were tagged), second, and third spawning seasons after tagging. The Oregon State Fish Commission (1948 A and 1948 B) and Gharrett (1950) have reported on tagging activities. Shad tagged on July 2, 1946, were recovered in the Umpqua River shad fishery in 1947. A shad tagged on its way to the spawning ground in the Umpqua River on June 9, 1948, was caught on the following June 22, in the Siuslaw River, which enters the Pacific Ocean about 20 miles north of the Umpqua. Two tagged fish made a similar migration in 1949. Another tagged in 1947 in the Umpqua was recovered in 1948 in the same river, and another was recovered in Coos Bay, about 30 niiles south of the Umpqua in May, 1949. Shad were tagged extensively as part of a mark-and-recapture method of estimating the population of shad runs and fishing mortality (see account of Oregon shad fisheries). Although the various tagging experiments cited above are not as complete as might be desired to formulate some pattern of dispersal, they generally suggest the following: (1) In general, aduIt shad seem to return to the same stream in which they have once spawned, although there is as yet no measure of the percentage of repeaters that return to the same stream as compared to those that do not. (2) There seems to be a general northward migration of shad after spawning in Chesapeake Bay tributaries and the Hudson River, some shad swimming northward as much as 900 miles. On the other hand, there is evidence that some shad move southward from the two areas and also from the St. Lawrence River. It is not known, for example, whether these shad had already made a northward migration (because of the lack of information of dates and otherwise). (3) There seems to be a general mixing of shad populations from different regions around the Gulf of Maine during late summer and early winter months. In winter they presumably separate and move southward (or northward to Quebec). Shad tagged in the Gulf of Maine dis- tributed themselves in coastal rivers from Massachusetts to Georgia. (4) Very few shad have been recovered which could be classified as "lost" or deviating from the "typical" migration routes. One Quebec shad was classified as "lost" in the Atlantic when it was recovered at the head of the Bay of Fundy. Shad tagged one year in the Hudson River were recovered during the next in the mouths of several streams 1 or bays of North Carolina, Chesapeake Bay, Delaware Bay, and Raritan Bay. Signifi- cantly, none was returned from a spawning ground other than that of the Hudson River. (5) Whether tag recoveries are a good index of the wanderings of shad is a 1 moot point at this time. A variety of tags was used in the different experiments so that it is not known if some are more selective than others when tagged shad enter , areas that are being fished by specific gears. In some cases tagged shad are snagged almost immediately after being released. Another controversial point is ,that tag recoveries seem to come from the centers of extensive fisheries so that such records may not represent a fair sample of the actual distribution of tagged specimens, (6) The following provisional theory can be developed from the scattered tagging recoveries: Shad move north from the major rivers of Eastern United States aftep spawning, spend the summer off the New England coast in the , migrate south along the Continental shelf during early winter to Georgia, move shoreward, and once again migrate northward, entering the rivers in which they have previously spawned. The factors causing such a circular nlovement may be seasonal temperature and current changes of the offshore waters. It is known, for example, that in certain sections of the Gulf Stream the current is relatively strong during the early summer, falls off rapidly in strength to a minimum in October or November and then increases rapidIy until January or later. A secondary mini- mum is called for in April or May, but it is not clear from the available transport determinations whether or not the prolonged winter maximum usually surpasses the summer maximum (Storamel, 1950). (7) It is believed that the northward movement of shad is a feeding migration, while the southern movement is a spawning migra- tion, a situation somewhat analogous to some movements on the Pacific Coast. The rate of migration of shad has only been observed incidental to regular tag- ging studies. Vladykov (1950) indicated that the speed of 18 fish was from 25 to 50 miles per day. Earl Walker, formerly of the U. S. F. W. S., tagged a small male during May, 1950, at Haddam Meadows, Connecticut, and it was recaptured eight hours later below Essex about 20 miles downstream. D'Ancona (1,9128)decIared 9 that Alosa fintu swims only about five kilonieters (or a little over three miles) each day in the Tiber River in Italy. The rate of speed is probably dependent on fluctua- tions of current flow and temperature. Whether there is a distinction in the rate of speed of buck and roe shad, and whether sexes begin schooling at the same time at sea is not known. In the Potonlac River, for example, of 61,000 shad comprising the first of the run received at Washington, D. C., from March 19 to 24, 1897, 90 percent were males. This percentage is higher than usual, but the males definitely precede the females, and obviously they assemble efther at sea or they aggregate gradually during the upstream movement. Morrow (1948) pointed out that the clupeoids exhibit the most perfect of schools and show a complete lack of external sex charac- teristics. He made the statement, erroneous with regard to shad, that they spawn in indiscriminate schools without apparent heterosexual orientation. Ile pointed out, however, that the basic factor of schooling is the premise of visual reference points that clupeoids establish with one another, but no information is available on the mechanisms ifivolved. Factors Affecting the Migrations of Shad The factors influencing the movements of anadromous fishes are little known. Details of why shad migrate to the same general area for spawning every year and how they reach these points are lacking. No investigations are available to shed light on the qualitative reasons for shad migration in North America, although Roule (1922) and D'Ancona (1928) have begun such studies on the European shad, .Alosa finta. Two problems present themselves (assuming that in general shad obey the parent-stream theory) : (1) how do shad discriminate between various rivers encoun- tered during spawning migrations; i, e., do they employ visual, olfactory, or tactile methods of identification; and (2) what environmental mechanisms attract them and cause them to ascend the rivers to breeding areas? It is possible to postulate a theory from a perusal of studies made on salmon migrations and from the results of certain laboratory experiments. Little is known concerning the mechanism causing the return of Pacific salmon to the parent stream, although a vast amount of work has been published in this field. Huntsman (1945, 1948, and 1950) has summarized some pertinent information regarding factors affecting fish migrations, particularly those of the Atlantic salmon and the sea-running brook trout. He believes that strong freshets stimulate fish that have gathered at the mouth of a waterway to ascend a river. Well-fed and sexually mature fish which usually gather near the bottom of a stream swim upstream more rapidly than the current carries them downstream due to a strong rheotactic reaction. , Conversely, lean and spent fish descend a stream because they are not usually stimu- I lated by strong currents at the surface. Down-running shad leaving spawning grounds often are observed with their backs and fins jutting out of surface waters of a river because of their somewhat lower specific gravity after spawning. Hunts- man pointed out that the warming of shore water may also cause the early ascent of salmonoids. He has indicated that shad behave similarly to the salmon in relation to freshets. Ascent occurs chiefly as the freshet is subsiding. Descent of salmon at all stages occurs with freshets but chiefly at the height of a flood. It is part of the response of the fish, effected through sight, to contact with the bottom and possibly to the action of turbulences on the lateral line organs. He concluded that, "Freshets effect migration of fish in three ways: (1) they stimulate the fish to ascend streams; (2) they carry fish down streams; (3) they tend to break up the 'homes' to which the fish come to be related." D'Ancona (1928) generally held the same ideas. He declared that migration of A Eosa jintu in the Tiber River in Italy is due to physio- logical conditions which modify the sensibility of the shad confronted by the environ- mental conditions, i. e., "rheotropisrn, so to speak, directs the migration." It has long been suspected that the solution to the parent stream theory is that the physiology of anadromous fishes has been adjusted to certain physico-chemical I factors of waters. Shelford and Powers (1915) demonstrated that clupeoid and salmonoid fishes respond to a COa tension gradient in sea water. Powers (1941) pointed out the possibility or probability of a C0a tension gradient from the mouths of spawning streanls to the salmon feeding grounds. He believed that the physico- chemical behavior of mixed fresh sea water produces a COa gradient which is the dominant factor in determining the path or migratory movements of the fish in the I sea. In fact, he found that red salmon migrate to the spawning grounds along paths of fresh-salt water gradients. He stated (1939) that the chemical receptors sensi- tive to this factor are found in the branchial arches or in structures derived from branchial arches. Notwithstanding his interest in fish migrations, Powers does not 10 subscribe to the parent stream theory as such. Scheer (1939) commented that Powers did not take into consideration the fact that a run of fishes, whether in the sea or in a river, niay divide, some passing into one river or tributary while others continue in their previous course independent of a COz tension gradient. The most stinilulating discnssion of the parent stream behavior of fishes was pre- sented by Nasler and Wisby (1951), who believe that sensory niechanisnls may play an important part in the orientation of fishes. They cited the work of Craigie (1926) who released 500 sockeye salmon, in half of which the olfactory nerves had been severed, and found that the migratory behavior had been somewhat interfered with. In a series of experiments to determine the presence of some characteristic odor of the stream which guides the returning migrants, two distinct problems arose: (1) do streams have characteristic odors to which fish can react, and if so, is the odor organic or inorganic in nature, or a combination of both; and (2) can salmon (or any other fish) detect and discriminate between such odors, if they do exist? In an attempt to answer the first question the authors set up a conditioned response train- ing program with bluntnose minnows (Hyborhynchz~s). The fishes were able to discriniinate successfully between chemical differences of two Wisconsin creeks after two months' training. Extinction tests indicated that these minnows would respond to the stream odors after a "forgetting" period, which was longer in fishes trained when young than in those trained in senility. Heat cautery of the olfactory epi- thelium produced fish which were no longer able to respond to the training odors, indicating that olfaction was the sole means of discrimination in these tests. Chemical analysis of the stream waters showed a total absence of COz, proving that this com- pound was not the factor that was detected. There is a strong indication that the odorous stimulant is a volatile, aromatic snbstance. The authors postulated that the nature of the guiding odor be such that it must have meaning only for those fish conditioned to it during their fresh-water sojourn. Preliminary tests with salmon proved that they could detect and discriminate between stream odors. It is believed, in summary, that the mechanisms that might enable shad to begin their spawning migrations froin sea to fresh water are: (1) a physical stimulus such as freshets and perhaps temperature changes that stimulate them to ascend a river; and (2) a chemical stimulus such as a COz tension gradient in sea water, although Hasler and Wisby (1951) have shown that experimentally this factor did not enter " into the problem with the discrimination of stream water by minnows. In fact, they consider the common chemical sense for Con and the ability to discriminate tempera- , ture differences among fishes as crude senses when compared with the highly sensitive , olfactory system. More important, the mechanism that might enable shad to find I their home stream seems to be inherent in a stream's odor which is derived from the vegetation and soils of the watershed. These aromatics must remain constant over 1 long periods and lend a distinctive smell which can be perceived, learned, and recog- nized again after a protracted period of non-exposure. Finally, it is conceivable that each stream sends its characteristic odor as a discrete flow into the ocean until it loses its entity through mixing and dilution. Although shad may wander out of a given flow for three or four years, its physiological makeup probably is so geared that the characteristic stream flow may be picked up during migration up or down the coast in preparation for spawning. Once it has picked LIP the path of the natal stream with its characteristic odor the shad ascends the course to the spawning grounds. Shad in the Ocean Shad spend most of their existence at sea, and little is known of their habits and niovements when out of the rivers. Leach (19215) presented some of the pertinent inforniation that is known about their oceanic activity. The ocean areas to which they resort are unknown, although it is believed that they stay within the confines of the continental shelf. The food of shad in salt water has been ascertained only in a few cases. In the Gulf of Maine, the shad is known to associate in large num- bers with nlackerel and sea herring during June, September, and October. It has been taken in North Truro, Massachusetts, in fall, when the ocean temperature was from 43 to 49 degrees F.; in November it has been found on the west side of Sakonnet River, Rhode Island; in May and June it has been captured with mackerel a few miles northeast of Cape Cod Light. Hildebrand and Schroeder (1928) stated that, "To our knowledge no shad ever have been captured in the open sea off Chesapeake Bay, although menhaden purse seines are used in this region throughout the summer and fall. In the Gulf of Maine, however, adult shad not only are present (at times in abundance) from October until into December, but according to Bigelow and Welsh 11 (1925) schools of immature shad from one foot long and one-half pound in weight up to 2 and 2% pounds are reported every year at Provincetown for a short period in June. These authors report the capture of numbers of shad about 14 inches in length in the traps at Magnolia and Beverly from June 20 to July 6, 1921." The same authors quote Atkins (1887) who reported large numbers of immature shad feeding about the bays and mouths of rivers along the coast of Maine during the summer after the main body of spawning fish had ascended the rivers. These immature so-called "sea shad" belonged to the group ranging from about one-half to 2% pounds in weight. The only knowledge of the shad from the time (fall of the year) the young leave the rivers and bays, in which they were hatched, until they return as mature spawning fish is obtained from the immature fish, probably two or three years old, that are found during the summer in fairly large numbers along the shores of the Gulf of Maine and in smaller numbers south of Cape Cod. Vladykov (1936 and 1950) discusses the movements of shad in Atlantic Ocean waters off eastern Canada. Leach (1925) stated that under certain conditions the adults may remain in fresh-water rivers for a whole year. In November, 1890, ,600 were taken in Chesa- peake bay, although these may have been a very early spawning migration from the sea. They have been found in the occasionally in limited numbers in August and September, and even during the last week in December. Their move- ments are apparently largely controlled by the water temperature. It is believed that they seek to occupy an area having a temperature of 60 or 70 degrees F., and that their migrations are determined by the shifting of this area. Shad in Rivers The migrations of shad from sea proceed in successive intervals of time according to the latitude of their occurrence. They first appear in the St. Johns River, Florida, about mid-November, the season of greatest abundance being February and March. In the , Georgia and the Edisto River, South Carolina, the run begins early in January and ends the last of March. In the North Carolina rivers these stages of the migration are a little later. In the Potomac River, advance individuals appear late in February, but they are most numerous in April or early May. In the , the maximum run occurs in early May. The shad reaches the Eudson River the last of March, and is found in the Connecticut River toward the end of I April, is most abundant the last of May, and leaves the stream late in July. In the I Kennebec and Androscoggin Rivers, Maine, it is first taken in April and has left by the middle of July. In the St. John River, New Brunswick, it appears by the middle ' of May, and in the Miramichi River, New Brunswick, late in May. In the Columbia River at Bonneville Dam, Oregon, it arrives in July, and reaches a peak in mid-July. Carl and Clemens (1948) stated that shad entered the Fraser River, British Columbia, in July. Erkkila, et a1 (1950) stated that shad migration was discovered in the I Sacramento-San Joaquin Delta, California, during midJune and Iasted through early July. Leach (1925) maintained that the main body of shad ascends rivers when the temperature of water is from 56 to 66 degrees F., the numbers diminishing when the temperature is over 66 degrees. The movement of the shad up the rivers is not con- stant, but in waves, causing a rise and fall in the commercial catch. Successive schools enter the Potomac River, for example, from February to June, the males preceding the females, the latter becoming most abundant toward the close of the season. Sexual Maturity Greeley (19317) stated that shad may spawn at three years of age, whereas Leitn (1924) believed that some began spawning at four years, while others began at five years of age, as Cable (1944 C) indicated. It is generally believed that buck shad mature one year earlier than roe shad. Spawning marks are not visible on shad that are entering a breeding area for the first time; the females of this initial run are known as "virgins." It is believed that since spawning shad do not feed extensively in fresh water, the stored fats in the flesh and some material near the outer edges of the scales are taken up to mature the spawn. Thus the outer rim of the scales becomes thin and frayed in appearance. When growth of the scales is resumed the frayed area forms a distinct, ragged line which is known as a "spawning mark." Atkinson (1951) has shown that adult shad definitely feed in fresh water so that it is possible that non-feeding behavior may not he an important factor in the formation of a spawning mark. Shad whose scales have one or more spawning marks are called "repeaters." Repeater shad obviously have escaped the nets one or more times, and 12 they form the "escapement" or "spawning reserve" necessary for the restoration of the fisheries. Courtship and Spawning Grounds Shad in spawning condition may be found anywhere above brackish water, and under favorable temperature conditions spawn wherever they happen to be. In some river basins they exhibit a well-defined choice of spawning places, preferring locali- ties below the mouths of creeks, where the warmer water of creeks mingles with the colder channel water, according to Leach (1925). The favorite spawning grounds, or "shad wallows," as they are termed by the fishermen, are on the sandy flats which border the stream and the sand-bars that are found at intervals high up in fresh water of the river, according to Goode and Gill (1903). Shallow areas near the mouths of fresh water creeks appear to be preferred in some waterways. Shad are not definitely known to seek spawning grounds in brackish water, although Leim (1924) suggests that ". . . the rapid increase of the shad in California has been due to the fish spawning in brackish water where the eggs had the best chance of developing succe~sfully.'~The European species of shad behave like the American one in spawn- ing a relatively short distance above the head of the tides (see Hoek, 1900, and Roule, 1920). D'Ancona (1928) stated that the spawning of Alosa finta took place at night in deep water near the confluence of the Paglia with the Tiber River in Italy at 22-25 degrees C. When shad have reached suitable spawning grounds, and are ready to extrude their eggs, they move up to the flats seeemingly in pairs, according to Goode and Gill (1903). The time of this movement is usually between sundown and 11 P.M. "When in the act of coition they swim close together and near the surface, their back fins projecting above the water. The vigorous, spasmodic movements which accompany this operation produce a splashing in the water which can plainly be heard from the shore, and which the fishermen characterize as 'washing' ". Leach (19285) declared, "In a state of nature the shad deposits its eggs loosely in the rivers without building a nest, the two sexes running along together from the channel toward the shore and the eggs and milt being ejected simultaneously. The shad lays its eggs during the . highest daily average temperature, a condition realized about sunset, when the warmer shoal water commingles with the colder channel water, establishing a baI- ance." He observed further that principal spawning occurs from five to 10 P.M. In I the Potomac River the eggs of shad caught in a seine in the morning amounted to only I 11 percent of those taken between midnight and noon, the percentage being 14 one I year and eight during another. Leim (1924) stated that spawning took place in the evening after sundown and continued on until midnight and even later. At this time the fish could be observed swimming rapidly over a somewhat erratic course usuaIly in places where the current was neither sluggish nor swift. In some of the wild rushes the fish swam close to the surface so that the wake was plainly seen or occasionally they would "break" the surface when the splashing made their presence audible as well as visible. Newly fertilized eggs were found by placing a net near the bottom downstream from the disturbance. Ovulation A progress report on shad production problems in North Carolina waters, issued in 1938, contained a number of interesting observations on the development of shad eggs. It stated that eggs developed gradually in the ovaries as the temperature of the water rose. At higher temperatures the eggs matured more rapidly and uni- formly. When the temperature become excessive, "rotten ripe", useless eggs resulted. At 55 to 62 degres F. the ovaries apparently developed a portion at a time, and that Portion is spawned under favorabIe conditions. Numerous roe shad observed to have spawned out the contents of a small part of the ovaries, and then opened were found to contain one or both ovaries in a state of partial development. As the temperature rose from 6k? to 68 degrees the ripening of the ovaries was rapid and as the water temperature rose from 68 to 75 degrees the development of the ovaries was more rapid and complete with some of the shad producing "rotten ripe" eggs. The latter invariably resulted beyond 77 degrees. Thus shad spawn intermittently, as the ovaries ripen. When many shad are taken in a seine haul, some are not ripe, some have spawned a part and have no ripe eggs, and some have completed spawning or devel- oped "rotten ripe" eggs, all of these stages depending apparently largely on water temperatures. Very few completely ripe roe shad are taken with fully developed ovaries (see United States Bureau of Fisheries, 1938). 13 Leach (19-25) described the compact mass of eggs in the ovaries that distend the whole abdomen. The average weight of the ovaries is about 1'3 ounces. Shortly before spawning, transparent eggs of large size, differing from the opaque hue of less mature ones, are to be found scattered through the still compact ovarian mass. The mature eggs are finally freed from the remaining portion of the ovary and extru- sion begins, a liquid stream of eggs and mucus flowing from the oviduct on the slightest pressure of the abdomen. When eggs are freshly deposited they are trans- parent pale amber or pink color, and are about 1/14 inch in diameter. They are somewhat flattened and irregularly rounded in form, but immediately after fertiliza- tion the eggs become spherical through the absorption of water, then measuring about 1/7 inch in diameter. This gain is only the distended egg membrane, the vitellus or true germinal and nutritive portion not having increased. The vitellus is slightly heavier than water, hence causing the ovum to fall to the bottoln. Eggs are deposited loosely, and after reaching bottom some are buried under silt carried downstream by spring rains, and perhaps some may be eaten by predators. Hildebrand and Schroeder (1928) stated that the period of incubation varies from six days and four hours in an average water temperature of 57.2 degrees F. to a little less than three days at an average temperature of 74 degrees. The average hatching period at temperatures that prevail during normal shad seasons varies from six to ten days. Leim (1924) stated that shad spawned in Canadian waters after the temperature had reached 53 degrees F. From 1/6 to 118 of the eggs died within a few hours after their disposi- tion due either to lack of fertilization or to unfavorable conditions. The relation between temperature and rate of development of the eggs can be represented by a straight line. Number of Eggs Shad are prolific spawners, but much less so than many other food fishes. The quantities of eggs taken by spawn-takers employed by hatcheries do not represent the actual fecundity since it has been a cominon observation in the Maryland hatcheries that many eggs are cast in advance of stripping. The average number of eggs taken from one shad is not over 30,000. Single fish have been known to yield 60,000, 80,000, 100,000, and 115,000 eggs, and on the Delaware River, in 1885, one shad yielded 156,000 eggs. Recent studies by the U. S. F. W. S. (see Lehman, 1953) indicate that the total fecundity per year may be almost twice the latter figure. Many eggs fail to be fertilized, as was suggested by Leim (1924), and but a small percentage of those impregnated are hatched. After being extruded and fertilized the eggs sink to the bottom in relatively still water. In flowing water the current may carry them some distance since they are only slightly heavier than water. L'D~~n-R~nnes~9' After spawning, the emission of reproductive material leaves adult shad in an emaciated condition with thin, sharp bellies and thin sides, thin head, sunken eyes, and with a general overall whitish appearance. They are very lean and hardly fit for food, hence are known as "down-runners," "racers," and, more appropriately, "spent fish." Fishermen frequently refer to them as bucks, failing to differentiate between the spent female and the normally slender male. Although fishermen occa- sionally catch down-runners, so-called because they are caught on their way to sea, they are not conimercially popular. Their low specific gravity may cause them to swim downstream in surface waters so that they are occasionally observed during their descent. Cable (1944 B) believes that there is a possibility that some shad may die in South Carolina waters after spawning for the first time because of the big number of dead down-runners that was observed. Down-running shad in more northern waters begin to feed soon after spawning and become quite fat by the time they reach salt water in summer and fall. Youag Shad Leim (1924), who investigated the life history of the shad in Canadian waters in great detail, pointed out that the number of shad larvae produced was extremely small in proportion to the number of eggs deposited. After hatching, the shad larvae were mostly carried downstream to tidal water where they were collected and carried about by tides. Young shad when hatched are scarcely 2/5 inch in length, but they grow rapidly, reaching an average length of about 1-7/8 inches during the first half of July in the Potomac River; 2-5/8 inches by the last of August; and 2-3/4 inches by the last half of October, according to Hildebrand and Schroeder (1928). On the other hand, they recorded yearling shad that had attained, under especially favorable 14 u E F FIGURB 5 Egg and larval atnges of the shad, dlosa sagidissin~a. (After Leach, 1924). A. Freshly extruded egg enlarged, showing its envelope much wrinkled. B. Shad eggs, showing vitellus and distended ega membrane, natural size. 0. Sho~vsthe gradual accumulation of germinal matter at one pole of egg, the polar prorr.inence externally, and presence of plasmie process extending down through the vitdlnc D. ~mbryishad in position in its spacious enveloping membrane. From a photograph. E. Diagrammatic representation of an embryo tc show course of begmental ducts sd and extension outward of pectoral plates PP, which are ir~timatelyconcerned in tlfe develop- ment of pectoral fms. F. Side view of a young shad 13 days old, viewed as a transparent object. ab, rudimen- tary air bladder: L, liver; Gb, gall bladder. G. An embryo in its envelope, on the third day of development, nearly ready to hatch. FIGURE6. Em-- and larval stages- of the shad. Alosa sawidissinza. (After Leach. 1024). H and I. Two views of an egg after the blastoderm has spread considerably and the embryonic area e is well defined. K. View of unhatched embryo, which developed in a temperature of 450 F., producing distortions of tail. L. An egg envelope with its contained embryo, 44 hours after impregnation, viewed as a transparent object. M. An egg envelope with its contained embryo at the beginning of the third day of development. From a photograph. N. Anterior portion of a young fish on fourth day. To show relations of liver L to yolk Y, over which the portal vessel pv passes forward to empty into the venous sinus, in common with the anterior and posterior jugulars j and j, ba bulbus aortae, ve ventricle. 0. View of fore part of a young fish 17 days old, from ventral side. Fraum 7. Larval stages of the shad, Atom sapid

