ORIGIN AND DISPERSAL OF THE ALEWIFE, ALOSA PSEUDOHARENGUS, AND THE GIZZARD SHAD, DOROSOMA CEPEDIANUM, IN THE GREAT LAKES

ROBERT RUS• MILLER Museumof Zoology,University of Michigan Ann Arbor, Michigan

ABSTRACT

It is possiblethat ,the alewife is native 'to Lake Ontario and the gizzard shad to Lake Erie, alth•oughcouclus,ive evidence fo• this 'i.s looking. T.here appear to 'be no recordso.f the alewife from Lake 'Ontario prior to 1873, after the •lake had been stockedwith Banericanshad. T'hat speciesmay still o,ccurin Lake Ontario occasionally.A tabular comparisonof these speciesis given ,to faoilitate identification.The alewife is referred to 'the genus Aloha (rather than Pomo- lobu•) becauseno reliable characteristicsare available,to distinguishthe species of these two groups. Willmot recordeda herring-like fish (not the Banericanshad) from Lake Ontoricoabout 1837 which probably was the gizzard shad, though it may have been the alewife. The first record of gizzard sh,adin Lake Ontario is for 1913, but 'the specieswas rep,oi:tedfrom Lake Erie in 1848, 18 years after the com- p'letion of the first canal •to connectLake Erie and 'the .Ohio River. It is hy- ,pothesizedthat the gizzard shad entered Lake Erie in pre-.Co.lumbiantimes during a warm-dry period. A chronologicalreco,rd of the appearanceo,f alew.ivesin the upper Greaet Lakes shoresthe first specimenreported fro,m Lake Erie was taken in 1931, Lake Huron 1933, Lake Michigan 1949, and Lake Superior 1954; actual dates of first appearancein eaGhlake are unknown,but .the speci,esobviously moved from Lake Erie to the other lakes via Detroit River and Lake St. Clair. The alewife did not beco•ne a'bur•dant in Lake Erie ,or Lake Huron unti'l around 1950 or in Lake Mi,½higanun'til about 1956; it is widely distributedbut not yet abuudant in Lake Superior. The phenomenal increase in albundanceand spreadingo,f .the alewife may be .correlatedwith the upsetin populationbalance by the sea lamprey; .catch s•tatisticsfor *h'e ale•vife and shallow water cisco (Coregonusaraedii) from South Bay, Lake Huron, suggesta direct or indirect causal relationship during 1954-1956. The gizzardshad does not o,ccurnorth o.f lower New York Harbor (and there only rarely) on the A•tlantic slope, al'tho,ughit has been reported from New Brunswick,where it was probablymisidentified for 'the alewife. Its inland range has .beenextended By canal connectionsand perhapsit reachedLake Erie in thi's way; it definitelyentered Lake Michigan throughthe ChicagoRiver canal. A referencein 1874 'to the "saw•belly"from the Great Lakes undoubtedly representsthe gizzard shad,also knoxvn,by that name.The specieshas eviden,fly spreadto Lake Ontario and 'the vicin?tyof Quebecby moving through the WeIland Canal, and to Lake Huron via 'the Detroit River and Lake St. Clair. The 'first specimenknown to b.etaken in Lake Michigan was caughton October 13, 1953, west o• Muskegon,Mich'igan; anotherwas taken near Gary, Indiana, on November 2, 1953. The gizzard shad has not been reported from Lake Superior yahere cold, deep water may prohibit its establishment.It has on ,occasi.ongained some commercialimportance in Lakes Erie and Huron, where 97 98 AmericanFisheries Society

31,600 pounds valued at $930 were taken in 1949. The gizzard shad may have been aided in expandinguorthward by the warming climatic trend of the past half century. The ,t•o species,particularly the alewife, are steadily ir•creaslngin a,bundance and methods of utilizing thi's new resourcesl•ould be devised.

INTRODUCTION

Three speciesof the herringfamily, Clupeidae,which is predominantly marine,have been reporeed from the GreatLakes drainage. The American shad, Alosa sapidissimaWilson, has been i/seedas native ,to Lake On- thattarioit (e.g., first byappeared Hubbsehere and Laglet,when planted 1947, P•y34),the U.S.although Fish others Commission claim in the 1870's (Smith, 1892, pp. 193-194;Evermann and Kendall,1902, p. 213). The specieswas las• recordedfrom Lake Ontario,in 1931, but it occursin 'the lower Ottawa River and stragglersmay occasionallyreach the lake (Radforth, 1944, pp. 94-95). As de•ailledbelow, ehe alewife, Alosa pseudoharengus(Wilson), may have penetrated.the Great Lakes drainagefrom •he St. LawrenceRiver as far as Lake Ontario.It hasbeen ableto surmountthe :barrierof NiagaraFalls 'by passing through the We'l- land Canal'into Lake Erie, from which it has spreadthroughout the Great Lakes.The gizzardshad, Dotosoma cepedianum (LeSueur), probably en- teredthe Great Lakesdrainage natura'lly from •he MississippiRiver basin via Lake Erie and then moved eastward into Lake Ontari,o. Its recent occurrencein Lake Michigan,on the other hand, seemsclearly the result of immigrafi,onthrough the ChicagoRiver canal; it is unknownin Lake Superioror in •he notehemparts of LakeHuron and LakeMichigan. Becausefishery bi.ologists and ochershave difficulty in distinguishing thesethree species, some of ehedistinctive external characters are indicated (Table 1) in order to facilitateidentification, particularly of youngfish. Leim (1924, p. 227, figs. 28-31) gaveadditiona'l characters that distinguish shadfrom alewives,including photographs of earlylife-history stages. An adult gizzardshad (unlessinjured) is easilyrecognized by ehe .greatly lengthenedlast dorsalray, as well as by the small,inferior mouthand deep'body. The positiono• ehe dorsalfin, relativelength of the anal fin, and comparativesize of the upperjaw providefield charactersfor distinguis'hingsmaller specimens .of the three species.Veveebral counts do not overlapbe•veen the a,lewifeand the shador be•veenthe gizzard shadand the shad.As Graham(1956, p. 20) hasshown, popula[ions of the alewife from Lake Ontario have fewer ver[ebrae than do l•hose of Atlantic coastxl streams. Alewifeand GizzardShad 99