I Florida, flows north nearly parallel with the coast a distance of 375 miles, and enters , the ocean near the northeastern corner of the state. It is very broad, the width in I the lower 100 miles being from one to five miles, and for two-thirds of its length it is 1 over a mile wide, often expanding into spacious lakes. There are no permanent obstructions to the passage of fish, and shad ascended nearly to the sources of the nver. The St. Johns differs from all other streams on the Atlantic coast of the United States in that its sources are in warmer latitudes than its entrance into the ocean. As a result, shad are said to appear several months before the spawning time, and the fisheries are therefore prosecuted during the entire winter. Steve?;on (1899) quotes Marshall McDonald, one of the early f3ommissioners of Fisheries, By reference to tables giving the temperature of the St. Johns River at Jacksonville, it is seen that the temperature of the water gradually descends, reaching 60 degrees F. at Jacksonville about the last of November. This date is coincident with the first appearance of shad in the St. Johns. * * * In all other streams of the Atlantic coast the fish appear to wait until the temperature of the river has risen above that of the salt-water area into which the river empties before they ascend in the spring. The migration of shad into the St.Johns River is clearly not for the immediate purpose of spawning, as the operation is not performed for months, but in order that they may keep within the limit of the hydro-isothermal area appropriate to them." Stevenson claimed that on the St. Johns was of a recent origin, beginning after the middle of the 19th century. McDonald (1887), quoting R. E. Earll, stated definitely that a Connecticut harvested shad in Florida in 1858. The absence of extensive fisheries before the sixties was due to inadequate shipping facilities and lack of local enterprise. Drift nets were introduced about 1860, most of them being operated by fishermen from Connecticut and New Jersey. The Civil War interrupted all fishing except for local use, but at the close of hostilities the operation of drift nets was renewed and greatly increased . Stevenson pointed out that during 20 1873, from the bar at the mouth of the St. Johns River up to Palatka, 75 to 80 nets were fished during the shad season. His observer, Dr. C. Koch, stated, "These nets are 200 yards long and 10 feet deep and are set in from 10 to 182 feet of water. In 1873 the product of the sh&d fisheries amounted to 250,000 shipped or consumed. The average price at Jacksonville was 15 cents apiece." The early yield of shad has been given by Stevenson as follows: YEAR NUMBEROF SHAD YEAR NUMBEROF SHAD 1864 None reported 1878 200,000 1873 250,000 1880 71,914 1876 160,OO~O 1890 872,09.4 1877 280,000 1896 456,281 The legal season extended from December 1 to the end of the following March, with close time "between sundown on Saturday afternoon and sunrise of Monday morning of each week." Actual fishing operations began at the opening of the legal season, fully a month before shad were caught in any other water on the coast, and ended about the second or third week of March. An interdiction existed against the use of "any gill nets for the capture of shad with mesh less than 5 inches from knot to knot measured lengthwise, that is to say, a mesh when brought to a square, the sides of which are not less than 2-19 inches" or any seine with a mesh less than 3 inches. Fishermen during the late 19th century believed that shad spawned at the very head of the river, and that they never returned to sea, since they had never observed a spent fish. They believed that weak and helpless spent fish were easy prey for alligators and catfish. None of these allegations has been proved. Stevenson stated that there were three distinct geographical sections in the shad fisheries of the St. Johns: (1) From the sea to Jacksonville: In this lower section of the river, covering a distance of 218 miles wholly within Duval County, the drift net was the only form of apparatus used. Ranging from 500-600 yards in length, they were set directly across the current, one end being buoyed and the other attached to a boat, allowing them to drift with the current. Most of these nets were operated by fishermen from Connecticut and New Jersey. Nearly all of these fish were shipped to New York City and other distant markets. Between Jacksonville and Bridgeport, a distance of 46 miles, no fisheries were operated because of the two to five miles width and the sluggish nature of the river. During 1896, 291,111 shad, valued locally at $47,720, were harvested. (2) The Palatka section: This area extended from Bridge- , port to Welaka, a distance of 35 miles located wholly in Putnam County, but the fishery worked principally by drift nets was centered at Palatka. During 1896, 37,300 shad, worth $5,222, were harvested. During Stevenson's time, the widespread growth of the water hyacinth, Eichomia (Piaropus) crassipes, threatened to hinder the operations of gill netting since the floating plant entangled in the nets. It was believed that further i growth and distribution of the plant would cause an abandonment of the shad fishery in this section. (3) From Lake George to Lake Harney: This portion, known as the Upper St. Johns, consists of a series of connected lakes, the most important being Lakes George, Dexter, Rlonroe, and Harney. Sanford, on the shore of Lake Monroe, was the center of the shad fisheries. Seines were the principal apparatus used, with a few drift nets being employed. An interdiction existed against the use of seines3 the lakes forming part of the St. Johns; consequently, most of the seining was confined to the channels connecting the lakes. During 1896, 127,865 shad, valued at $8,982 were harvested. In the early part of the season shad were sold to the Palatka shippers as high as 50 cents each, but the price fell rapidly and when the season was at its height the price became frequently less than $7.50 per 100. Sf. Mmys River.-This river has its source in the Okefenokee Swamp, and forms the boundary line between Florida and Georgia for a distance of 175 miles to its entrance into the Atlantic. Stevenson stated, "Owing to its proximity to the St. Johns, and the greater abundance of fish in that stream, as well as to the meager shipping facilities on the St. Marys, very little attention has been given to catching shad in the latter river." The fisheries were confined to the use of a few drifts nets and stake nets by men living between Crandall and King's Ferry, most of the catch being landed at Oakwell, Georgia and King's Ferry, Florida, where it was carted to the nearest railroad station and shipped to distant markets. Few shad were taken with bow nets for home consumption, and very little seining was carried on because of the narrow channel and swift current. The number of men employed in the shad fisheries in 1896 was 110,40 of whom lived on the Florida side of the river and 70 on the Georgia side. During 1896,10,193 shad worth $1,754 were harvested, of which 3,933 were taken by men living in Florida and 6,260 by residents of Georgia. 21 PRESENT-DAYSHAD FISHERIES IN FLORIDA The St. Johns River is still the principal shad-producing area in Florida; only a few shad are taken in the St. Mary's River. Although the shad of the Atlantic Coast has not been taken commercially from the west coast of Florida, the Fishery Landings for Florida, jointly issued by the U. S. F. W. S. and the University of Miami Marine Laboratory, listed the following pounds of shad: February, 1951- 2,775; April, 1951-112. These were probably the Alabama shad, Alosa alabamae, limited to the Gulf Coast (Rivas, 1949), or perhaps not even of the genus Absa (Ralph W. Yerger, in Litt.) In addition, a few Atlantic shad are said to be taken from the Timoka River, near New Smyrna Beach. Freeman (1950) has compiled the only up-to-date information relative to the status of shad in the St. Johns River. He points out, for example, "During the 1940-50 season the first shad caught in Lake George by the haul seine, which operates the year round (though illegal for shad), was on October 13, 1949. Lake George is a large lake through which the northward flowing St. Johns River passes and is located at the southern end of the area most heavily fished for shad. The season opened on December 1, 1949. Usually the season opens December 15 and extends to April 15; however, it closed 'March 31 this season because of the early opening. Fishing is conducted at all hours of the day by the various methods employed in taking the shad. The first method encountered is drift gill nets and beach seines near the mouth of the river. These are not highly signscant in the industry as they have taken an average of approximately 15 percent of the shad caught since 1944." From 12 to 15 drift nets operate between the mouth of the river and Jackson- ville, although drift netting is not considered extensive. About 25 percent of the total catch of shad is from the Jacksonville area. Freeman (1950) continued, "As the shad proceeds up the river it encounters the next fishing area in the vicinity of Palatka. Drifts are employed here and are fished at the beginning and end of each tide. If the shad succeeds in escaping the drift nets, it encounters a section of approximately 15 miles where shad nets are used. These are operated on the ebb or down-tide; hence, they take up where the gill nets leave off.'' From Jacksonville to Pelatka approximately 10 drift nets were in operation during 1950, and about 6-7 were used around Pelatka. South of Pelatka to Welaka at Dunn's creek shad and herring seines are of 2% inch stretch mesh. "There are still two more minor obstacles the shad must surmount before it reaches the spawning areas. One, the haul seines in Lake George, has been there , for many years; the other is a newcomer. It is the newly found sport of with light tackle. It is believed the latter methods take less than five percent of I the catch." Shad also migrate up to Lake Crescent (two or three nets), and south 1 ward to Lake Monroe, where, Freeman stated in conversation, there is reason to : believe that spawning takes place. During 1948 and 1949 Freeman (1950) made spot checks of the catches made I by shad nets in the area extending from Dunns Creek to Welaka. He noted that the percentage of shad has shown an increase every year while the total production of the river decreased. He noted that some shad fishing grounds located close to one another typically produced more shad than others. He declared, "This is ap- parently due to variations in types of bottom and amount of current. Some bottoms are such that it is almost ilnposible for a shad to escape under or around a net, while others leave ample room for escape." Sex ratios indicated the runs are made up of 67 per cent males and 33 per cent fe;males. The average weight and length of both males and females decreased as the season progressed, indicating that the largest ". . . which are also probably the oldest, move into the river first." Freeman further reported that it was the belief of fisherman that ". . . at one time the average shad was much larger than it is at the present. Some fishermen claim that this was due to the introduction of shad fry from some of the northern streams. The fish dealers have attempted to get the Fish and Wildlife Service to bring in fish from the northern areas once again in an attempt to bring the average weight up." A discussion of the wisdom of planting shad fry given later indicates that such measures are not effective. Freeman encountered the usual difficulty in interpreting the age from shad scales. He states, however, "It appears that most of the fish, both males and females, are making their first spawning run." He did not observe spent roe shad and he generally substantiates Stevenson's story that shad never returned to sea after spawning at the head of the river. He advised that a tagging program be instituted in order to determine the number of shad that have lived long enough to make more 22 than one spawning run. ,He concluded, "It is believed the closing of at least two days a week to all methods of fishing for shad would help increase the runs."

SPORTFISHING FOR FLORIDASHAD The shad has been a sport fish around DeLand, on the St. Johns River, since 1942, according to Corson (1951). One fisherman captured nine and seventeen shad, respectively, on two occasions. The sport has increased in popularity during the last few years. In 1948 the U. S. Army Engineers cut a barge canal through what fishermen called the spawning grounds of shad at Blue Springs on the St. Johns River. The river became muddy and shad allegedly moved south, or upstream, to an isolated spot known as Leomon Bluff. The tackle for sport fishing for shad on the ,St. Johns consists of either glass or bamboo rods which are used for trolling. Flyrods are frequently used with good results, particularly with spinning reels. Long lines up to 100 or more feet are used with a small silver spoon for deep trolling. A landing net is essential since shad are frequently lost on landing because of the soft mouth parts. A skiff powered by an outboard motor is run several miles up and downstream, trolling at moderate speed. During March, 1950, an Annual Silver Shad Tournament took place near DeLand and a large number of people participated. Snyder (1949) presented some of the interesting aspects of sportfishing for shad in the St. Johns River. The rise in popularity of shad fishing in the St. Johns River is attested by an article in the New York Times (Wright, 1951), which appeals to the potential tourist in that it describes the sport as a new and exciting attraction. Thus the article points out that DeLand and Sanford are the important shad-fishing areas. It describes the runs of shad that have obviously been gleaned from the accounts of sport and commercial fishermen as follow-s: "Studies of the St. Johns River shad show that they come in four separate runs annually. The first arrives in the early part of November. There is usually a lapse of about three weeks between the departure of one run and the arrival of another. The fourth and final school remains until about the middle of May and, like the others, disappears as mysteriously as it arrives. The November school runs larger in size than those which follow. The fly in the ointment is that sportsmen contend that some are taking too many shad just for their roe. Shad roe during the winter months has become as common as ham and eggs in DeLand and vicinity." This account provides sufficient fuel for speculation as to the ultimate fate of the shad fisheries. The popularization of sportfishing for shad, with a concurr,ent decline in the commercial aspects of the fishery due to changes in the economy of fishermen, may be emphasized by the more articulate and vociferous sportsmen and may lead to a gradual overshadowing of the commercial fishery.

Eco~oMrcsOF FLORIDASHAD Florida fishermen have been greatly influenced by the decline of shad, according to Freeman (1950). Prior to the Civil War shad were not harvested to a great extent in Florida waters, but shortly afterwards New England fishermen began their seasonal migrations in Florida to begin shad fishing from south to north along the niajor shad-producing rivers of the Atlantic Coast. Even the Mattaponi Indians from Virginia took advantage of the early season at the turn of the century by beginning in the St. Johns River and gradually working their way north to the Hudson River. As shad fishing became popular the number of fishermen and the fishing intensity increased, a fact that partly accounts for the tremendous production figures at the end of the 19th cencury. Freeman stated, for example, "Unfortunately, the aniount of equipment operating during the various years is not known, thereby leaving a question as to whether the increase during the years 1889-90 and 1902 and 1908 are real increases in the shad popuIation or increases due to methods of har- vesting. It is suspected that more equipment was largely responsible for this increase and possibly to sonie extent an improvenient in the methods of fishing." Methods of fishing in the St. Johns River have gradually become more effective and the operations are carried on seven days of the week through most of the time the shad are in the river. Since gears are not licensed, the number operating is difficult to establish. Permits are issued to the fishermen, but they arc not required to report their catch. The personnel employed in shad fishing consists of fishermen who operate on catfishes. mullet and crabs after the shad season. The drift nets are usually 300-600 yards long, about 15 feet deep and are fished from sniall outboard 23 or rnboard motor skiffs. The fishermen recognize two peak runs, one in January, the other in March. Immediately alter fishing their nets the fish are taken to one or another of a small number of dealers at Pelatka, MTelaka,or Jacksonville. There the fish are immediatly iced down and shipped to the major market in Philadelphia, Pennsylvania. According to Mr. Freeman, very few shad are sold in Florida; when they are, they are sold for 204 to 256 a pound. Fishermen usually receive an average of about 7d for buck and 14# for roe shad. Prices start off lower early in the season and as markets are expanded they increase. Shortly after shad from northern rivers enter the markets, Florida shad lose ground on prices. Mr. Freeman stated in an interview in 1951 that in general there has been no increase in shad fishing; that is, large numbers of new individuals have not entered the fishery. In some cases a decrease has taken place, especially where fishermen were unable to make sufficient profit above expenses, as was the case during 1940. Stevenson prophesized in 18916 that the spread and growth of the water hyacinth would cause an abandonment of the shad fishery in the Pelatka section of the St. Johns. The plant is prolific even today, but is a source of annoyance rather than an economic hazard to inost fishermen. During windy days drift nets entangle with the floating plant, and fishing efficiency is decreased. The Corps of Engineers have attempted to control the plant by spraying with 2-4-I), but the results have been inadequate. Helicopters and barges for spraying the chemical are being used at Orange Lake by the local sportsmen's association. In the summer of 1950 spraying operations were carried out to rid an area of from 1,800 to 2,300 acres of hyacinth- infested waters, and Freeman (in liit.) thinks that the result was fairly satisfactory. Shad fishermen have supported the work with some enthusiasm. Beginning in 1948, the Florida Game and Fresh Water Fish Commission began investigations in the St. Johns River to determine the effect of on population. Dr. John F. Dequine, chief fisheries biologist of the Com- mission, found that commercial methods of taking freshwater non-game species have little harmful effect on game fish populations under given conditions, and that it was econonlically possible to conduct commercial fishing operations under these "given conditions."

REHABILITATIONOF FLORIDASHAD Florida followed the example of its northern neighbors by joining the shad hatchery school of thought at the turn of the century. In their early reports State authorities pleaded for funds to establish shad hatcheries, since they believed that this measure would replenish the declining production of shad. On several occasions tanks of shad hatched in northern states were shipped for planting in Florida. In some cases the high temperatures caused the fish to die when the cars were at a standstill. According to the Report of the Fish Commission of Florida for 1901- 1902 (1903), two cars of shad fry, aggregating five million were successfully dis- tributed as follows : River Locality Number St. Mary's River At Glen St. Mary 416,866 fry

.-Suwanee ~- ~~-~-- River-- Ellaville 416.666 Aucilla River Aucilla Ochlochonee River Ochlochnee Little River Quincs Chattahoochee River River unction 416,666 Suwanochee River Dupont 200,000 Toms Creek AlexanderviIle 375,000 Ichelucknee River Branford 375,000 Sante Fe River High Springs 375,000 Blue Springs River Juliette 175,000 Anclote River Tarpon Springs 125,000 Hillsboro River Tampa 325,000 Peace River Barton 175,000 Miami River Miami 375,000 From the above list of localities it is obvious that an attempt was made to establish shad runs in niost of the rivers of peninsular Florida. This experiment indicated that stocking with shad fry indiscriminantly in any waterway was a poor rehabilita- tory measure in Florida. In many of the rivers the environmental conditions were apparently unsuitable for shad, because subsequent planting of shad in waters of southern Florida during the early 20th century also failed. On the other hand, shad 24 stocked on the Pacific Coast prospered in strange waters, indicating that shad will propagate in foreign waters that possess qualities suitable for them. Florida authorities wrote in the Third Biennial Report of the Florida Shell Fish Commission for the years 1917-18, that, "Shad ascend to its head waters [of the St. Johns], [and] spawn but little, if any, until they have reached Lake Harney. It is quite evident that the greater portion spawn much further up the river where [it] dwindles into holes of various depths and sizes. Here the fishermen, during the spawning season, gather and with short seines daily drag these holes and thereby not only take the fish, but often take them before they have deposited their spawn, and also drag out and destroy many little shad that may have been hatched, and cover with mud such eggs as have been spawned. The law on closed night fishing for shad, and Sunday fishing, is not enforced. The enforcement of the law is not the duty of the Shell Fish Commissioner but is under the officers of the several counties of the State. The result is that shad fishing . . . is only a small percent of what the Commissioner believes it should be. * $ * " The account further stated that in the lakes constituting a part of the St. Johns River, gill nets are used but the catch is light when compared with that taken by seines which are dragged across but a few of the narrowest points of the river. Authorities argued that the only way they could counteract the abuse of shad fishing was to increase the numbers of the fish by artificial propagation. During 1929-30 shad hatcheries were operated near Lake George. At the Welaka Hatchery 707,000 shad fry were propagated; at the John W. Martin Hatchery, about 25,0100 shad fry were hatched. In the Tenth Biennial Report of the Shell Fish Division of the Department of Agriculture for 1931-1932, it was stated that, "In 1931 the Department erected a new shad hatchery in connection with the pond hatchery at Welaka. This hatchery consists of a battery of fifty-ounce glass jars, vats and fry tanks. Adult shad are caught in nets for the hatchery and are stripped of ripe eggs while alive. They are afterwards transferred to the glass jars in the hatchery. After the eggs have been in the hatchery jars for forty-eight hours mortality has practically ceased. The usual period of incubation is from four to twelve days. As the eggs hatch the 'fry' gently overflow into vats and tanks where they are held for a period of from one to six weeks, at which time they are allowed to overflow into the St. Johns River and begin their journey downstream. . ." The First Biennial Report to the newly organized Florida State Board of Con- servation (Biennium ending December 31, 1934), stated that "At the St. Johns River at Welaka in 1925, bass, bream and shad were successfully produced in quantity." It was described as follows: "A unit has been built and equipped for the hatching of shad. The design suggested by Superinkendent Dunson has not only proven its efficiency but is an object of interest to those who visit the hatchery. It is equipped with 24 fifty ounce jars where shad eggs, stripped when the roe is ripe and in prime condition and adificially fertilized by milt taken from the buck shad, are placed for hatching. Each of the 24 jars will contain from four to five quarts of shad eggs. Each quart represents 28,000 eggs. The Welaka shad hatchery is so constructed that the process of incubation from the time the eggs are placed in the jars, through the different stages of development, to when the eggs incubate and are released in the river may be observed. The tiny fish may be seen to pass from the jars into a receiving trough and to drop from one plane to another until they reach the final descent into the receiving vat from which they are released direct into the St. Johns. * * * . . . the shad hatchery . . . is a guarantee for maintenance of the supply of shad in the river, though great numbers can be taken by the com- mercial industry. During the 1934 season thirteen million shad fry were planted at this point." The report continued, "The output for 1934, due to the fact that construction was in progress, was far below what will be its normal yield. The hatchery delivered to Florida's lakes and streams . . . 16,000,000 shad. 'The fact that Putnam County received the largest allotment from the Welaka Hatchery during the 1934 season is justified by the fact that, but for the Putnam County Board, improvements and developments at Welaka through the use of labor and funds supplied through C W A and F E E A probably never would have been accomplished. A wider distribution of the output from the 1935 season is scheduled. The operation of the Welaka Fish Hatchery from Juiy 1, 1933 to June 30, 1934, for bass, bream and shad purposes, cost the state $12,941.95. The Second Biennial Report (Biennium ending June 30, 1936) stated that the Welaka Hatchery was enlarged with funds from the Federal Government. "On 25 April 21st the Hatchery Superintendent wrote that the shad hatchery had been rebuilt and made four times larger and that the demand for fingerlings for planting was increasing daily." Over 13,000,000 shad fry were hatched and planted. An analysis of the subsequent commercial production of shad during the years following the planting of shad fry and fingcrlings in the St. Johns River points out the ineffectiveness of hatcheries. During the 1925 season approximately 650,000 shad were harvested commercially. At this time thousands of shad fry were released. Three, four and five years later, when the returning adult shad would theoretically be making their way into the Sishery, the production had hardly increased; 1928- ti91,000; 1929-700,000; and 1930-880,000. In 1930 approximately 750,00(l shad were released in the St. Johns; the commercial production for the third, fourth and fifth years was : 1933-479,402 ; 1934-782,000 ; 1935-800,000. During the 1934 season 13,000,000 shad fry were released; 1937-450,400; 1938 commercial production- 343,000; 1939-354,000 shad. The slight increase in commercial production that occurred during 1928 and 11930could have been due to more intensive fishing or some other economic factor, or natural conditions that might have increased recruitment of shad. The subsequent landings listed above show conclusively that planted shad did not contribute to an increase in the shad population. On the contrary, shad production decreased during the years that more and more shad fry were propagated, although the decrease is probably not due to plantings. These facts apparently were considered critically by conservation authorities in Florida, for in 1942 shad hatcheries were discontinued. Aside from the regulations controlling fishing, no current rehabilitatory pro- grams are in operation for shad. Extensive fisheries for other species have been developed in Florida so that shad have become relatively unimportant, it being 19th in number of pounds of the 50 species reported during the 1950 Recapitulation of Food Fish Census (Published by the Florida State Board of Conservation). The efforts of the State Board of Health in the Jacksonville area to clear up domestic sewage and pulp and paper mill effluents during 1947 through 1949 was considered an indirect effort to help reestablish the shad fishery. The Florida Game and Fresh Water Fish Commission is concerned at this time with maintaining the present pro- duction of shad on a sustained basis, and is attempting same by preventing pollution. and by changes in fishing intensity and other aspects of the environment.

COMMERCIALPRODUCTION OF SHADIN FLORIDA One of the problems in determining the annual catch statistics of shad in Florida waters has been ,the difficulty in collecting raw data. Fortunately, records for many years have been preserved. There is the additional problem of deciding how reliable the records are after they have been collected. Furthermore, it is impossible to draw a continuous curve to show the trend of commercial production of shad simply because for certain years such statistics do not occur, although they range from 1873 to date. The records from the 1880's to the 1930's were collected by the U. S. 'Bureau of Fisheries (U. S. B. F.). After the 1930's the Florida State Board of Conservation began collecting its own statistics independent of the Federal agency. Unfortunately, glaring discrepancies in the returns of total poundage occur in some cases. The Florida agency began collecting statistics by requesting that wholesale fish dealers send in a monthly report of the total poundage of every commercial species they sold. For example, during the 1933-34 Biennium the following is recorded: Shad handled by 65 wholesale dealers-479,402 pounds; estimated yield based on reports from 30.8 percent of dealers in Florida-1,556,500 pounds. By 19316 whole- sale dealers were required by law to return adswered questionnaires regarding total poundage of shad handled through their outlet. An estimate of 1,232,972 pounds of shad was recorded, although during 1936 only 440,000 pounds were actually deter- mined directly froin the returns. During the 1931-32 Biennium the State agency recorded 2,754,592 pounds of shad, whereas the U. S. F. W. S. has recorded 1,167,000 pounds. Another discrepancy occurred in the biennium 1929-330 when the State figures were 1,195,000 pounds and the Federal 1,581,OOC pounds. One of the weaknesses in the statistics collected by the Florida State Board of Conservation is the fact zhat the figures are only partial returns from the various fishing areas. The nuinber of counties in which shad are reported as harvested varies annually. For example, in the poundage listed from Florida's returns of shad, the following exhibits an obvious incompleteness: 1939-six counties; 1943-seven counties; 1945-eight counties; 1946-six counties; 1947-six counties; 1948-eight counties; 1949-seven counties; 1950-three counties. Changes in the nuinber of 26 counties reporting catches may be due in part to changes in regulations. It is possible also that fishermen from one county dropped out of the fishery and began work in another county. The trend of annual catches of shad in Florida, particularly in the St. Johns River, has been downward since 1908 (see Figure 8). A cursory examination of production figures based on incomplete statistics readily shows this. The curve begins at a little over 800,000 pounds in 1873, the declines to a little over 250,000 pounds in 1880. This is the period in the early history of the fishery during which: the low production was due to exploration, the introduction of crude fishing methods, and few fishermen (especially native) to exploit the fishery. Beginning in 1888, when the production had reached almost 1,500,000 pounds. the harvests soared to an all time high of over 2,800,000 pounds in 1908. More fishcrnien and greater fishing intensity (with probably more Floridians participating), better fishing gear and snioother operation, a fuller knowledge of the shad's habits and spawning grounds, the popularization of shad as a delicious article of food, and the broad- of the fact that a source was available for shad several months before the runs began in the Northern markets, probably contributed a great deal to the large production. The numbers declined and leveled off from 1918, when about 900,000 pounds were taken, to 1950, when only 300,000 pounds were taken. During the intervening period annual production figures have fluctuated, sometimes more, some- times less. However, it is difficult, if not impossible, to estimate even roughly the ratio of harvested fish to the total natural population of shad on the basis of available information.