T^BZE 1.--•omparison hetu,eenthree specieso] herringsreported ]rom the Great Lakes drainage

Character American shad• Alewife"Gizzard shad s

ast ray of dorsal fin... Not modified Not •nodified Modified into a Ion filament (inconspicuou or absent in yotlng)

rigin of dorsal fin ..... Well before insertion of Well before insertion of Over or behind insertio pelvic fins pelvic fins of pelvic fins

nal fin ...... Relatively short, with Short, with 15-20 rays Relatively long, wit 18-24 rays 25-36 rays

entral scutes...... Typically 22 d- 16 = 38 Typically 19 d- 14 = 33 Typically 18 4- 12 = 3 (range, 34-39) (range, 32-37) (range, 27-32)

•ngitudinal scale rows. 52-64 45-54 52-70

ertebrae (including hypural) ...... 52-60 47-50 48-51

pper jaw ...... Long, slender, its tip Shorter and wider, its Short and moderatel: reaching posteriorly to tip reaching to front of wide, its tip scarcel middleof pupilin young pupil in young and to reaching front of eye i: (25 ram. T.L.) and to rear of pupil in adult; young and front of pupi beyond eye in adult; no notch in adult; a slight notc] no notch (indentation) along ven tral margin in young becoming a deep notc in adult

ower jaw ...... Strong, longer than up- Strong, projecting be- Relatively weak, equa perjaw, not fitting into yond upper jaw, not to (young) or shorte notch in upper jaw fitting into notch in than upper jaw, fittinl upper jaw into notch in upper ja•

[outh ...... Terminal Terminal Subterminal to inferio

illrakers...... Long, slender, about Long, slender, about Very long, fine, close-set 25-75 on lower limb in 20-40 on lower limb in about 90-275 on 1owe young to adult young to adult limb in young to adul

•Ve.rtebral counts t.aken from Vlaxlykov and Wallace (1938); nunxber oi anal rays in p•rt from [,elm (1924, p. 224); g-iill•r•ker counts ap,pvo)zimated from YIilde- brand and Schroeder (1928, p. 94). •-Counts o.rig'i,na2, based on Great L•kes and Onta•vio m•terial (m•ly, exce.pt /'or g'i,llrake•r num!ber which is modified fr•rn YIi•de.b.rand •nd Schro,eder (1928, p. 89). Leim (1924, p. 20.6) re,corded • variati,o,n ot 46 to 51 vertebrae in A_t•,an•ic C(•s,t aJ.ewives.

SAl,1 ao.unts taken from Milllet (1950) except number of g'il.lrakers, which is orlg'inal. 100 AmericanFish eries Society