YEAR FIGURE8 In the 1918-1950 period a rough estimate of an average of 300,000 pounds of shad was taken annually. However, the fluctuations were accompanied by periods of decline followed immediately by an apparent recruitment in the stocks of fish. The cycle usually is somewhat erratic, but even in small amounts the trend is often mis- interpreted to mean that the actual numbers of shad are either increasing or decreas- ing on a large scale. In reality, however, the production figures may vary con- siderably when the natural population over a long period is relatively stable, in- creasing or decreasing only slightly in response to the effects of natural conditions such as environmental changes, interspecific competition, and available food. In the final analysis the shad production in Florida waters will probably remain at its present general level, and will increase or decrease significantly only under extra- ordinary conditions. 27 FACTORSAFFECTING DECLINE OF FLORIDASHAD Freeman (1950) has discussed some of the popularly assigned reasons for the decline of shad in the St. Johns River. He skates, "The three most probable causes of this great decrease are (I) pollution, (2) overfishing, and (3) destruction of the breeding areas. Other factors which might have some bearing are weather con- ditions, such as temperature or rainfall, and changes in the river due to dredging which might have affected the current, thereby creating confusion in finding the stream in which they were born. Most of the theories can be partially discarded by one means or another, for example, it is almost impossible to blame pollution since the decline started before the factor became serious. At the present there are two paper mills that are being blamed for the poor runs, but both have been installed since the low production period began. There are a group of ships which belong to the Navy's Mothball Fleet anchored at Green Cove Springs. Oil pumped out of the bilges of these ships is another object upon which the shad fisherman heaps his wrath, but these have only been tied up there in recent years. Ellis, et al. (1947) found in their work on the Delaware River that the critical point for young shad as far as oxygen is concerned was 5 p.p.m. A survey conducted by the Florida State Board of Health (1949 and 1950) revealed the oxygen did not drop below this point at any location in the areas where the pollution supposedly affects the runs. It is possible that [domestic and industrial] pollution has had its effect and will increase if the industries and towns continue to add to the amount of waste products dumped into the River, affecting fish life of all types. However, pollution does not appear to be the major cause at the present." Siltation as yet does not appear to be a major factor. "As far as can be determined, the only destruction the breeding grounds have incurred is that done by illegal fishing which occurs to some extent in the areas which are considered the breeding areas, but this is not extensive. The breeding habits of the shad in this river have not been thoroughly checked and observation of com- mercial fishermen has been relied upon for spawning information." Freeman belieyes that the correlation of weather conditions and currents with the abundance of shad is impossible on the basis of present evidence. There are no dams on the River to affect the migrations. Fishermen in the River believe that the Corps of Engineers7 dredging operation have prevented shad from returning to the spawning grounds. Sports fishermen state that a channel cut in 1948 at Blue Springs caused shad fishing to decline so much that the sport was pursued further at Lemon Bluff. Freeman concludes, "By the process of elimination, all but overfishing has been discarded. It is believed that this is the major cause of the decrease that has been noted." 'Some biologists believe that overfishing is not the major cause of shad decline in general and, in any case, Freeman's analysis of the St. Johns situation is incomplete so that argument by elimination is not valid. Freeman infers, as did Stevenson (1899), the possibility that the annual shad harvest is made up largely of virgin spawners which may be indicative of high rate of mortality due to fishing. Cable (1944B) believed that a large number of virgin spawners were harvested in the South Carolina fishery, and stated that many of them die after spawning. There is a possibility that the apparent abundance of virgin spawners in the fishery may be due to two factors: (1) the inability of tech- nicians to read the spawning marks on scales with any degree of accuracy, thereby erring on the age of fish that may have returned to spawn on more than one occasion; (2) the environmental conditions are such in the waters of the Southeastern States that the life span of shad is shorter, and that spawning may then take place only once in a fish's life. No definite information is available on these two points.

SHADREGULATIONS IN FLORIDA Shad regulations, although not very strict, were widely flouted in Florida waters. The regulation for bar and mesh of shad nets in 1915 was as follows : "No person shall place out in the rivers of the State of Florida any gill net for the capture of shad of a less size than two and one-half inch bar from knot to knot, or five-inch stretched from knot to knot. * * *" In addition to disregarding this law, fishermen occa- sionally fished during the closed period from April 1 to November 29, and also ". . . between sundown on Saturday afternoon and sunrise on Monday morning of each week" during which periods shad fishing had been prohibited since 1896. The closed seasons for shad were but slightly modified in 1936. Shad fishing was prohibited between the first day of April and the first day of December of any year. To be in possession of or to transport iced shad between the seventh day of 28 April and the first day of December of any year was aIso prohibited. The weekend interdiction was apparently taken off the list of regulations. At present no fishing is allowed south of Lake George on the St. Johns. According to Mr. B. 0. Freeman, the seasons can be set up or set back two weeks, and a regulation in the Wildlife Code of the State of Florida (1950) states that the Commission has the authority to close any body of fresh water from any type of fishing in an emergency, so that conceivably they could experiment with rehabilitating shad by declaring a closed season to commercial fishermen. In addition, the Commission requires all fresh water fish dealers to report at the end of each month the amount of fish they have sold, or else the next year's license may be denied. The latter regulation is the only means through which the State collects catch statistics of shad. On the lower St. Johns, which comes under the jurisdiction of the Department of Conservation, no license is required and there is no operating means of obtaining a report of shad catches. ANNUAL CATCSl STATISTICS OF SHAD IN FLORIDA Year Pounds Year Pounds* 252.000 * 1934 782,000 1,448,000 * 1935 800,000 *** 2,05'1,000 * 1936 440,OO *** 2,654,000 " 193'7 450,000 *** 1,299,000 * 1938 343,242 ** 1,011,000 * 1939 354,785 ** 1,819,000 * 1940 345,362 ** 2,833,000 * 1941 256,501 ** 964,000 " 1942 323,449 ** 503,000 * 1943 666,348 ** 655.000 *** 1944 810.838 **

* Figures from the published report of the various agencies of the Federal Government. ** Production from statistics gathered by the Florida State Board of Gonservation. *** Pruduction figures estimated from biennial returns of the Florida State Board of Con- servation.

SHAD FISHERIES OF GEORGIA FORMERSHAD FISHERIES IN GEORGIA During Stevenson's (1899) time the shad fisheries of Georgia were rather im- portant. He stated, however, that "Although requiring a greater number of fisher- men the shad fisheries are not as extensive as those of Florida as regards the value of boats, apparatus, etc., and the quantity and value of shad taken." He presented three tables of information regarding the magnitude of the fishery, and divided the various fisheries into the seven major river systems of the State, of which the shad fisheries of the St. ,Mary's River had been described in the section of Florida shad fisheries. Satilla River.-This is the most southerly river in Georgia. It rises in Irwin County, flows 200 miles through a level sandy region, and enters the ocean 18 miles north of the outlet of St. Mary's River. The commercial shad fisheries of the Satilla were of very recent origin in Stevenson's time, dating only from 1894, although prior to that year many shad were taken by bhe river men for their home use. Shad were taken by drift net and sent to Brunswick, Georgia by boat to market, except those used for home consumption. Altarnaha River.-This river is formed by the junction of the Ocmulgef! and Oconee rivers, between Appling and Montgomery Counties, and, after flowing a distance of 150 miles, empties into the ocean a few miles below the town of Darien. The river, with its many tributaries, is situated entirely within Georgia. Its width varies from 150 to 800 feet, averaging about 300. The season is much later in the Altalrlaha than in the rivers south of it, since the temperature of the water is gener- 29 ally lower due to the fact that the river has its source in the mountains, whereas the other streams rise in the sandhills and swamps near the coast. Stevenson (1899) stated that although shad were abundant in the Altamaha, the shipping facilities were so unsatisfactory that the fisheries had never been developed beyond the extent of local consumption. These local fisheries extended from the mouth of the river to the junction of its two principal tributaries. They were most extensive in the vicinity of Darien and Doctor Town. The forms of apparatus used were drift nets, set nets, and bow nets, named in order of their importance as determined by Steven- son. Of 25,030 shad caught during that year, 17,510 were obtained in drift nets, 7,010 in set nets, and the remaining 710 in bow nets. Ocmulgee River.-This river is formed by the union of South and Yellow rivers about 20 miles below Covington, and thence flows about 300 miles to its union with the Oconee. Shad ascended the Ocmulgee as far as Macon, but few were taken above Hawkinsville, and nearly all those caught on the rivers were obtained below Abbeville. The catch in 1896 was reported at 1,502 shad, valued at $632, of which 892, worth $368, were taken in set nets, and 610, worth $2'64, in bow nets. Oeonee River.--This river rises among the hills of northeastern Georgia, the main stream being formed by the union of North and Middle forks, which unite below the town of Athens, from where it flows 280 miles to its union with the Ocmulgee. Shad were seen occasionally as far up as Milledgeville, but very few were caught (principally by means of set and bow nets) above the town of Dublin, 108 miles above the mouth Ogeechee River.-This river, the only one of importance between the Altamaha and Savannah, is located entirely in Georgia, rising in Green County, and after flowing a distance of 350 miles, empties into the Atlantic 38 miles above the entrance of the Altamaha and 16 miles below the niouth of the Savannah. Nearly the whole river is located below the escarpment line, and it drains extensive swamps; con- sequently, the temperature of the water is generally much higher for corresponding dates than in the two adjacent streams, the Altamaha and Savannah. At the Shoals of Ogeechee, 200 miles from the ocean, where the river crosses the Fall Line, there was a wooden dam 225 feet long and 8 or 9 feet high. Four miles above was a second wooden dam 280 feet long and 15 feet high. The foot of these shoals was the upper- most limit of the shad range, and very few passed above Millen, 100 miles from sea. The Ogeechee ranked first among the shad streams of Georgia, surpassed even the Savannah in the number and value of shad taken and yielded nearly twice as many as the Altamaha and tributaries. Owing to the higher temperature of the water, shad ran up the Ogeechee somewhat earlier than in the case of the Altamaha or Savannah, but later than in the St. Johns River. Practically all the commercial fishing was carried on with drift nets in the lower 22 miles of the river, the best fishing being near Harvey's Cut, about 10 miles from the sea. Most of the fishermen were non-residents of the river basin, many coming from Savannah and New England and living in house boats during the shad season. Savannah was the headquarters for the fishermen and principal market for the catch. Sava~znahRiver.-The Savannah has its sources in the mountains of western North Carolina, but the river proper is formed by the union of the Tugaloo and the Seneca rivers at Andersonville, South Carolina. For a distance of 325 miles it forms the boundary line between Georgia and South Carolina and it empties into the sea a short distance below the city of Savannah. It averages about 300 feet in width in the lower 170 miles; then to .4ugusta the width ranges from 500 to 800 feet. Seven miles above Augusta at the Fall Line a dam extended diagonally upstream for 1,000 feet, then 720 feet directly across. During Stevenson's time the dam overflooded during freshets, the water sometimes standing seven or eight feet above its crest, Because of its injurious effect on the shad fisheries, the residents above Augusta complained; in 1883 a fishwap was constructed in the South Carolina end of the obstruction, but it was not used by shad. Shad entered the Savannah River about the first week in January, and by January 20 the fishing season was well-opened and continued until mid-April, when shad became scarce and garfish so numerous and destructive to nets that the fishery was abandoned. The commercial fishermen of Georgia fished almost exclusively with drift nets, catching 51,325 shad in 1896. Set nets, fall traps, and cast nets, which were operated mainly for other species of fish caught a few shad. Residents of South Carolina caught 2,974 shad by drift nets, and a few by cast nets. The total fishery in 1896 was worth $19,236. The principal fishery was in the lower portion of the river along the Georgia shore and tributary to the city of Savannah. Drift 30 nets averaging 350 yards in length and 30 feet in depth (5?k inch mesh) averaged 1,000 shad to the net. From the northern liinit of the Savannah fisheries to the Augusta dam, a distance of 175 miles, the fisheries were prosecuted mainly for local consumption by South Carolinians. An interdiction existed against fishing with drift nets "from the rising of the sun each Thursday until the rising of the sun on each Monday," but no special effort was made to enforce this regulation. Stevenson (1899) stated that several valuable seine fisheries formerly existed on the Savannah below Augusta, but none had been operated during the 187m6-1896 period. He wrote, "Compared with twenty or more years ago, the yield of shad in the Savannah is small, except in the lower end of the river." During the early part of the 19th century shad annually migrated to the headwaters of the Savannah and through the Tugaloo, 49 miles in length, and for a distance of 10 miles up the Talluhah, where their further progress was barred by Talluhah Falls, 3184 miles by the river course from the sea. After 184'6 the dam above Augusta had acted as a barrier to the further ascent of most shad that found their way to that point. Those that passed through the sluices were not sufficiently numerous to warrant commercial fisheries, but they were occasionally taken in gears set primarily for other species a distance of 80 miles or more above Augusta.

PRESENTSHAD FISHERIES OF GEORGIA The waterways utilized by shad in Georgia have remained relatively unchanged during the past half century. In general shad production has declined ta a marked degree, a condition probably due to a number of factors. Increase in industrializa- tion, poor farming practices, and population have necessarily resulted in appreciable pollution and has changed slightly the water character of rivers, but not enough to halt the annual shad migrations. Mr. Fred Dickson, Chief, Fish !Managenlent, during the past two years, and Mr. Joe Stearns, editor of publications, Georgia Game and Fish Commission, have pro- vided a general appraisal of the shad fisheries of the various rivers. No detailed information on the present status of the shad fishery in Georgia is available. They point out that shad fishing has continued as intensely during recent years as during the past half-century, but shad production has been less, although complete catch statistics are not available for recent years. Shad still ascend the same areas in the major rivers to spawn and shad operators harvest the fish at the same localities with almost the same kind of gear and boats as during the early 1900's. Except for the immediate vicinity of towns and industrial areas that have expanded along rivers, discharging sewage and foreign matter into the water, niost shad are taken at the same fishing grounds as yesteryear. The St. Mary's River is considered a relatively productive shad body, and since the water is generally warmer than the rivers immediately north of it, shad fishing begins almost as early as in the St. Johns River. St. Mary's River fishermen have complained about the decline in nunlbers of shad over the years. They state that the river has produced less shad in 1953 than in former years. There is no extensive population on the river to create domestic sewage, and no dams exist. A few drift nets are used, but owing to the rugged nature of the bottom, set gill nets are popu- larly used. A few paper and pulp mills have been established, and anti-pollution measures have been ad~rocated. Although St. Mary's River is technically considered a part of Florida, shad are harvested priinarily by Georgia fishermen. A few are taken by Floridians, but no record is kept by the State of Florida of this catch. The Satilla River is a relatively clean shad area, although the upper tributary receives domestic sewage from the small towns bordering it. Set and drift nets are used. Fishermen today complain that carp, alligator gars, and other predators con- tribute to the decline of shad, but there is no direct evidence that this is true. In addition, siltation has altered the character of the river at certain localities. The Altamaha River, and its two tributaries, the Ocmulgee and Oconee Rivers, are among the iliost important shad streams in the State today, although they are not as clean as in past years. Extensive domestic pollution, siltation, and paper and pulp mill effluents have changed the water quality of the Altamaha system, but not to the degree that shad runs have been stopped. KO dams are present, but the runs of shad up the tributaries have lessened, according to reports from fishermen. During the next year the fish management division of the Georgia Game and Fish Coinmission expects to nlahe extensive investigations on the Altamaha River system with Dingell-Johnson appropriations. Their major effort will be spent in developing control measures to improve the conditions resulting from the polluti-n, and to 3 1 determine factors affecting game and production, with particular emphasis on shad conditions. The Ogeechee River is one of the most important shad producing streams in Georgia, although evidence of some pollution, principally from sewage and siltation, is everywhere. No effort has been made by civilian or industrial authorities to correct such conditions. Dams have not been built during the last 50 years. Except near the larger towns, the shad fishing locations have not changed. Most of the fishing still is concentrated in the lower part of the river by means of drift nets. The Savannah River, according to Dr. E. P. Creaser, former Supervisor of Coastal Fisheries, has virtually no shad now. Sewage and paper and pulp mili effluents are abundantly evident. What fisheries remain still are located in the lower 20 miles of the river and the catch is made by means of drift nets. The Clark Hill Dam recently constructed near Augusta on the Savannah River is above the upper limits of the spawning beds and will not affect shad migrations.

FACTORSAFFECTING DECLINE OF SHAD Shad production has declined considerably in Georgia. Stevenson (1899) analyzed the problem during the late 19th century, and stated that the decrease, in part, was ascribed to the large amount of drift-net fishing in the vicinity of Savan- nah, where the stream is quite narrow, and the amount of twine used therein almost completely obstructed the passage of fish to the spawning grounds. A second cause, according to Stevenson, was found in the limitation of the available spawning grounds. The quantity of muddy water rendered the lower length of the stream unfavorable for spawning purposes, and the dam near Augusta prevented the utiliza- tion of the area above that point, thus limiting the spawning grounds to a few miles just below the Augusta dam, and within this restricted area the eggs allegedly were eaten by the predaceous fish attracted there. In recent years poor law enforcement and overfishing have been considered the reasons for the shad decline, although no objective evidence is available. The fact that nets are not taken out for the three day weekend period has no doubt lowered the escapement of brood stock. Law enforcement authorities in Georgia consider it almost an impossibility to control even a small percentage of the violations of shad laws. Runoff causes extensive siltation, but during the height of shad fishing in spring the waters are clear. Domestic and industrial pollution has increased within the last three decades, and yet adequate measures have not been taken to alleviate these conditions. Anti-pollution programs have been outlined, such as on the St. Mary's River, but no concrete results have followed. Federal authorities (U. 8. Corps of Engineers) are considering several flood control dams on the Altamaha River system within the next few years, and a severe effect on shad migrations and fishing is fpared. The study to be made next year by the Gorgia Fish and Game Commission on the Altamaha River system is especially pertinent, in that it will directly or indirectly concern shad from the following viewpoints: (1) an analysis of fish populations, with an effort to manage effectively the ratio of rough and sport fishes; (2) a tagging program to determine whether Georgia shad are solely virgin spawners, or whether repeaters make up a portion of the catch: (3) factors in the river system contributing to the fluctuation of fish populations. The Shad Investiga- tions group of the U. S. F. W. S. expects to center their activity in the Ogeechee River at some time during 1953 to study the habits of shad.

SPORTFISHING FOR GEORGIASHAD Within the last three years sport fishing for shad has increased in popularity among (Georgians along the Ogeechee River. The sports fishing is carried on mainly during February on the Xing's Ferry Bridge. When the water temperature ranges from 55 to 60" F, fisherinen troll with Barracuda No. 2 spoons, and the usual catch is five to ten shad a day.

Owing to the seasonal nature of shad fishing in Georgia waters, residents must conduct their activities during later winter and spring months, consequently, the majority of the ,fishermen are engaged in the business only temporarily. Many of them are farmers, thus deriving their living from occupations other than fishing. As the season progresses, most fishermen believe that shad increase in size from the early somewhat smaller migrants; then their sizes become smaller at the end of the run. As a result of this belief, fishing intensity is increased during the height of the migration. Few shad are shipped north compared with the larger number locally consumed. During March at King's Ferry Bridge the shad are caught, iced, and offered for sale at a point known as Fish Camp. Although shad ascend the rivers of Georgia for considerable distances, most of the shad fishing is conducted from the mouths to about 30 to 60 miles inland; consequently, many of the shad fishermen are residents in the coastal regions. According to Dr. Creaser, the fish companies regularly subsidize fishermen to the extent of boats and nets and then purchase the catch from them.

Statistics for Georgia fisheries are very disconnected so that only an incomplete picture can be drawn of the general trends. Early in the shad fishery a uniform produotion of about 250,000 pounds (1880-1889) was apparently maintained, proba- bly due to stable fishing conditions and personnel. Beginning in 1890, an abrupt upward turn in statistics indicates a remarkable increase in production, starting roughly at about 400,000 pounds in 1897 and culminating in 1,300,000 pounds in 1908 (see Figure 9). Such an upsurge is difficult to interpret for Georgia, but probably it is closely associated with an activated economy that was part of the State's rapid growth during the pre-World War I days. From 1908 to 1918 no figures are availa- ble, but a drastic decline, 92.4 percent of the 1908 fishery, occurred. That is, only about seven percent of the fishery remained. Between 19118 and 1845, the latest date for which figures are available, the fishery fluctuated a great deal, reaching a high point in 1929 and then declining to less than one-sixth of this high figure in 1939. The later, perhaps somewhat more reliable, figures in which pounds of shad are cited, seem encouraging from a commercial viewpoint, but authorities who have observed the fishery believe the actual catch to be far in excess of this figure. The general fluctuations might suggest biologically inherent cycles in the shad rather than the influence of man on natural populations, but there is no direct evidence of this from research. 1.4 - - #ICOMMERCIAL HARVEST s - / -

8 '.a-- 2 3 - 0 0 - 8 -

3

- REHABILITATIONOF GEORGIASHAD During the last five decades production of shad has declined in Georgia waters, and fishery authorities have sought methods of rehabilitating the fish. The annual yields of shad have fluctuated and the trend has been downward since Stevenson's extensive investigations of the early Georgia shad fisheries. The apparent poor harvests have been blamed on a variety of conditions, and in a sense they have justified the different rehabilitory regulations that were recommended. A discussion of this niatter in the Georgia Department of Game and Fish's Report and Recom- mendations of the State Game and Fish Commission, 1912-13, is as follows: "The shad laws of the State are excellent and can be made effective. They seem to be all that could be desired, although they have been so long a dead letter on the statute books of the State that the most strenuous and harsh means seem to be necessary to their adequate enforcement. The past shad season was disappointing from the standpoint of the shad fishermen. On account of the weather conditions the shad came early, and in much smaller nunibers than the average year, and the fishermen near the coast were not prepared for the enforcement of the provisions of the law enacted many years ago, and the result was disastrous to the business of fishermen who take shad in Georgia waters for the market." An anonymous writer in the Georgia Wildlife Conservation Bulletin (May, 1931, Great Decline in Shad Pro- duetio?~),stated, "Probably no one species of our important commercial fishes has suffered as much from overfishing as the shad. * * * In considering the present status in Georgia we can observe a concrete and most impressive example of the great value and importance of protection and propagation." Unfortunately, during this period no objective investigations were initiated to provide a basis for action. Most fishery authorities grasped for general reasons for the production failure of shad without realizing the more important inlplications in their pronouncements. The Georgia Department of Game and Fish reported that in 1926 a small portable shad hatchery was operated on one of its patrol boats from which it propa- gated about a million baby shad which were planted in the waters of the Satilla and Ogeechee rivers. In addition to this, a note in the Atlanta Journal (March 3, 1936) stated that a 75 acre fish hatchery was built in Bryan County for the artificial propagation of shad and black bass. The fish were to be liberated in the Ogeechee River and adjoining streams. According to Dr. Creaser, this hatchery is not now used for shad. Moreover, between 1875 and 1900 the U. S. B. F. liberated shad fry in Georgia waters. Accounts of these operations may be found in the reports of the Bureau. Although shad hatcheries are considered today to be inadequate for the rehabili- tation of the fisheries, the experience of Georgia fishery authorities seemed to be just the opposite. Actually, the increase in commercial production of shad may have been due to a diversity of factors. The Georgia Wildlife (1931) writer stated, "Just what may have been the practical result of this propagation work is problematical and difficult to prove, but, inasmuch as it is the common belief that the adult shad returns to the waters of its birth, some encouragement is to be found in the fact that the shad production has increased substantially in these particular rivers during the past two or three years, while there has been a continued decrease in the total shad catch of other Atlantic waters. As compared with a catch of 133,750 in 1923, the catch in Georgia for the past three years has been as follows: 1928, 317,267 pounds; 1929, 417,807 pounds; and 1930, 522,212 pounds. The 1928-29 production of North Carolina and South Carolina shows a continual decrease from previous years, whereas the FIorida production has remained practically the same. The Georgia shad is favored in many Eastern markets and is specified on the bill of fare of many of the most popular hotels and other eating places of the country." Today's evaluation of such success in a recruitment of the runs of shad would hardly give credit to stocked shad fry. The same writer stated that among the plans for increasing shad production in Georgia, two things were needed to insure a big shad production: (1) absolute pro- tection of the roe shad in their natural spawning grounds during the spawning period, failure to provide this protection being held responsible more than anything else for the unfortunate plight of shad in Georgia; (2) assistance to nature in re- plenishing the shad supply by artificial propagation. As a fupther management measure, the Georgia Fish and Game Commission shortened the shad season in 1938 to help build the diminishing production (Atlanta Journal, January 14, 1938). "Except for the St. Mary's River, which borders both Florida and Georgia, shad fishing will be permitted only from February 1 to April 1 34 this year, Previously the season extended to April 20. All waters, except St. Mary's, will be closed to shad fishing between sunrise Friday mornings to sunrise Monday mornings." One of the strongest recommendations for the rehabilitation of shad fisheries in Georgia was made in 1945 by Dr. Creaser, who called for a closed season. His statement appeared in Ot,itdoor Georgia (March 1, 1845) in an article entitled "Poor Catches of Shad Lift Prices High"; "Veteran shad fishermen report very poor catches for a late season. As a result of the weak catches, the price of this famous food fish with its delectable roe is expected to skyrocket to a new high of $3 to $3.50 each. Buck shad will bring from $1 to $2, depending on size and weight." The article went on to say, "Dr. E. P, Creaser, . . . called his shot this year by predictiyg many months ago a very poor season and is a strong advocate of a closed season. Georgia shad are sadly in need of a rest,' Creaser said, 'and if they do not get a closed season, the catch will decrease each coming year. Few shad are able to escape the many nets to reach headwaters for spawning.'" He also pointed out that illegal shad operators were not controlled adequately in Georgia. PRESENTSHAD REGULATIOWS IN GEORGIA The present shad regulations in Georgia were instituted in 1943 (See Public Information Bulletin No. 1, 1949-1950), as follows: "The season for taking shad in the St. Mary's River shall be from December 15 to April 15. In all other streams the season shall be from January 1st until April 15. Nets shall be 4% inch or 5% inch-mesh sizes when stretched. Nets shall be set so as to allow one-third of the stream width free for passage for fish. Nets shall not be set within 150 feet of a net previously set. During each week of such open season there shall be a closed time during which no shad shall be taken from the waters of this State in any manner and no shad nets shall remain in or be placed in such waters, beginning at sundown Saturday of each week, and extending until sunrise on Tuesday fol- lowing. There is no closed season on taking shad with rod and reel, fly rod, or pole and line. The daily limit, however, is five shad per person, and they cannot be sold." ANNUAL CATCH STATISTIC'S OtF SHAD IN GEORGIA * Year Pounds Year Po~~nds 1880 252.000 1928 317.000

* Figures from the published reports of the various agencies of the Federal Government. "* Figure from a published report by the Georgia Department of Game and Fish.