ALEWIFE The scien,tificname of the alewife is generallygiven as Pomolobus pseudoharengus(Wilson). However,Hildobrand and Schroeder(1928, p. 93) 'indicatedthat Pomolobusand Alosaare scarcelydistinct and Bailey, Winn and Smith(1954, p. 118) gavefurther reasons why it seemsbest to unite them.No clear-cutmorphological characters have 'been proposed to distinguishas a groupthe speciesusually placed in Pomo/obasfrom th,oseassigned to Alosa. Moreover,the name Pomolobztsprobably rests on uncertainnomencatural grounds, for it may be questionedseriously whetherRafinesque's description of P. chrysoch/oris(type speciesof Po- mo/obm)is basedon 'thisgenus and species,or .on the Ohio shad,Alosa a/abamaeohie,sis Evermann (S. F. Hildebrand,unpunished manuscript on the Clupeidaeof the WesternNorth Atlantic). Use of A/osa,the older of the two genericnames, therefore seems preferaNe. There is no conclusiveevidence that the alewife .occursnaturally in LakeOntario. Bean (1884, p. 589) andothers bdieved it wasintroduced there•by mistake in .the 1870'swhen s'had were plan:ted by the U.S. Fish Commission.This has beendenied (see Smith, 1892, p. 188), but evi- dencefor nativity is weak sincethere appearto be neither specimens norrecords of thespecies from Lake Ontario pr'ior to 1873.The possibility that alewives did reach this lake from .the St. Lawrence River should not be dismissed,however. Rad.forth (1944, p. 96) felt 'thatthey may have enteredLake Ontarioduring the invasionof the ChamplainSea, in the later stagesof the glacial historyof the Great Lakes.Toner (1934) pointedout 'that its presencein certainlakes whose outlets pass over barrier falls in Frontenacand Leedscounties, eastern On:ratio, disproves the claim thatthe alewifewas planted in LakeOntario. Possibly the speciesreached its limits of toleration in Lake Ontario and in favorable seasons was able to establishsizxble populations which were decimatedduring inimical years,when presumablyonly the hardiestindividuals persisted. Since around1890, the Mewirehas intermittentlyappeared in Lake Ontarioin almostincredible numbers, as noted during periodi.csummer die-o,ffs (Graham,1956, p. 21). Whethersuch comparative abundance .reflects recencyof introductionis problematical. The historicalappearance of the alewifein the upperGreat Lakes, as evidenced'by capturesknown to the author,is givenin chronological order (Table 2). The first specimenwas takenin Lake Erie in 1931, in Lake Huron in 1933, in Lake Michiganin 1949 and in Lake Superior in 1954.This clearlyestablishes the sequenceof its invasionof these!akes but the datesof first appearancein eachlake are not known.As Dr. M. B. Trautmanpointed ,out (i, litt.), theremust have been thousands of alewivesin Lake Erie in 1931 'beforeone could be caughtin a net. Never- theless,the speciesapparently did not reallybecome abundant there until about1950, althoughcommeroal fishermen caught alewives infrequently, in Ohio waters,after 1940, and therewere similarreports from the Cana- Alewifeand GizzardShad lol

T^BL• 2.--Spread of the alewife in the •pper Great La•ea

Date Locality Remarks •

Sept. 23, 1931 .... L. Erie, at E. end off Adult, 7a• in. long (Dymond, 1932). Nanticoke, Ont.

Mar. 28, 1933 .... L. Huron, 12 mi. E. of Adult, from about 70 fathoms; others Outer Duck Id., Ont. reported (MacKay, 1934).

April 15, 1935 .... L. Huron, 40 mi. from Adult, 188 mm. S. L., from 70 fathoms; Rogers City, Mich. UMMZ No. 109109 (Van Oosten, 1935).

Dec. 26-31, 1940.. L. Erie, off South Bass ld., Adult, 9 in. T. L. taken in gillnet by Ohio K. H. Doan (Trautman, 1957).

Dec. 27, 1942... L. Erie, near Vermillion and Two adults taken in gillnet by fishermen Lorain, Ohio (Trautman, 1957).

May 5, 1949 ..... L. Michigan, W. of S. Adult, 176 mm. S. L., from 60 fathoms, Manitou ld., Mich. Charles Anderson; UMMZ No. 157215.

Mar. 20, 19512 . . . L. Michigan, off Whitehall, Adult, 197 mm. S. L., from 60 fathoms, 14 mi. SW. of Montague, Mich. James Grover; UMMZ No. 160969 (see The Fisherman, Vol. 19, No. 5, 1951, p. 6).

Mar. 12, 1952 .... L. Michigan, 10 mi. ENE. of Adult, 177 mm. S. L., Frank Miller; Milwaukee, Wisc. UMMZ No. 162861.

July 13, 1953 ..... L. Erie, on beach W. of Young-of-the-year taken with gizzard Colchester, Ont. shad (Langlois, 1954, p. 207).

July 14, 1953 ..... L. Erie, South Bass ld., Ohio Young-of-the year (Langlois, 1954, p. 207).

Dec. 22, 19535 .... L. Michigan, 12 mi. WNW. Adults, 180 and 188 ram. S. L., from 32 of South Haven, Mich. fathoms, Chris Jensen; UMMZ No. 166514.

Sept. 25, 1954... L. Superior, off Pendill's Cr., Adult, 168 mm. S. L., Robert Gordon; Chippewa Co., Mich. UMMZ No. 167872.

Nov., 1954 ...... L. Superior, Chequamegon Bay, Adult, 164 mm. S. L., obtained by off Long Id., Wisc. Russell Daly; UMMZ No. 167945.

June I & 14, 1955. L. Superior, Whitefish Bay, Adults, 162 and 175 mm. S. L., from about I mi. N. of mouth of pound net, Ernest L. Pomeroy. Tahquamenon R., Mich.

June 3, 1955. L. Huron drainage: Ocqueoc R. Adults, 179 and 182 mm. S. L., Albert Weir near Rogers City, Mich. E. Hall, Jr.; UMMZ No. 169062.

June 10, 1955 L. Superior, Whitefish Bay, Adult, 197 ram. S. L., sent in by Mich. L. R. Anderson.

June 11, 1955. L. Michigan, East Bay of 19 juveniles, 55-76 mm. S. L., bait dealer Grand Traverse Bay, Mich. for Stanley Lievense; UMMZ No. 170945.