SHAD FISHERIES OF SOUTH CAROLINA FORMERSHAD FISHERIES IN SOUTHCAROLINA Stevenson (1899) has left an extensive report on the extent and location of the shad fisheries in South Carolina. At that time he noted "The unusually large number of persons employed for the small number of shad caught . . . the average yield per man being less than 100 shad." He divided the fisheries as follows: Combahee and Ashepoo Rive?-s.-These are small streams located between the Savannah and Edisto Rivers. The former was ascended by shad to Walker for a distance of 85 miles, while the latter was ascended by shad for a length of 50 miles to Walterboro. Set gill and stake gill nets were the principal means of harvesting shad although bow nets were also used. A few shad also were taken in the New, Colleton, and Coosawhatchie Rivers, but not in commercial quantities. Edisto River.-Stevenson stated that the Edisto was second in rank among the shad streams of the State. It is formed by the junction of North and South Forks in the southern part of Orangeburg County, and thence to the ocean it measures 35 l,83miles, although in a straight line this distance is not over 75 miles. There are no falls of note on the Edisto, and aside from the trees and shoals there are no obstructions to the ascent of fish from the mouth of the stream to Davis Bridge on the South Fork, and on the North Fork to Jones Bridge, a distance of 49 miles above Orangeburg and 98 miles from the junction of the two streams. The com- mercial shad fisheries on the Edisto extended from near the mouth of the river to Orangeburg on the North Fork, yet they were most extensive near Jacksonboro, at the crossing of the Charleston and Savannah Railroad. They were prosecuted largely by Negroes from Charleston, but the number of local people engaged had increased during the latter part of the 19th century. Set gill nets were the principal method of harvesting shad, but a few bow nets and haul seines were also utilized. The principal gill net grounds extended from 12 miles below Jacksonboro to two miles above that station. At that time gill nets were used only at night on account of the clearness of the water. This allowed a free passage for shad to reach spawning beds unmolested during the day. The season opened on the Edisto about January 10 and closed the end of March, the greatest portion of fish being taken in February. Charleston Harbor and Tributaries.-Between the Edisto and Santee Rivers are several streams having a common outlet at Charleston, the most important being Cooper, Ashley, and Wando, each of which is small and navigable only for short distances from Charleston. A few shad ascended each of these rivers, but the fisheries were extremely local and limited in extent. Shad were taken by means of gill nets, haul seines, bow nets, and pump nets, a type of bow net. Santee River and Tributaries.-The Santee is formed by the junction of the Congaree and Wateree at a distance from the ocean of 90 miles in a straight line, but following the meanderings of the stream a distance of 184 miles, and was navi- gable its entire length. There were no villages, nor considerable collections of houses along the river, the banks being lined with extensive swamps and forests; hence, during ~Sltevenson'stime the fishery resources of this river had never been developed. Planters and woodmen along the banks of the stream caught a few shad with bow nets and set gill nets for local use. Near the crossing of the Northeastern Railroad at St. Stephen there was some fishing for distant markets. In the Wateree (or Catawba) River, the principal tributary of the Santee, rising among the Blue Ridge Mountains of North Carolina, shad ascended in some numbers as far as Camden, 252 miles from the ocean, and a few were taken at times at the foot of Great Falls, 20 miles farther up. In the Congaree, formed by the junetion of Broad and Saluda rivers, 49 miles above its union with the Wateree, shad were formerly taken in the shoals near Columbia in considerable numbers, and some ascended each of the main tributaries many miles farther. On Broad River a few stragglers ascended as far as ,Green River, 141 miles above Columbia and 28 miles above the boundary line between North and South Carolina, or 374 miles from the ocean. During Stevenson's time the dam at Columbia prevented shad from going farther up the river. In 1883 a fishway was constructed ov,er the Columbia dam, but shad did not appear to use it. Winyah Bay and Tributaries.-During Stevenson's time this region, formed by the confluence of Waccamaw, Pee Dee, Black, and Sampit Rivers, was one of the principal shad-producing locations in the Southern States. The Bay is approxi- mately 12 miles long and four miles wide. From the outer entrance of the harbor to Georgetown, a distance of 14 miles, also for 30 miles up the Waccamaw and at the mouths of Sampit and Black Rivers, the water was well filled with drift nets from the middle of January to the end of March during Stevenson's time. He pointed out that, "The shad fisheries of Winyah Bay are of comparatively recent development, being the result of the tendency in the fisheries on all of th Atlantic Coast streams toward concentration at points nearest the mouth possessing the most convenient shipping facilities, and yet where the river is sufficiently narrow to render very costly apparatus unnecessary." The tributaries of Winyah Bay were all more or less important as shad producing areas. They are described separately below : Waccamaw River.-This river has a length of 149 miles, the lowest 216 of which forms the main channel of Pee Dee River. In its lower portion, below Brook Green, the shad fisheries of the Waccamaw were of much importance, being fished mainly by drift nets. Stevenson stated that in 1896 the greater part of shad were taken near the ends of nets, the fish crowding the river banks rather than following the channel. Above Brook Green shad were taken by means of bow nets as far up as Conway, and occasionally at Red Bluff, 30 miles farther; and a few sometimes were 36 taken even beyond the North Carolina line, over 100 miles from Georgetown by the river course. Pee Dee River and ~ributaries.- his river is known by many names; among them, Waccamaw, Bull Creek, Pee Dee, Great Pee Dee, Little Pee Dee, Yadkin, but generally the name "Pee Dee River" refers to the entire stream above Waccamaw River. It rises on the eastern slope of the Blue Ridge in Watauga County, North Carolina, and flows a distance of about 485 miles. Stevenson conveniently divided the system into three sections: (1) the lower 204 miles, being from the Waccamaw to the lowest dam near Cheraw; (2) from Cheraw to the Narrows, a distance of 71 miles, containing numerous dams and other obstructions to the passage of fish; and (3) the 210 miles above the Narrows, known as the Yadkin. The lower portion of the Pee Dee is swampy, and shad were obtained throughout its length, but in no great abundance at any point. Bow nets, drift nets and haul seines were used to harvest shad. From Cheraw to the Narrows, between which the current is swift, there were numerous fishing dams from one to two feet high. While these dams frequently extended across the stream, there were generally thoroughfares around them, and at high water, which usually prevailed during the shad season, they were partly submerged, permitting many fish to pass over. Numerous sluices were left for the passage of water, in which were placed fall traps and wheels for catching the fish. Very few shad were caught in the traps during the upward run. Shad wheels were introduced in the Pee Dee about 1846, antedating by over 30 years those used in the Columbia River salmon fisheries in Oregon. Stevenson stated that during 1896 between Cheraw and Narrows, a distance of 71 miles, there were 16 wheel and fall-trap fisheries, which more or less effectively obstructed the free passage of fish. All of these were originally constructed to catch shad, but in 1896 not a single shad was taken in the upper six fisheries, and very few in the others. From the narrows to the headwaters numerous fish-dams yielded many shad before Stevenson's time, but up to 1896 had declined rapidly or had been compIetely extirpated. Shad ascended far up the Lynch River to Tilley Ferry; in the Little Pee Dee River, however, none were recorded. Shad were taken as far up the Black River as Mouzans (over 130 miles from Georgetown) where the shad season began about February 10, one month later than in Winyah Bay. In the Sampit River fishermen caught shad for local use. CURRENTSHAD FISHERIES I According to Dr. Robert Lunz, Mr. Alonzo Seabrook, and a Carroll (Charleston) attendant, the bulk of the shad produced today are taken in the Winyah I Bay area. Mr. Seabrook stated that shad are taken in the Pee Dee River, where fish-wheels are still operating as far north as the North Carolina line, but the ,I fish are not taken in commercial quantities more than 40 miles from the coast. The fishery on the Black lRiver and Sampit River seems to be negligigle. It was stated by the fish dealer that shad were taken in some quantities from the Santee River, i but there is reason to believe from the statements of Lunz and Seabrook that these I were largely hickory shad. There are apparently no shad taken in the CharIeston region. They all agreed that shad had declined in the Edisto River tremendously, although it is recognized as a comparatively silt-free and unpolluted waterway. Seabrook stated that shad are still taken up to the Branchville region on the Edisto River. The Ashapoo River was closed in 1936 for a 10-year period to all fishing by a joint agreement between the Federal and State authorities. It was opened to shad fishing in 1946, and according to Mr. Seabrook good catches were made there. Shad are still known to ascend the Combahee River as far as the swamps at the head of the river, but there are no commercial records from this river. According to Seabrook, a few shad are taken on the Savannah River, but the greater number is taken on the Georgia side of the river. In general, the bulk of South Carolina shad are taken from the Georgetown County region. Few shad of any consequence are taken from the Edisto. Shad taken in November and December in the St. Johns River in Florida are transported up to Jacksonboro, where they are sold prematurely for a premium price as "Edisto Shad." Due to budgetary limitations the statistical section of the Branch of Com- mercial Fisheries of the U. S. Fish and Wildlife Service has not been able to make a general canvass of South Carolina shad fisheries since 1945. During that year they noted that the catch of shad by gear was as follows: haul seines-none; anchor gill-18,400 pounds; drift gill-49,500 pounds; stake gill-21,000. The catch by counties in 1945 was as follows: Beaufort-9,300 pounds; Charleston-33,300; Colle- ton-8,200; Georgetown-351,900; Horry-6,300. 37 DECLINEOF SOUTH CAROLINASHAD According to Stevenson (1899) shad fisheries in the Winyah Bay area extended as far up as Bean Shoals, on the Upper Pee Dee, a distance of 382 miles from the ocean. About 18416 the use of dams for catching shad was introduced in the river and it resulted in shutting off the fish from the upper reaches of the river. The stream all along its length is so narrow that it was almost completely obstructed by an amount of twine which .would have had little appreciable effect in retarding the run of shad up the broad tributaries of the Chesapeake or Delaware. Even as early as 1896 shad had declined so alarmingly that Stevenson recommended that artificial propagation would be essential to the prosperity of the fisheries. South Carolina has suffered a marked decline in shad since the peak in 1896, when 146,627 fish were taken. In 1947, only 28,570 fish were taken. In recent decades, authoritative observers in South Carolina state, actually more shad were taken than were reported. If this source of error in the statistics be uniform, then we have a more consistent set of returns that can be interpreted with a certain degree of discretion. The decline of shad may be more apparent than real if several factors are taken into consideration: 1. Some fishermen no longer take shad and do not con- tribute to the local statistics, thus the returns are lower. This would not necessarily mean that shad have been reduced in proportion to the returns. 2. Some fishermen have turned to more lucrative fields and others have not talien their place, as shown by figures on capital investment in gear on Winyah Bay (Lunz, et al., 1944), thus no further decline would be expected. 3. The laws restricting the season and gears (19132-1936)are much more stringent than they have been in the past, and this causes less shad to be taken, thus reflecting a decline in catch, but not actually in numbers of fish. In the past, particularly during the depression (1930-1937), overfishing was considered the real cause of the shad's decline. Fishing intensity was heavy, and many people out-of-work turned to fishing as a method of livelihood. Cable (1944B) has devoted a great amount of research to shad conditions in South Carolina. She stated that, "There has been a steady although, irregular decline of the shad fishery in South Carolina since 1896 when the fishery was de- veloped to its fullest extent . . . it is believed this decline is due to the effects of overfishing. The most important effect of overfishing has been that each year the I escapement of spawning fish from the nets of the fishery has been inadequate to I maintain the fishery at any level of abundance." She declared further that, "The decline of the shad fisheries of South Carolina I has been ascribed to a combination of several causes, the most destructive of which, I and the one cause which can be remedied with profit to all concerned with the fishery, I I is overfishing. Formerly it was thought that pollution from domestic and trade I wastes injured spawn and fry and thus was a major factor in reducing the abun- I dance of the fisheries. Obstructions, in the forms of dams which prevent adult fish from reaching spawning grounds in the upper portions of rivers, have received much blame, also. These factors undoubtedly have played a part in the decline of some of the shad fisheries of South Carolina, but their effects are obscured by the effects of overfishing." 'Terhaps the most convincing proof that overfishing and not pollution, or obstruction of a river-unless very near its mouth-is responsible for the decline of the shad in that stream, is that one of the worst cases of depletion in South Carolina occurs in the Edisto River which is unpolluted and unobstructed. * * * Examination of the daily commercial catch in the Edisto River has shown that approximately 65 percent of the weekly catch in 1938 was made on Monday night, 20 percent on Tuesday night, 10 percent on Wednesday night and five percent on Thursday night. From Friday until Monday, incoming fish made their way up the river and were fished out at approximately the same rate as on the previous week. The fact that most of the week's catch was made the first night of the weekly fishing period indicates that the fishing rate was so high very few fish escaped that first night of fishing. By the end of the week shad were so scarce in the river fishermen declared it was unprofitable to fish. The few fish that did escape the commercial 'fishery in the lower part of the river still had to run the gauntlet of seines and dip nets above the 40-mile limit for an additional month. The fishing intensity in the other rivers of the State was almost as high'as in the Edisto. As a result, only a very small proportion, not more than 10 percent, of the total runs in those rivers was allowed to spawn. Consequently the number of young produced was too small 38 to increase appreciably the population of shad in the various rivers. During the succeeding years before the war, the fishing rates in all the rivers continued to be excessively high." BIOLOGICALEVIDENCE OF SHADDECLINE Cable (1944B) has found that spawning marks on the scales of shad in most northern rivers have proved to be a reliable tool for measuring the escapement in those rivers. "Spawning marks are lacking, however, on the scales of nearly all shad from South Carolina rivers. * '': * The scales from less than one percent of the shad from South Carolina rivers bear a spawning mark. Hence, it appears that in addition to being anadromous like salmon, the shad populations in these rivers also resemble salmon in that many of then1 die after spawning." Further evidence that this is so is seen in the dozens of dead and dying shad which have been observed along the shores of the Edisto River south of Jacksonboro during a single afternoon in the early part of summer. Of the shad that were tagged in the river after the spawning season, not one has been recaptured in a subsequent spawning migration. Occasionally many dead shad are observed in northern rivers during the summer after a prolonged period of intense heat." She also attempted to assess shad escapement from evidence to be gleaned from eggs and larvae collections. Changes in egg abundance reflect changes in the number of spawning fish. "In the Hudson River, N. Y., . . . scale studies and tagging experiments have shown the spawning escapement to be 40 to 50 percent of the annual run of shad, an escapement which appears to be adequate for the mainte- nance of the fishery. If then, for the sake of comparison, an escapement of 50 per- cent of the annual run is considered adequate, we may compare catches of shad eggs and larvae on the same basis . . . collections in the Edisto River in 1940 and 1941, when compared with similar collections from the Hudson River, give startling evi- dence that the present spawning escapement in the Edisto is far short of that needed to build up the fishery. The concentration of shad eggs in the Edisto, as indicated by the catch of eggs per unit of gear, after adjusting for the greater extent of spawning grounds in the Edisto, is about l/s of that in the Hudson River. The spawning escapement in the Edisto, therefore, being 9" of 50 percent is around 10 percent of the run which is wholly inadequate. The fishing rate responsible for a 10 percent escapement is, of course, 90 percent." Cable (1944B) also made the following interesting observations, "It is not known why the shad in South Carolina rivers do not return to the ocean immediately after spawning. They do not take food after entering the rivers, from January to April, nevertheless they tarry on the spawning grounds a month, or even two months, after spawning is completed before attempting to go down to salt water where they might feed again. The fish are then so emaciated they are called 'slabs' by the fisher en. Slabs are often seen floating or swimming feebly with the current. Possibl they are unable to withstand high temperatures when in this weakened conditi n. Whatever the cause of mortality among spawners, very few of them surviv to spawn a second time. For this reason, recovery of the fisheries in South Garoli a will be slower than in rivers where the shad, allowed.to spawn, live to return o the fishery many times if not caught." i POLLUTION Lunz, et al. (1944), coninienting on Cable's report, state ". . . it is also true that pollution in some instances plays a part. Fishermen and dealers have stated that dead young shad often litter the slips and the water under the docks in the Sampit River. These fish were said to have been killed by pollution on their journey to salt water." The Sanitary Engineer of South Carolina reported in the Thirty-ninth Annual Report of the South Carolina State Board of Fisheries (19459, on the following condemned areas for pollution. These areas are as follows: (1) Daufuski Section, half of Savannah River along South Carolina Shores, Mud Creek, Wrights River, and New River; (2) Beaufort Area, portions of Beaufort River, Coosaw River, * Pearson (1944) in his report on shad to the South Carolina Board of Fisheries apparently was not aware of Cable's observations, or else had made subsequent analyses of scales to disprove her statements. He stated, "The fishermen need have no fear that the portion of the run allowed to escape the fishery to spawn wilt he a total loss to them. In fact, most of the escaped fish will return to the rivers the following year as much larger fish. If not caught shad may return many times. Some of the shad of .South Carolina waters population have spawned five or six tlmes. * * " Archers Creek, dattery Creek to Broad River, Chowan Creek; (3) Charleston Area, Cooper River Bridge from Pinopolis Dam to Charleston Harbor, Ashley River from A. C. L. Railway Bridge to Charleston Harbor. Intercoastal water from Hamlin Creek to Charleston Harbor, Charleston Harbor to the Atlantic Ocean; (4) Stono River area; (5) Georgetown Area, including Winyah Bay, Pee Dee River, and Sampit River. In 1944, the Sanitary Engineer stated that, "The increase of pollution in our waters, no doubt has been caused somewhat by the increase in our population, [thus] the actual causes of pollution in our area are from sewerage conditions and the disposal of waste materials from pulp mills, fertilizer factories, etc., and other poisonous materials that find their way into the bays, rivers and sounds."

REHABILITATIONOF SOUTHCAROLINA SHAD Cable (1944B) declared that, "As early as 1869 it was generally recognized that too much gear was being fished and that the shad fishery was in danger of depletion even while the catch of this fish continued to rise. In that year, the State availed itself of an offer from the United States Commissioner of Fisheries to turn over; from the Federal hatchery 'as many shad fry as could be cared for and supported. For a number of years thereafter millions of shad eggs and fry, obtained from Federal hatcheries, were liberated in South Carolina shad streams. These plantings were augmented by the output from a State hatchery erected in 1880. At times when the annual catch of shad increased several years in succession confidence was felt that the increased catches were the result of the plantings, but subsequent declines in the catch have shown that productivity cannot be maintained by this method alone." Stevenson (1899) did not mention hatcheries in his report on the shad fisheries of South Garolina, so that apparently they were non-existent for some time. Dr. Robert Lunz (in litt.) indicates that hatcheries were running before 19316 or 1937, after which time they ceased operation. John C. Pearson, in his shad report to South Carolina State Board of Fisheries (1944), Thirty-eighth Annual Report made the following observations concerning shad hatcheries in South Carolina. "The hatchery at Orangeburg, S. C., demonstrated that shad fry could be reared in ponds with varying degrees of success depending on the size of the ponds, intensity of population, the amount of natural food in the pond and whether other food was intm- duced. The mortality ranged from 50% for fish held two months or less to more than 99% for fish held for six months or longer. The young shad made normal growth in the ponds. Fingerlings that were a month old were 1% inches long, at five months old they averaged about three inches long, the following spring the survivors were from four to five inches long. Mortality of pond reared shad is not unduly high during the first month and the growth rate is very satisfactory. Finger- lings 1% inches long are large enough to school and swim rapidly. At that time they are quite as able to evade their enemies as they are at any time during their first summer, and since the food supply in the ponds is usually insufficient to carry a large number of fingerlings for a longer time, it seems that they should be liberated at that size. The sanctuary of the pond will have served its purpose protecting the young while they were incapable of evading their enemies. I would suggest that as many fish as possible be reared in these ponds and be placed in our rivers at the time mentioned above. Pond rearing of shad fry should not be substituted, in the conservation program, for the reduction of the fishing rate as the principal [sic] for restoring abundance to the fisheries. The pond-raised fish can never be numer- ous enough to supplant natural reproduction, but they can help to hasten recovery where it is already in progress as the natural result of moderate fishing intensity." For several consecutive years Pearson in various reports (1943, 1944, 1945) on shad to the South Carolina State Board of Fisheries recommended, ". . . that you enact a law requiring that shad nets be placed 400 yards apart in lieu of the present law that allows them to be placed 200 yards apart. Unquestionably, the change in this law would permit large quantities of spawning fish to get by and go into the spawning area. It would also reduce the number of fishermen operating in our waters, and in the course of time increase the number of shad." This statute was never accepted by the legislative body of the State. Cable (1944B) recommended in great detail and without reservation the Mary- land Fishery Management Plan for reducing the fishing rate. She stated, "Since 1941, a number of South Carolina's shad fishermen have gone into the armed forces, some have found employment at the Charleston Navy Yard and still others are 40 engaged in construction work at cantonments. Unless new fishermen have taken their places or the remaining regular fishermen are fishing harder than usual, the fishing rate for shad in this state must be somewhat lower than it was from 1938 to 1941. Now is the opportune time to obtain detailed records of the catches, the amount of gear, and the number of fishermen to compare with prewar employment figures, as a basis for a comprehensive management program. These data will show the extent of the present reduction in fishing intensity and will point out any increase which may occur during the post war period. Since shad do not return to the rivers for several years after their first summer there, increased production due to large spawning reserves resulting from reduced fishing rates cannot be observed until the young reach maturity." To date no effort has been made to accept her proposals, and the only "management" measures accepted for rehabilitory purposes was the few changes in restrictions governing the amount and kind of gear a fisherman may use and time of fishing. Dr. Robert Lunz (in litt.) stated that "commercial fisher- men have requested easing the restrictive laws on shad fishing, but apparently no one is particularly interested in rehabilitating them by proper management." In addition to the intensive shad survey made by the U. S. Bureau of Fisheries from 1937 to 1941, and reported upon by Cable (1944B3, the only additional work has been made by the shad investigations group of the U. S. F. W. S. in the Georgetown area during 1951. The U. S. F. W. S. (1952) reported that its shad investigation group in coopera- tion with the State of South Carolina made a study of the shad fishery of Winyah Bay and tributaries. One biologist was stationed at Georgetown, South Carolina, from March 1 to 25, 1951. Daily catch records were obtained from about half of the fishermen of Winyah Bay for this year as well as records of past daily catches. The catches were sampled throughout this period for sex of fish, scales, length, and weights of fish. This and related data has been analyzed and tabulated for future use to determine any changes in the fishery in future years as a result of changes in fishing regulations or intensity of fishing. The present shad fishery authorities of the U. S. F. W. S. have been hesitant to recommend any specific measures. They have reexamined much of the research work concerned with scale analyses, escapement of brood stock, and overfishing of shad and with a new interpretation of this material feel that the present facts do not justify any type of management plan. Their viewpoint is simply that not enough objmtive data is available for any action. Although they do not specifically dis- courage the enactment of restrictive legislation or the inauguration of plans (similar to the Maryland Fishery Management Plan), they do not endorse them because of the lack of facts to justify their acceptance. Some state officials commend this attitude, but others are anxious to put measures into action and also to experiment with management plans that are not inherently harmful. They feel that the latter approach accomplishes two things: (1) it assuages bitter criticism from fishermen and conservationists who are alarmed at the shad decline; (2) it might have a bene- ficial effect on the fishery even though the initial facts would not justify the accept- ance of such plans or laws. On the other hand such plans, although not harmful and perhaps beneficial for recruitment of shad, may have a deleterious long range effect on associated fisheries and predators. This in itself calls for careful, long range study of shad and its ecology.

ECONOMICSOF SOUTHCAROLINA SHAD FISHERIES Cable (1944) pointed out that as early as the interval between 1860 and 1887 the use of an increased number of nets by shad fishermen began to contribute to the "depletion" of the fishery. As is shown later (section on laws), the various regula- tory statutes which arose as a result of this situation have been uneconomic in their effects, especially that of restricting the fishing week to four days. Many com- mercial fishermen believe the regulations are working at cross-purposes, and they refuse to respect the statutes. According to authoritative observers, some fishermen even keep their nets in operation during the weekend closed period. They fail to report the fish caught during this period. Since 1941 and during the recent Korean conflict a number of South Carolina's shad fishermen have gone into the armed forces, while many have found employment in defense operations outside their normal occupation. From a management view- point some biologists have hailed this as a possible method of allowing a greater escapement of brood stock. Since this situation existed during the World War I1 period, it was believed that a noticeable improvement in shad production would 41 result during the 1946-1949 period. Figures that are available do not support this premise. Dr. Robert Lunz has indicated that shad fishermen in general have not suffered appreciably from the decline of shad. They have been cognizant of the loss, but since 1920 and particularly through the 1930's they have switched over to the shriinp fisheries to augment their livelihood. Although they were not active in the shriinp fisheries during its early development they have since derived a major source of revenue from it. Many of them pursue shad fishing to a limited degree, since the shrimp fishery begins during late spring. Lunz, et al. (1944) have pointed out an important aspect of the shad fishery, "That shad fishing is becoming unprofitable, not only because of the decrease in numbers but because, . . . the fishing time is limited, is clearly shown in Winyah Bay. In this area during the 1920-30 period fishermen owned approximately $30,000 worth of equipment as compared to about $5,000 today. During that period there was no limit on the permissible fishing days. Fewer fishermen in the whole shad industry working under a license plan similar to that of Maryland, with good equip- ment and more working days, in addition to rigid enforcement against night fishing and closure of streams, would no doubt work to better advantage. The run of shad is more or less dependent upon weather conditions rather than upon the particular days of the week." Cable (1944B) demonstrated the importance of shad in the economy of of South Carolina. "Although the catch of shad in South Carolina is small compared with that of other sections of the Atlantic Coast, the value to fishermen and retailers is higher than at most other points along our coast. Shad from the rivers of this state have a particularly fine flavor. They are large fish and their roe is firm when they enter the fisheries. They reach the niarkets early in the season and bring top prices which, in normal times, are often 50 to 60 cents per pound. In, 1925, a catch of 79,131 shad sold for approximately $160,000, about two dollars each, in northern markets. In recent years local demands for this fish have exceeded the supply. It was estimated that the 1940 catch of 14,932 shad brought to the fishermen of the state about $11,000, an average price of 75 cents per shad. The price to local consumers usually is from one to two dollars each for roe shad and seventy-five cents to one dollar each for buck shad." For an illustration of the trend of shad production in South Carolina, see Figure 10. Laws GOVERNINGSOUTH CAROLINA SHAD Cable (1944B) sulninarized the South Carolina statutes on shad conservation. She declared, "Laws intended to correct the situation have not accomplished their purpose. Although restrictions have been placed on the amount and kind of gear the shad fishermen may use and his work week has been shortened in order to reduce fishing intensity, the fisheries have not been restored. The shad season, for many years, has opened January 15 (February lsrt, since 1941), when the shad were just beginning to enter the rivers. The commercial fishery, located below the 40-mile limit, is closed on March 25th. By that time most of the shad making up the run for the season have entered the rivers and those not caught by the commercial fishery have already passed into the upper section of the rivers above the 40-mile limit where fishing was permitted until April 25th (April 20th since 1941). At all points, throughout the fishing season, a weekly closed period is in force from sunset on Friday to sunset on Monday leaving a short four day week for shad fishing with three days of enforced inactivity. Spawning in the Edisto River, where observations were made, begins as early as the middle of February in normal seasons, but does not reach significant proportions until the last two weeks in April and the first two weeks in May. The principal spawning, therefore, comes after the rivers are closed to all shad fishing and is affected by the closed periods only to the extent that spawning fish are allowed to escape the fishery." The principal method by which shad production is annually recorded in South Carolina is by means of a $.05 tax levied on each shad caught and sold by a fisherman. Consequently, fishermen must purchase tax tags that must be displayed upon each fish that is not destined immediately for human consumption. Each county clerk maintains a record of all such tags sold each year. Unfortunately, tax tags are also sold for hickory shad for $.02 apiece, and the distinction between the two is not often made. Authorities state that large numbers of shad are taken illegally or are not recorded, consequently the statistics are generally unreliable. Shad buyers and shippers must purchase an operating license for $25.00, and fishermen must pay $.25 for each shad net (100 yards or fraction) ; there is an additional fee for gill nets , ($1.50) and pound nets ($3.00). 1 ANNUAL CATCH ISTATIISTLC.~OF SHAD IN SOUTH CAROLINA Yec~rs Pounds Year Pounds 1880 2088,000 '# 1930 214,0,00***% 1883 260,000 "* 1931 152,001) "*"" 18'87 3866,000 * 1932 123,000 ":k"" 1888 433,O'OO " 1933 219,000 *:k*" 18.8 9 577,000 " 1934 209,000 **"* 1890 5863,000 " 1935 199,000 **** 1896 672,000 * 1936 g7,0,00 *"*" 1897 560,000 " 1937 89,000 *"** 1902. 434,000 * 1938 83,000 *"*** 1907 355,O'OO "** 1939 81,000 ":*** 1908 464,000 *:R* 1940 71,000 "*":k 1909 485,000 **::" 1941 101,000 "**" 1910 550,000 *** 1942 12,5,000*:s"" 1918 1867,000 * 1943 126,000 *"** 1923 184,000 " 1944 10,1,000 "*"" "*:kh 1924 326,000 1945 68,000 "**" ' 89,000 * 1925 396,000 """ 1946 135,oo~**** 1926 330,000 **** 1947 143,000 **** *%"* 19E7 182,000 ' 1948 100,ooo **"" 1928 320,000 "*** 1949 108 O,O,O :k"*" 1929 3160,000 *""* 1950 98~ooo*"*" - * Published figures of U. S. Fish and Wildlife Service. *' State estimate, from Cable (1944). '" Calculated from state tax receipts, from Cable (1944). ***" Annual &port of South Carolina State Board of Fisheries (number of shad converted ta poundage by multi~licationof 5 pounds per shad). '*** Catch records solicited from fishermen by Oliver Duff,from Cable (1944). SHAD FISHERIES OF NORTH CAROLINA FORMERSHAD FISHERIES OF NORTHCAROLINA Stevenson (1899) states that the North Carolina shad fishery was the most important on the Atlantic Coast, basing his position on work done in 1896. For the entire State he gives 6,947 as the number of fishermen employed. Boats were valued at $80,388, drift nets at $6,063, stake nets at $73,874, pound nets at $132,083, seines at $56,561. With the addition of shore installations ($243,882) and a few minor types of gear, the total investment came to $597,757. He indicated that the shad fisheries of the sounds was of recent origin compared to the fisheries on the rivers south of the sounds, dating from about 1873 and receiving their greatest development in the ten years prior to the date of his report. It is evident, however, from other statements in the report that there were fisheries for shad in the rivers flowing into the sounds much further back in time. The fisheries on the sounds were conducted by men from the inland who came down to the islands for that purpose. Stevenson (1899) made a detailed survey of the rivers where shad occurred, and has separated them as follows: The Cape Fear River: Here he gave Smiley's Falls as the practical limit of the shad in the 1890's and stated that dams at Battle and Buckhorn 12 or 14 miles farther up the river were impassable for shad. On the Black River, a tributary, the limit was Taylor's Bridge. On the North East River, another tributary, the run went to Kornegay. Stevenson stated that no special attention was given to shad in the rivers and estuaries between Cape Fear and Cape Lookout but that some were taken in the mullet fishery and ". . . doubtless profitable fisheries could be established." In Pa~nlicoSound the principal fishing stations for shad were the marsh islands and points in the northeast third of the Sound. They were operated both by men from the Banks and men from the "uplands," most of the fishing places being unin- habited at other times of the year. The Neuse was considered the ". . . most important shad stream between the St. Johns and the James." Local fisheries then still existed above Smithfield but the bulk of the catch was obtained within 22 miles above New Bern, and beIow. It was stated that the shad did not run far up the Trent, a tributary. They ascended the Contentnea to Stantonburg above which were abrupt falls. On the Little River shad ascended to Whitely Mills where numerous mill dams obstructed their further progress. Parnlico-Tar:-The shad fishery on the Pamlico-Tar extended to a short distance below Rocky Mount. At Rocky Mount a falls of 20 feet blocked further passage. The bulk of the shad entering Albemarle Sound did so through Croatan Sound. Very few used Roanoke Sound and consequently nets were set mainly in the former. A1bemnrle:-Stevenson stated that Albemarle Sound with its arms, the Pasquo- tank and Perquimans rivers, were among the most important shad fishing areas on the Atlantic Coast. Pound nets were introduced here about 1860-70 and by 1896 seines were disappearing. The muddy waters of the Roanoke River sometimes injured the pound nets and seines, but proved somewhat beneficial to the stake net operations. The commercial fisheries of the Roanoke River were confined to the extreme lower end from the mouth to Williamston. Some fish were found above that point. At Eton Falls above Weldon the river descends 100 feet in 13 miles. The waters of the Chowan were stated to be stained and clear in contrast to the muddy waters of the Roanoke. Most of the fisheries were between the mouth of the River and Harrellsville. In 1938 it was stated (U. S. Bureau of Fisheries, 1938A) that the waters of Albemarle Sound were the most important for shad in the State. Stevenson in his statistics for the shad fishery in 1896 stated that drift nets were the principal gear in the Cape Fear River, and the only kind below the mouth of the ,Black River. Above the Black River seines and bow nets were used. In the Pamlico and Albemarle Sound areas, however, drift nets were unimportant compared with stake nets, seines, and pound nets. Wheels were operated only in the Roanoke. Judged by the number of fish caught the gear for the State as a whole had the following order of importance: stake net (944,000 shad), seines (533,000 shad), pound nets (479,000 shad), drift nets (83,000 shad), and bow nets (56,000 shad). Smith (1907) gives a table showing the shad catch by gear in North Carolina for six different years from 1887 to 1904. This table clearly shows that during this period the importance of seines declined drasticaliy, the importance of gill nets declined considerably, while the importance of pound nets rose from approximately one-twelfth of the catch to about one-half of the total catch. 44 In the Cape Fear River, Stevenson says there were formerly four dams below Smiley's Falls but they had long since disappeared. Above Smiley Falls two dams were said to be in existence but badly broken down in places. The North Carolina Institute of Fisheries Research points out that there are now three dams or locks on the Cape Fear, at Arcadia, Elizabethtown and Tarheel, respectively 50, 70 and 90 miles about the mouth of the river. They are supplied with fishways but shad are not known to use them. Stevenson lists no dams on the Neuse. There is now a dam on that stream at Raleigh which is 150 miles from the coast. There is no information on whether there is a fishway. There apparently have never been any dams on the Pamlico-Tar. Stevenson mentions no dams on the Roanoke and there is apparently none in North Carolina today. However, a hydroelectric installation is now under construction on that river at Bugg's Island, Virginia. References to this developnlent in the reports of the Department of Conservation and Development have concerned the distribution of the power to be produced and no information is available concerning the installation of fishways. No dams are mentioned on the - Chowan. The waters of those North CaroIina streams which enter the ocean in South Carolina are not considered here, since there is no appreciable commercial shad fishery in their North Carolina portions. PRESENTDAY SHADFISHERIES OF NORTHCAROLINA Captain John Nelson, retired Commissioner of ~CominercialFisheries, supplied most of the information on the present extent of the shad fisheries of the state. The Cape Fear River is still an important shad river as in the past. Most of the fishery is in the lower part of the river. The three dams of the main streams are locks and some shad must go through them for they are reported on the river above. How far up shad now run is not clearly known. Captain Nelson stated that shad ascend the Neuse as far as Goldsboro and It was stated by Dr. Eugene Roelofs that probably shad were not found there because the water is too salty and there are no large streams entering these sounds. Con- sequently, Stevenson's expectation in 1896 that shad fisheries might be established there could not be realized. Captain Nelson stated that shad ascend the Neuse as far as Goldsboro and in the Pamlico-Tar as far as Greenville. In there is shad fishing both along the inner side of the "Banks" and also on the mainland side. These fish are believed to enter the Sound through Hatteras and Ocracoke inlets. Smaller inlets I such as Whalebone Inlet are said not to admit shad because they are too shallow. I Captain Nelson stated that there are nets for shad all along the inside of the banks from Hatteras northward. The chief run of shad now comes through Oregon Inlet. They also formerly came through New Inlet, but that entrance is now closed. Oregon Inlet is also closing up but it may be dredged out. The chief North Carolina shad fishery is now in the Albemarle Sound region. Croatan Sound is a location of a very important I shad fishery, and there are nets strung all along Albemarle Sound. The Roanoke is a muddy river and the fishery does not extend above Jamesville. The Chowan is a blackwater river, and shad now ascend it to Winton. The statistical division of the Branch of Commercial Fisheries of the U. S. F. W. S. made their last survey of the extent of the shad fisheries in 1945. The catch of shad by gear is as follows: Common haul seines-61,000 $ounds; long haui seines-4,500 pounds; anchor gill nets-123,300 pounds; drift gill nets-10,500 pounds; stake gill nets-246,000 pounds; pound nets-450,000 pounds; fyke nets- 22,600 pounds. The distribution of the conimercial landings of shad by counties for 2945 will also give an indication of the location and magnitude of individual fisheries. The counties and landings are as follows: Beaufort ...... 9,400 Ibs. Martin ...... 500 lbs. Bertie ...... 18,500 New Hanover ...... 5,000 Camden ...... 1,600 Onslow ...... 5,000 Craven ...... 28,900 Pamlico ...... 11,000 Currituck ...... ,.. 25,000 Pasquotank ...... 414,400 Dare ...... 682,400 Perquimans ...... 36,800 Gates ...... 3,500 Tyrrell ...... 25,00'0 Hartford ...... 3,000 Washington ...... 12,000 H. F. Taylor (1951) has made an extensive survey of the major fisheries of North Carolina from a geographical viewpoint. He noted that although the quanti- ties and values of poundage of fishes could be handled with efficiency by a small 45 number of dealers, "say, a dozen or so," the problem is complicated by the varied configuration of the coastline which stretches about 300 miles in length. He divides the whole coastal region naturally into three productive areas: (1) Northern Production Area extends from the Virginia line inland to Pamlico River, and along the Outer Bank to Ocracoke. Shad are taken principally in western Albemarle Sound and Chowan River. Shad are transported to markets by the Norfolk and Southern Railroad and the Atlantic Coast Line railroads. Railroads are less used in the Southern areas to move fish. (2) Central P~oductionArea extends from Pamlico River in the north to New River in Onslow County, embracing the most important fisheries of the State. How- ever, owing to the generally high salinities of the waterways in this area, few shad are harvested here. (3) Southern Productio?~Area includes Pender. New Hanover and Brunswick counties, but few shad are taken. ECONOMICSOF SHAD FISHERIES OF NORTHCAROLINA The fishermen in the Cape Fear area are full-time fishermen who make their entire living from the fisheries. A commercial fishery is centered in the New River and at Morehead City but this fishery is almost entirely outside-fishing and shad do not figure in the catch. The shad fishermen on the Neuse are mostly farmers who take the fish mainly for their own use in the spring. There seems to be no information of this nature on the Pamlico-Tar. The fishermen on the outer banks and on Roanoke Island as well as those on Albemarle Sound are full-time fishermen and to them the shad fishery represents a very significant portion of their yearly Income. H. F. Taylor (1,951) made a highly intensive study of the economics of fishermen in North Carolina. Although he did not specifically study the shad situation, many of his observations and impressions are valuable in any overall consideration of shad fisheries. As an indication of the relative position of fisheries in the employment picture, he noted that the average number of fishermen was 6,909 (of whom about 1,500 were employed casually) which may be compared with 27,991 farmers and farm managers. "The number of fishermen in North Carolina was about the same in 1940 as it had been in 1890," but fisheries for specific species have changed. Dr. Taylor declares that the wealth produced by agriculture, forestry, and manufacture t results probably from the employment of both labor and a substantial amount of capital, while that produced by the fisheries is mainly produced by labor. The greater I part of the value of fish production is in four counties, Carteret, Dare, Brunswick, and Pamlico. In the first two counties fisheries are the leading source of income. L In commercial fisheries Dr. Taylor further states that the counties with best access to the source are Carteret, Pamlico, Brunswick, Dare and Onslow, while Pasquotank, Beaufort and Craven are well situated as possible primary market centers. By comparing the prices of shad in North Carolina with those elsewhere, it has 1 been found that North Carolina fishermen have consistently received a ht~herprice than the average for that produced elsewhere. Roelofs (1951) stated, . . . shad runs occur here early in the season while the price is high. When the later large runs of the Chesapeake and other northern waters reach their peak, the price of shad drops as much or more than 50 percent, because of the larger supply. But by this time the. bulk of the North Carolina supply has been marketed." This difference, according to Taylor, is in the season of abundance; that is, North Carolina is able to offer shad before they become ypntiful as the season advances further north. He qualified this statement with, . . . it should be noted that the North Carolina production of shad shrank from 16 percent of all shad in 1890 and 1908 to 10 percent in 1930 and eight percent in 1939; the greatest shrinkage in production occurred in the season of normally high price; the price behavior of shad was there- fore somewhat better in North Carolina than it was for the country as a whole." North Carolina's position over the more northerly states in producing shad for market somewhat earlier in spring prompted Dr. Taylor to remark, "Such advantages as these should obviouslv" be uressed for all thev are worth." COMPARISON OF NORTH CAROLINA WITH THE UNITED STATES, EXCLUSIVE OF NORTH CAROLINA, IN QUANTITIES, VALUES, AND PRICES TO FISHERMAN OF SHAD. 1. 1890 1908 1930 * 1940 ** TT-. -.C! TT S TT s u. s. N. C. EX-N. C. N. C. E~-ON:C. N. C. ELN~C.N. C. EX-N. C. Quantity Pounds 5,768,000 35,877,000 3,942,000 23,699,000 1,323,000 13,507,000 897,000 11,295,000 Value Dollars Q 306,015 1,458,000 373,000 1,740,000 233,274 1,644,000 140,462 839,000 Price Per Pound $ .0530 .0407 .0947 .0735 .l760 .I217 .I570 .0742 1 H. F. Taylor, 1951 * Average 1929-1930-1931 C* Average 1938-1939-1940 46 North Carolina's cut-up coastline has caused the fisheries to be so distributed that scarcities in some regions cannot be made up by abundance in others. Dr. Taylo> remarked that shad migrations in the northern section cannot economically be used to fill in the off-season in the Morehead-Beaufort region. This has brought about the existence of a very large number of producer-dealers, each one handling the produce of a small number of fishermen during the limited season, and at a small amount of money value, making for general inefficiency and high costs. In determining the primary distribution of Korth Carolina sea products, Dr. Taylor found that there are two finfish peaks among the annual average shipments to Fulton Market, New York City, of which shad was responsible for the March-April peak. In a recapitulation by species as annual averages for seven years (1940-1946) of shipments to New York from four South Atlantic States, North Caro- lina leads in the production of shad. Thus, the comparison is as follows: AVERAGE ANNUAL RECE1PT:S OF &HAD LN POUNDS AT FULTON MARKET, NEW YORK FRlOM 1940 TtO 19416, INICLUWIVE North Carolina Maryland Virginia Florida 256,261 26,341 56,197 18,860 Taylor (1951) also gives figures showing the comparative economics of the shad fisheries for three periods since Stevenson's report. The figures are given in several different forms. The quantity in thousands of pounds are: for 18189-92 (composite) : 41,645, for 1908: 27,641, for 1929-31 (average) : 14,830, and 1938-40 (average) : 12,192 (These figures are for the entire North Carolina shad catch). He also dis- cusses the factors which affect the price paid to the fisherman, contrasting the behavior of prices between the shad, where the doubling of price has not compensated for the decrease in take, and the lobster, where the decline in take has been accom- panied by a sixfold increase in price. The following figures come from tables in Taylor, pp. 511-533. They show the value of North Carolina shad in comparison to the total value of all food fishes (shell- fishes included with fin-fishes) in the years when canvasses were made: 1880-39.070 1931-14.0% 1887-39.5% 1932-16.8% 1888-38.970 1934-14.7% 1889-29.8% 1936- 8.37. (surpassed by gray trout, I 1890-30.31% mullet and crabs). I 1897-28.0% 1937- 7.4% (surpassed by 5 other : 1902-~22.5% species). 1908-22,l % 193&10.8% (surpassed only by gray 1918-22.6% (surpassed by alewives) trout). ! 1923-2'7.9% 19139-10.3% (surpassed only by 1927-20.7% shrimp). I 1928-216.1% 1940- 8.6% (surpassed by gray trout 19129-19.3% and shrimp). 1930-16.2% 1945- 4.4% (surpassed by 7 species).