Sept. 14, 1955... L. Huron, along shore in the Fingerlings, about 2 in. T. L., bait dealer; Les Cheneaux Channels, ident. by L. R. Anderson. Mackinac Co., Mich. Oct. 17 and Nov. 12, 1955. L. Superior, Gros Cap area near Adult male (210 mm. T. L.) and adult female Sault St. Marie, Ont. (273 mm. T. L.), ident. by K. H. Loftus and W. B. Scott; ROM No. 18157 (male).

Feb. 13, 1956 .... L. Superior, off Two Harbors, Adult from herring net, Adolph Ojard; Minn. ident. by John G. Hale and Samuel Eddy. 102 AmericanFisheries Society

T^BLE 2--Continued

Date Locality Remarks •

March, 1956 ..... L. Superior, off Knife R., Adult taken by Ivar Pederson; ident. by Minn. John G. Hale.

June 6, 1956 ..... L. Superior, at mouth of Blind Adult, 93-• in. long, caught by George Sucker R., Luce Co., Mich. McGary; 2 other adults also taken. Ident. by L. R. Anderson.

July 31, 1956 ..... L. Superior, Split Rock R. Adult, 161 mm. S. L., from herring net, area, Minn. Ragnivald Sve.

Aug. 31, 1956 .... L. Superior, off Shaganance Id. Adult male, 166 mm. S. L., taken by (near entrance to Black Bay), Melville Gerow; ident. by R. A. Ryder. Ont.

Sept. 10, 1956z... L. Michigan, East Bay of 341 postlarvae to young, 19-40 mm. S. L., Grand Traverse Bay at taken in minnow seine for Stanley Lievense; O-at-ka Beach, Mich. UMMZ No. 171308.

Nov. 5, 1956 ..... L. Superior, Keweenaw Bay, 1 mi. Adult, 181 mm. S. L., from surface gillnet off town of same name, Mich. over 17 fathoms, taken by Uno Weideman.

•Abbr•vio•tio.n.s .are: S.L. s,t&nd,•rd le,ngth; T.L., total length; UMMZ, University of Miehig&n Mtts,eum o,f Z,oo,logY; ROM, l•oy•l O•t•ri,o Museum. =Approximate date. •On April 24, M,ay 10, and eel. 20, 1954, 3 more •du.lts v•e.re t.a&en in 32-50 f•thorn•s by Mr. Jensen from the sanae general area; UMMZ No. 16770•. dian watersof Lake Erie. "After 1950 the youngand adults[alewives] were sufficientlynumerous so that they were distributedrather regularly throughoutLake Erie. In December,1954, theyclogged the gill andfyke netsin the vicini;tyof 'Conneaut,Ohio, followinga badstorm" (Trautman, in litt., Dec. 20, 1956). This informationis supportedby Dr. W. B. Scottwho wrote (in litt., March 5, 1954) tha't "... in the last four or five years'the alewifehas definitelyincreased in numbers5n Lake Erie." The Welland Canal 'has,been available for this highly migratoryspecies since1829, and yet we haveno knowledgeof its occurrencein Lake Erie until 1931 (or of the equallymigratory sea lamprey before 1921). The rathersensational increase in abundanceand spreadof the alewife duringthe pastdecade has probably resulted from a numberof changes particularlyfavorable to .thisspecies. Important modifications in the com- plex interrelationshipsof 'the native fauna have been .brought about by the sealamprey, Petromyzon marinus, and thesedisturbances may have influencedthe change'in alewifeabundance. We do not yet have,however, sufficientbiological information to be a•bleto draw conclusionson this problem.The lake'herring and alewife may compete. for foodand space at somestage in their life cyclessince the adultsrely h,eavilyon animal Alewifeand GizzardShad 103 plankton.Locally, as at SouthBay (Table 3), SaginawBay and Green Bay (informationfrom StanfordH. Smith), the alewifepopulations have recentlyincreased grea'tly while the lakeherring "stocks have decreased. Possibly,newly metamorphosedlampreys migrating across these bays to deeperwater have helpedto reducethe lake.herrings; or, the decrease (particularlynoteworthy in thesebays in 1954, 1955, and 1956) may be a resultof normalpopulation fluctuation. That it has occurredsimu,ltan- eouslyin the three bays,however, suggests some degree of causalrelation- ship, direct or indirect,between the alewife,lake herringand lamprey (and possiblyother species). The alewife.is now abundantin manyparts of Lake Huron and Lake Michigan.During the periodMay 6, 1954, to March 10, 1955, whenthe U.S. Fish and Wildlife Serviceresearch vessel M/V CISCO made thirteen cruisesin the southernpart .of Lake Michigan (mostlysouth of a line joining Milwaukeeand Grand Haven, but north to a line joining Manito- woc and Ludington),no alewiveswere caught.On the otherhand, during nine similar cruisesin the northern half .of Lake Michigan, over the periodMay 10 to November12, 1955, 21 alewiveswere. taken from 10 stations.Nevertheless, by late 1953 the alewife.had movedas far south- ward in Lake Michiganas the vicinityof SouthHaven (Table 2, item 11). Moreover,in a memorandumdated January 2, 1957, Max A. Hunt (Dis- trict FisheriesSupervisor, Plainwell, Michigan) reportedthat "Commer- cial fishermen,fishing out of Saugatuck,report the alewife.is beingcaught in their "chub"nets in up to 43 fathomsof water.There havebeen 200- 300 fish taken in a single.lift. The fishermenare complainingabout the alewife fouling their nets." Saugatuckis a'bout20 miles north of South Haven and well within the southernhalf of Lake Michigan. The authoris indebtedto LeonardS. J.oerisfor the followinginformation on the alewife in the Green Bay area of Lake Mi:chigan.The first indica- tion of spawningwas notedin October,1953, when two or threeschools of small alewiveswere presentin the carp-holdingponds near Duck Creek,Wisconsin (at the southerntip of GreenBay). Additi,onalreliable reportsof schoolsof youngwere receivedin the early fall of 1954 and

T^•-• 3.--Catch of alewife and lake herringin SouthBay, Lake Huron, Ontario (all fish takeh in 1• inch mesh five-pound nets fished in same location)

Year Pounds alewife Pounds herring Number oflifts

1951 ...... 0.2 (1 fish) 36,018 175 1952 ...... 1.0 (5 fish) 14,119 163 1953 ...... 6.0 (51 fish) 12,625 65 1954 ...... 2,316 8,731 97 1955 ...... 16,302 4,841 86 1956• ...... 22,428 3,476 93

•Figure.s for 1956 are provisional. subject to checking, but fina! figurers could vary only a few pounds froIn those given. 104 AmericanFisheries Society

1955. The numberof adultshas increased yearly from a singleindividual in 1952 to commonoccurrence by 1955 in all herringpound nets--yet not in sufficientnumlS•rs to causedifficulty for the fishermen.By the spring of 1956, however,fishermen in the Ellison Bay and Gills Rock area (on the easternside of GreenBay) were becomingquite concerned by the continuingincrease of .the alewife. A steadyincrease in numbers hasalso .occurred for the gillnetfishery along The northern part of Green Bay and Lake Michigan. Alewivesare now abundantin the SaginawBay area of Lake Huron. For example,during operations of the CISCO over the periodMay 3 to October25, 1953, 6 of 9 cruisescaught this species.While trawlingin a very shallowarea near the southwestend of the Bay, about6,000 ale- wives,1.5 to 3.0 incheslong, were takenin a 35-foot trawl in one 5-minute bottomtow, and 8,500 werecaught in a 5-minutetow just off the bottom. Over 50 percentof all fishescaught in trawlingoperations by the CISCO werealewives (information from StanfordH. Smith). Recordsup 'to the fall of 1956 suggestthat the aiewife may not yet have establisheditself in numbersin Lake Superioralthough i.t has rap- idly dispersedthere. The CISCO ran nine cruiseson the lake duringMay 3 to October25, 1953, and took no alewives,although their nets were fishedin someof the regionsfrom which the specieswas obtainedsoon thereafter(Table 2.) Van Oosten(1935) felt that the alewifeproba.bly entered Lake Huron through Georgian Bay directly from Lake Ontario via the Trent River- Trent Canal-LakeSimcoe-Severn River waterway,rather than 'by way of the Detroit River and Lake St. Clair, sincethe specieshad not beenre- portedfrom the lasttwo areas.In response'to my queryregarding this possibility,Dr. W. B. Scottwrote (Jan. 2, 1957) that thereare no records of this species(or of Petromyzon,Osmerus or Anguilla) from Lake Sim- cot, or any other major portionof the waterway,even though the habitat there seemsto be excellentfor all of thesefishes. The proposedroute is a very involved,even .tortuousone, and the water flows throughhydro- electricplants at two marinerailways at the westernend of the system. Failure to take alewivesin Detroit River is not surprising,since organized collectingthere (as by the Michigan Institutefor FisheriesResearch) has been carriedout chiefly in shMlowerwaters than thosewhich the alewife habituallyseeks. Radforth (1944, pp. 95-96) also concludedthat the alewife reached Lake Huron via the Detroit River and Lake St. Clair. It seemsevident that the alewife will becomeincreasingly abundant in the Great Lakes and that utilization of 'this new resource should be en- couraged.The speciesmay well establishitself, by distri•butionin bait' buckets,in suchinland lakes as CrystalLake, Benzie County,Michigan.

•On July 6, 19'56, an alewife 139 min. S.L. was taken by I. A. l•odeheffer and J. 142. Day in Mullet Lake, Cheboygan Co., Michigan (specimen in UMMZ collec- tion). Alewifeand GizzardShad 105

Accordingto RobertHeffner, bait dealerfrom Beulah,it occursnearby in Betsie River, near Frankfort, and at the mouth of Pla,tte River; he has observeddense schools of alewivesat sizesup to 4 inches•t Frankfort. Althoughthey do not survivewell when co14eetedin large numbers,a few do live and it is difficultto sortthem all ou.tfrom 'baitspecies. Ale- wiveshave 'been used in LakeOntario as a readysource of cheapfood as wel'l as for fertilizer, and on 'the Atlantic coastthey are marketedfresh, smokedor pickled.