DECLINEOF NORTHCAROLINA SHAD H. F. Taylor (1951) found that shad shows signs of "depletion" or biological scarcity in North Carolina. He stated, "The shad is probably at a permanent bio- logical disadvantage and may never be as abundant or important as it was in former years." A rather complete discussion of the possible factors that have contributed to the decline of shad in North Carolina has been advanced by Roelofs (1951). Until 1935 it was the most important food fish of the State in terms of value. Roelofs pointed out that, "The history of the shad decline in North Carolina parallels that of the entire Atlantic coast. Shad were once so abundant as to be sold for fertilizer but the runs became smaller during the period 1800-1850. Stevenson (1899), how- ever, gives records for the Greenfield seine fishery in Albermarle Sound from 1852 to 1896. These records show years of abundance and scarcity throughout the period, with a general increase from 1852 to 1874, a rapid decrease from 1875 to 1878, followed by a gradual increase to 1896, a year in which 50 million pounds were taken from the rivers of the Atlantic coast. It is not known whether the Greenfield records reflect accurately the picture of the entire coast, but they very likely follow the general trends in the coastal fishery. * * * " 47 Roelofs continued the discussion, "Much has been said and written regarding the causes of the decline of the shad fishery. . Overfishing, pollution, and dams are generally considered the three important factors involved. At first, it was thought that overfishing was chiefly responsible; therefore about 1880 a large-scale program of artificial propagation was initiated in an attempt to restore the shad population. Early indications were that the planting of small shad was accomplishing its purpose (a gradual increase in supply of adults from 1880 to 189m6) but in~provementsin fishing methods and greater fishing intensity were not considered in accounting for such increases. Then after 1896, while planting was continued, the shad decreased at a rapid rate. Arti'ficial propagation did little, if any, denlonstrable good; at least it could not maintain the supply." Roelofs has been cognizant of the pitfalls in the theory of overfishing. 'The effect of overfishing, or even the possibility of overfishing, is not completely under- stood." He quoted from Huntsman (1944) who stated that although ovefishing is believed to have resulted in under-replacement of the stock through decrease in numbers of spawning fish, this does not readily occur " since most species have a high reproductive capacity. * * :< Conceivably, overfishing might prevent full re- placement of stock, but it is desirable to have carefully documented experiments to establish the need for restriction of the fishery to assure replacement. * * * we have as yet been unable to learn of a clear, documented case of under-replacement through overfishing for this continent. + * *" Roelofs has further indicated that, "It is also difficult to place the full responsibility for the shad decline on overfishing, in the light of the Greenfield fishery data referred to above. This fishery, as were most other fisheries, was inoperative for four years during the Civil War, 1862-1865 inclusive. If the breeding stock previously had been maintained at too low a level, this four-year period should have provided ample time to increase the stock. Yet the annual production for the five-year period following the war was only 14 per- cent higher than the five year pre-war period (41,746 fish as compared with 36,609). The take for the next five year period (1871-1875) was about the same, but the following five years witnessed a drop to 17,185 fish annually. Production since that time has never approached the period 1857-1&611. Although this does not prove the impossibility of overfishing, it does show that controlled fishing may not provide the remedy desired, since if complete cessation of a fishery for a four-year period gave

I little benefit, controlled fishing could do no more." Roelofs continued, "The effects of pollution and construction of dams are much ! more easily demonstrated and evaluated. Many streams have been ruined for shad by industrial and municipal pollution and by the erection of obstructions of various kinds. Clearing of land has resulted in erosion and silting, rendering streams nn- I suitable for shad spawning. Shad formerly ascended many rivers to their head 1 waters. Stevenson (1899) reports shad fisheries in the early days on the Neuse River above Raleigh, but there are very few rivers today where the shad can reach , the head waters. In other words, shad and civilization are not compatible, and it is doubtful that the shad can ever be restored to the status of colonial days, in spite of the attempts to do so by well-meaning conservationists." In the 1938 report of the U. S. B. F. hatchery at Edenton, the role of predators in the decline of shad in North Carolina was somewhat uncritically discussed. The catfish, carp, and eel were cited as egg predators and it was stated that the stomachs of the first of these were found to be "packed" hard with eggs alleged to be those of shad and herring. Carp were examined and found in a similar condition though chiefly on the upper rivers rather than near Edenton. Eels were reported to pluck netted shad in the manner reported for the Chesapeake. Specific evidence is lacking in this report, and, in any event, many scientists believe that natural predation does not cause any considerable decline in shad populations. The report also cited the views of fishermen on the decline. "Much discussion prevails among fishermen as to the reduction of the annual catch of shad, each com- munity considering its own local problem, largely on surmise. !Many up river fisher- men lay their decline in shad catch to fishermen below them catching the shad before they get to their locality. They refer not only to the fishermen operating down sound and at the inlets, but often to a group of fishermen operating fishing gear of one sort or another fifteen or twenty miles below them, but the farther away the more certain they are that vast quantities are caught which would come to them if they were not caught below. They feel sure that the stream is blocked below, at times com- pletely by the fishermen's nets, which absolutely do not allow any passage of the fish." 48 "The fishermen fishing in the lower waters below the spawning grounds contend that since the spawning grounds are up river these waters should be closed to fishing so that the shad may have an opportunity to spawn, regardless of who it hurts, and that the fishermen along the spawning waters for the most part have other means of livelihood while they must depend entirely on fishing for a living. In addition, they say laws are violated at will in inland waters by sportsmen fishermen and the seines and gill nets sweep the entire river. . ." In reply to our 1951 questionnaire the Institute of Fisheries Research stated that commercial fishermen in Albemarle Sound are interested in promoting a rehabili- tation program. REHABILITATIONOF SHADFISHERIES IN NORTH CAROLINA Smith (1907) has given a broad picture of the history of shad hatcheries in North Carolina waters. He stated that in 1873 under the direction of U. S. Fish Commissioner Baird about 45,000 shad were hatched at New Bern. In 1875 shad hatching was attempted at New Bern by Mr. J. W. IMilner on behalf of the federa1 government, but no noteworthy results were obtained. In 1877 the State began fish cultural operations on its own account in compliance with an act of the legislature. The work was conducted at several points above New Bern but with little success owing to unfavorable seasonal conditions. The shad hatching in 1878 was noteworthy because it was conducted jointly by Mr. Milner representing the U. S. F. C., Col. Marshall McDonald representing Vir- ginia, Maj. T. B. Ferguson representing Maryland, and Colonel L. L. Polk, Com- missioner of Agriculture for Nortrh Carolina. The site of the operations was Salmon Creek at the head of Albemarle Sound and the season was the most successful up to that time, a million fry being produced. Shad hatching was continued by the U. S. F. G. in 1879 at the mouth of the Chowan River. the steamer Lookout beine. emaloved in the work. In 1880 the state constructed a shad hatchery at Avoca andUutil&ed"the eggs furnished by the Capehart seine fisheries at Sutton Beach. It was in 1881 that the steamer Fish-Hawk was first detailed for the same work with headquarters at Avoca and from that time down to a comparatively recent date that vessel was engaged in shad hatching in Albemarle Sound nearly every season. The S~tatecontinued to operate the hatchery at Avoca until 1884. In 1885 the State discontinued all fish cultural work. The , U. S. B. F. established a hatchery at Edenton in 1900. Fish cultural work with shad continued intermittently up until at least 1948 under the direction of W. C. Bunch, I Superintendent. Hatcheries received much support in their heyday, but they were beset with ; problems. U. S. B. F. personnel working at the Edenton hatchery said in a 1938 I report: "The shad spawn 'piece meal' and seldom ripen but a portion of their eggs ! before each deposit of eggs. This is the reason it is so difficult to obtain quantities of eggs for shad hatcheries, and why it would be impracticable to locate hatcheries on the fresh water coastal rivers of the state. The great number of shad that it would be necessary to catch to assure a reasonable number of ripe eggs for hatching sufficient to justify a hatchery is not available. . . . Artificially taken eggs assure protection from certain infections of eggs and protects them against the dangers that beset eggs deposited naturally in river bottoms through natural enemies of all kinds, to be released as fry, assuring a much greater percentage than in nature. This work is valuable and should be pursued wherever sufficient egg take warrants but due to the great difficulty in obtaining eggs for the reasons stated, we must look to the protections of the natural hatch in every reasonable way to increase our output of shad. . . ." According to a statement in an address by R. Bruce Etheridge, Director of the N. C. Department of Conservation and Development, because of the drop of the North Carolina shad harvest below a nlillion pounds in the early thirties the Department determined to attack the problem with great earnestness in the summer of 1936. A conference was held at Morehead City on August 20, 1936 and September 9, 1936. 6. C. Leach, Chief, Division of Fish Culture of the U. S. F. C. addressed a memoran- dum on shad operations in North Carolina to the Commissioner of Fisheries. From this memorandum we learn that the U. S. B. F. had discontinued operations at the Edenton hatchery except on a very limited scale. "The station had been operated by issuing counter-signed permits to the local fishermen for the purpose of catching shad above the railroad bridge at Edenton. This area is the natural spawning grounds of shad at the head of Albemarle Sound. Under the permits the fishermen were to deliver to the hatchery all shad eggs taken from the ripe fish. In many 49 instances they failed to deliver any eggs. This office was therefore of the opinion that if fishing were prohibited in the natural spawning area above the railroad bridge, greater results would be obtained than by hatching a very limited number of eggs at the station. Since the closing of the area to commercial fishing, a few eggs have been taken by seine hauls and these have been incubated at the Edenton Station. . . . It is quite probable that streams entering Pamlico Sound and Onslow Bay are inhabited by shad and that it might be possible to set up temporary hatcheries for the incubation of their eggs. It has been found that green shad eggs cannot be transported successfully for long distances. . . . It is believed that the state should appoint one well qualified man and the Bureau appoint a qualified man to make investigations of the conditions in North Carolina waters relative to the advisability of establishing independent stations. . . ." The memorandum then goes on to state that the Atlantic Coast states would have to take action to regulate the shad industry or there would soon be no shad to regulate. "Some steps must be taken to permit the shad to reach their spawning grounds. It is therefore recommended that there be an annual closed season and that no shad should be permitted to be caught possibly before the 151th of February and that the fishing season be closed shortly after the 15th of March. There should also be a weekly closed period from sundown Friday night until sunrise Monday mornlng. All pound nets must be made inoperative or the traps lifted so that the fish will not be impounded within the wings of the net. Fishing for shad should be prohibited within 20 or 25 miles of the mouth of a stream. It may be necessary to establish closed areas on the Cape Fear, Roanoke, or other streams so as to permit the fish to spawn naturally without being molested. If it is possible for the State to adjust the shad fishery and to enforce such regulations, it is believed that through natural spawning in protected areas and through the establishment of temporary hatcheries at points where the eggs are obtained in sufficient numbers to justify the operations, the shad run can be greatly increased in the State waters." At the meeting of the Board of Conservation and Development held in Raleigh on January 12, 1937, the sum of $1,500 from the Game and Inland Fisheries fund was appropriated for a thorough study of migratory fishes in cooperation with the U. S. B. F. In the address of R. Bruce Etheridge already quoted sotne statements were made concerning progress on regulations. Forty-five days had been cut off of the previous 130 day season. "It was not thought practicable to set aside open days each week because of the fact that it would be almost impossible for the fishermen to raise and lower their nets within the restricted period." Progressive tightening of restrictions over a period of years was recommended. Etheridge mentioned some additional points not considered in the recommendations oT the U. S. B. F.: (1) the role of predators, (2) the enforcement of regulations concerning offshore , which is illegal within the three mile limit for boats not licensed by North Carolina, and (3) alleged obstructions to migration in the form of debris from bridge construe- tlon, etc. Apparently as a result of the activity there were two reports on shad fisheries in North Carolina. Most of the findings and recommendations of the 1938 report have been summarized elsewhere. In December of 1939, Robert A. Nesbit, in charge of Middle and South Atlantic Fishery Investigations for the U. S. B. F. in a publication titled, "How Shad Can Be Brought Back in Chesapeake Bay and North Carolina," stated: " (1) The decline in shad production in Maryland, Virginia, and North Carolina is the result of over- fishing; (2) To correct overfishing of shad the annual toll taken by the fishery should be reduced to 60 percent of the present rate; (3) This should be accomplished by restricting the amount of gear operated so that the fishermen will be spared the uneconomic practice of purchasing and operating more gear than is required to take the annual harvest." Also, Nesbit recommended that no expenditures be made for permanent hatchery facilities and that existing hatcheries be operated strictly on a salvage basis. Follow- ing this report there was little done either from the point of view of further investi- gation or fisheries regulation and this condition has apparently persisted into the post-war years. In reply to a questionnaire sent out in the course of the present investigations, the Institute of Fisheries Research at Morehead City said that the State had at present no plan for shad rehabilitation but the 1951 General Assembly passed a pollution law which might have some effect on the shad. Further, it was indicated that the State contemplates no special shad studies. 59