GIZZARD SHAD Discussionregarding the northernlimit of this species,both for the Atlantic seaboard and for the Great Lakes-St. Lawrence basins, has been frequentlypublished. Early errorsin rangestatements and evidentlyin identificationhave led to subsequentconfusion. Some of the controversy, however,also hingeson the questionof whetherDorosoma is n•tive in the Great Lakesdrainage. A•t leastas earlyas the 1870's,Gill (1873, p. 811) erredby including CapeCod in the distributionof thisspecies. Subsequently, Jordan and Gil- bert (1883, p. 271) and othersperpetuated the error.The authorhas been unableto discoverthe basisfor listing CapeCod as the northernlimit of the gizzardshad on the Atlanticseaboard, 'but the localityhas per- sistedin the literaturealmost to the present(e.g., see.Vladykov, 1945, p. 36). D. ce?edianumis uncommonnorth of latitude40 ø N. in New Jerseyand southeasternPennsylvania, and reachesits northerncoastal limit in New York Harbor,usually called the LowerBay or SandyHook Bay (Brederand Nigrelli, 1934,p. 194; Breder,1938, pp. 23, 28). Hal- kett (1913,p. 46) statedthat .this species has been recorded from St. John River, Fredericton,New Brunswick,and this fo.rmed the basisof the in- clusionof New Brunswickin the range of Dorosomace?edianum by Hubbsand Lagler (1947, p. 34) and by Radforth(1944, p. 58). As pointedout by Vladykov(o?. cit.), the New Brunswickrecord is based uponthe remarksof an unknownauthor, who, in a reviewof Nash's Check List of the Fishesof Ontario, wrote of D. ce?edianum:•' . . .it is a nativeCanadian fish, and was recordedby 'thelate EdwardJack on the St. John River, at Fredericton,N. B.... "The S't.John River is a tributaryof the Bay of Fundyand its mouthlies more than 500 miles northeast of the northernmost known coastal limit of Dorosoma. This recordis thereforeunacceptable; the' fish thoughtto havebeen a gizzard shadvery likely was the alewife,which occurs as far northas Newfound- land and hasbeen confused with the gizzardshad. K'irtland(1850, p. 2) appearsto have'been the first to suggestthat Dorosomaen,tered the Great Lakes(i.e., Lake Erie) via canalconnections, and this belief has been variouslyaccepted and restatedby subsequent authorsduring the succeedingcentury. The prob•e'mis complicatedby the 106 AmericanFisheries Society very differentphysical conditions of Lake Michiganand Lake Erie, the only two of the Great Lakeswhich .offereddirect access to the gizzard shadin glacialor postglacialtimes. More •han a dozenspecies, representing three families,show a distri'bufionalpa•tern in Michiganthat clearlyin- dicatespenetration from the south into southeasternMichigan. These fishes invadedthe Lake Erie Ibasinby utilizing the former outlet (now MaumeeRiver) o• the glacialprecursor of LakeErie, or by way of other natural connectionsacross 'the very low divide betweenthat basin and the MississippiRiver system. Yet not oneof thesespecies is knownfrom the adjacentdrainage of Lake Michigan.I.t seemsevident tha't the colder, deeperwater of that lake.has served as an effective'barrier to thesefishes, as it evidentlyhas, until veryrecently, to the gizzardshad. Thus, although thisspecies was reported 'by Nelson (1876, p. 44) fromthe Chicago River canaland Lake Michigan,,there is no objectiveevidence to, show that zhe gizzardshad established itself in the lake by migrationfrom the Illinois River until recentyears. There are no specimensof this speciesfrom the southerntip of Lake Michiganin .theChicago Natural HistoryMuseum (.or any other collection),and Forbesand Richardson(1919, Map 8) gaveno recordsnearer Lake Michigan .than the junctionof Fox andIllinois Riversin La Sa'lleCounty, Illinois; furthermore, Gerking (1945, Map 8) did not find it in the Lake Michigandrainage of Indiana. Whether Dorosomais native to Lake Erie cannot'be solved conclusively because,to the author'sknowledge, there are neither specimens nor records of gizzardshad from this lake pri.or to thecompletion in 1830of thefirst canalconnecting the Ohio Riverwith LakeErie. The first specimensof Dorosomato be reportedfrom the GreatLakes were four fish recorded by Kirtland(/oc. cit.) in November,1848, from near the mouthof the CuyahogaRiver at Cleveland.He wascertain that this speciesdid not occurin Lake Erie or its tri'butariesin 1840, and hell'eyedthe specimens capturedin 1848represented recent invaders from the south. Kirtland was perhapsoverimpressed 'bythe winterkillof manygizzard shad during a hardfreeze in the Dayton,Canal, since he citedthis as evidence that Doro- soma"was a native of a warmerclimate." He was a•soimpressed with the faillureof fishermento recognizethe gizzard shad near Cincinnati much before 1840 and with their statedbeli,ef that it was an emigrantfrom the south.Klippert (in Jordan,1882, pp. 870-871)also believed that the gizzardshad was not native'to Lake Erie; he wrotethat '•'beforethe con- structionof the Ohio Canali.t wasnot foundanywhere north of the divide, or watershed 'between the lake [Erie] andthe OhioRiver." According to Jordan(op. cit., p. 742), Mr JohnH. Klippertwas % closeobserver and excellentamateur ichthyologist." In November,1877, Samuel Wilmot (1879) co}lec•eda specimen of D. cepedianumfrom Sarnia, Ontario, at 'thesouthern tip .ofLake Huron. It wasidentified by Bairdas Dorosoma cepedianum heter#rum and the author hasverified that it is a gizzardshad (USNM No. 20836). In transmitting Alewifeand GizzardShad 107 this specimen,Wilmot wrotethat "Fishvery similarin appearanceto this one have beenknown in Lake Ontarioand otherof our watersfor many years;I recollect.them forty years ago." This wouldhave been about 1837. Wilmotwas a pioneerfish culturist and supervisor of theCanadian salmon hatcheryat Newcastleon Lake Ontario (Milner, 1874, pp. 25, 58, 228, 231), and he surelyknew the Americanshad; his testimonystrongly sug- geststhat the gizzardshad inh•bited the Lake Ontariobasin as early as 1837, althoughit is possiblethat the 'specieshe referredto mayhave been the alewife.The firstpublishedrecord of the gizzardshad in LakeOntario that I havenoted is for 1913,when Nash reported •he speciesto be very rare (seeDymond, Hart andPritchard, 1929, p. 14). Sinceit is nowknown from an a•tual specimenthat Dorosomaoccurred in Lake Huron in the 1870's,it seemsobvious that the reference.by Milner (1874, p. 36) to the "saw-belly,"identified as Pomolobuscbrysocbloris Rafinesque, actually representsone of the early recordsof Dorosomacepedianum from the Great Lakes (which lake was not specified).This conclusionwas first expressedby Hubbsand Laglet (1947, footnote,p. 34). A'ithoughthe possibilitythat Dorosomafirst enteredLake Erie through canalscannot be ruled out, it is perhapsmore plausibleto regardthis speciesas a pos,tglacialmigrant .into Lake Erie. This view is favoredby Trautman(1957). On the northernfringe of its rangeDorosoma is near the limit of its thermaltolerance, forging northward during a seriesof favorableyears and ,then,under adverse conditions, becoming almost elim- inated(see, for example,Cox, 1896, p. 610). When numbersare killed and fishermensee the speciesfor the first time, theyfail to recognizeit and conclude that it is a recent invader from a more southern dime. Dr. Trautmanwrote me that this has happenedinnumerable times in Ohio sinceKirtland's day. Perhaps the gizzardshad gained access to LakeErie duringa warm-dryperiod in pre-Columbiantimes and then, with the re- trun to presentcondi,tions, it retreatedbut left relictpopu'lations. Doro- somais favoredto establishoutlying colonies because spawning adults enter smallstreams or ditches,provided they are of low gradient,and the young are later abundant,there. In early historictime, there were manyplaces along the divide 'be•Teenthe Ohio and Lake Erie watersheds(e.g., the dose approachof the Maumeeand Wabashdrainages near Ft. Wayne, (Indiana) whereDorosoma could readily have crossed f,rom one drainage to the other.It is alsopossible, as Gerking(1945, p. 33) suggested,that the gizzard s'hadmay have enteredthe glacial precursorof Lake Erie throughthe Ft. Wayne .outlet(Maumee River), but 'thisview seemsless acceptablebecause Dorosoma does not appearto be adaptedto very cold water,such as prevailedduring the meltingof the continentalglaciers in the evolutionof the Great Lakes.The speciesoccurs in the Great Lakes drainageof Indianaonly 'in the MaumeeRiver system'(Getking, op. cit., Map 8). Thereappears to be no needfor nor support'of the hypothesis (Radforth,loc. cit.) .tha,tDorosoma utilized the Mohawk-Hudsonglacial 108 AmericanFish eries Society outlet (or the Chicago.-U'blyoutlet) to enterOntario; if it uilizedglacial waterways'at all, .theMaumee outlet seems to •bethe onlypossible one that it might have entered.Vladykov (o D. cit.) concludedthat this species reachedthe environsof Quebecby descendingthe St. LawrenceRiver from the Great Lakes. Thus the gizzardshad, as perhapswith the alewifein LakeOntario, apparentlyneared 'the limit o.f its tolerancein Lake Erie. Its increaseand spreadin the lower Lakesmay well reflectthe knownwarming climatic trend (Baumand Havens,1956) or maybe a responseto changingbiotic conditionsof the GreatLakes in recentdecades--or both phenomena may be operative.That the specieshas enteredcanals and thus extendedits rangeseems well founded(e.g., seeWright, 1918, p. 544; Greeley,1928, p. 95). The first valid records(based on identified,preserved specimens) of Domomacepedianum from Lake Michiganare' the following:UMMZ No. 165645, an adult, 255 mm. S. L., taken by 'the McNa•bbrothers, .on Oc- tober13, 1953,in a trap net in 14 feet of water•bout 1 mile southof the channelconnecting Lake Michigan and MuskegonLake, west of Muskegon, Michigan.A secondspecimen, an immature.female in poorcondition iden- titled by the authorand then discarded,was obtained by D.C. Erlandson from a poundnet set in 25 feet of water,on November2, 1953,off Miller, 5 mileseast of Gary,Indiana. The evidenceclearly points to a southern origin for thesestocks, since no specimenshave been reportedin the northernhalf o,f Lake Michiganor in northwesternLake Huron. Dorosoma hasthus gradually extended i,ts range into. Lake Huron by emigrationfrom Lake Erie and into Lake Michiganthrough the ChicagoRiver Canal.The gizzardshad has not 'beenreported from Lake.Superior, where low tem- peratureand the absenceof shallowwater may precludeits establishment. This specieshas had a long historyof fluctuatingabundance in Lake Erie b•t it evidentlyhas rather recentlyestablished itself in numbersin certainparts o,f LakeHuron. For example,Bensley (1915) did not record it from GeorgianBay, and whenWalter Koelz collecteda specimenthere late in 1919 (UMMZ No. 55632), fishermeninformed him that the gizzardshad had ratherrecently appeared and that all thosetaken were small.Today the speciesis fairly commonin suitableparts of Georgian Bay. I't was so abunda=tin Detroit River and Lake St. Clair in the fall of 1932 that unemployedpersons caught them for food with dipnetsand hook and line. By October7, 1933, the gizzardshad had movedinland up the SaginawRiver drainageas far as Pine River at St. Louis,Gratiot Co., Michigan (UMMZ No. 101607, 1 adult); this is one of the few recordsof its occurrencewell inlandfrom the GreatLakes and populations havenot establishedthemselves at appreciabledistances inland. An attempt wasmade to establishthe speciesin WhirmoreLa!•e, Washtenaw and Liv- ingstonCounties, Michigan, in August,1935, when the MichiganInstitute for Fis'heriesResearch stocked 124 individuals(1.5 to 15.0 inchesS. L.) Alewifeand GizzardShad 109 from Lake St• Mary's,Ohio. It is estimatedthat 100 adult shadsurvived after oneweek, but if so, successfulrepeoducfion did not occurand the intrcductionfailed (IFR ReportNo. 315; seealso Cooper and Schafer, 1954). Abundancein lakes Erie and Hueon sinc,e about 1940 is indicatedby the commercialcatch statistics (U.S. Fish and Wild'life Serviceand Michi- ganDepartment of Conservation)which show that a totalof approximately 116,000pounds valued at $2,300were taken during the periodfrom 1942 to 1954. The larges'tcatch for a singleyear, 31,600 poundsvalued at $930, was takenin 1949. Occasionally,sudden mortality of vas•numbers (simi,larto that of •he alewifein LakeOntari,o) creates a publicnuisance, as it did at Erie, Pennsylvania,during January-February,1953. (This and other aspectsof the biology of the gizzard shad are summarizedby .me in a paperto 'be publishedby the U.S. Fish and Wildlife Serviceen- titled: "The clupeidfishes of the su•bfamilyDorosomatinae, wi,th special referenceto the gizzardshad, Dorosoma cepedianum.") The gizzardshad is not as widespreadin the Great Lakesas the ale- wife, probablybecause its preferredhabitat is lessextensive, and, excerpt locally,it is not likeyto attainor exceedthe popuation size' of thatspecies. Ie has been utilized in the Great Lakes as a source of food for Grout,swine and cattle,as fertilizer,and it hasbeen steamed and pressed for oil.

ACKNOWLEDGMENTS Splendid cooperationfrom many sourceshas made this 'paper posibl,e.I have tried to recognizeindividually the valua'blehelp given by •i'shermenby mentioning them in the text. The Institute o,f Fis'heriesReseaxch oœ the Michigan Dopar•mentof Conservati,onhas kept me informed o,f new records;Gerald P. 'Cooper,Kiyoshi G. Fukano, Walter R. Cro•we, Stanley Lievense, Leland R. Anderson, and T. H. Durling have been especiallyhelpful. The staff 'of the U.S. Fish and Wi],d]'ife Service's Great Lake Laboratory, particularly James W. Moffett, Ralph Hile, Stan- ford H. Smith, and Leonard S. Joeris, have made avail'able pertinent records and have discussedcritical aspectso,f the manuscrip't.Raymond E. Johns,onand John G. Hale have supp,liedMinnesola records.Jo'l•n Van Oo,stenhas been •of great help with lliterature references.Milton B. Trautman has given me the benefit o• his long experiencewith the a%veife and gizzard shad in Lake Erie and ,hasgenerously alloxved me to draw from material in press. My co,11eag•,e,Reeve M. Bailey. has critically read 'the manuscri'ptand offered valuable suggestions.A ,numlberof ,Cana- dian workers, especiallyW. B. Scott, F. E. J. Fry, John Bud& J. Murray Speirs, Richard A. Ryder, and Kenneth H. Lo,ftus,have fi•lled in recordsfor the .Canadian p,or•ionof the Great Lakes, and Dr. Scott has provi,dedstimulating discussion and useful referencesrenting •o t•heorigin and dispersalo• the alewife. All cf 'these people have my grateful thanks.

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