Synopsis of Biological Data on Smelt Osmerus Eperlanus (Linnaeus) 1758

'AO Fisheries Syhosiso. 78 FRm/S78 (Distribution restricted) SAST - smelt - 1,23(04),003,O1

SY LJ' OF BIOLOGICAL DATA ON SMELT Osmerus epe?kus (Linnaeus) 1758

19i'cpard by T.1. BeInina

FOOD AND AGRICULTURE ORGANIZATION OF THE UNITED NATIONS F ROME, 1969 DOCUMENTS OF THE FISHERY DOCUMENTS DE LA DIVISION DES DOCUMENTOS DE LA DIRECCION DE RESOURCES AND EXPLOITATION RESSOURCES ET DE L'EXPLOITATION RECURSOS PESQUEROS Y EXPLOTA- DIVISION OF FAO DEPARTMENT DES PECHES DU DEPARTEMENT DES ClON DEL DEPARTAMENTO DE PESCA OF FISHERIES PECHES DE LA FAO DE LA FAO

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OT Oceanographic data. 01 Données océanographiques. 01 Sinopsis sobre oceanografía. IT Limnological data. IT Données limnologiques. IT Sinopsis sobre limnología. and et y CART Information concerning fisheries CART Renseignements sur les pêche- CART Información sobre los recursos and resources of certain coun- ries et les ressources de cer- acuáticosvivos de algunos tries and regions (FID/S). tains pays et régions (FID/S). países y regiones (FID/S).

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SYNOPSIS OF BIOLOGICAL DATA ON SMELT

Osmerus perlanua (Linnaeus)1758

Prepared by

T.L BELYANINL

isltuta of Evolutionary Mopho1ogy and Ecology of Animals Academy of Soncoa Mocow, U.S,S.R

FOOD AN]) AOlJIWLUlDii O)GA'lZATION OF TUE UNITED NATIONS ìmiu9 Thoombor1969 PREPARATION OF THIS SYNOPSIS

The preparation of this Synopsis was promoted in view of the need to review the abundant und much scattered information on the biology and exploitation of Osmerus eperlanus which is the main object of a number of f:ishories in the northern hemisphere.

The details set out in this paper are based on data oollected by the author in the course of personal research work on the species and also, on information received from various sources, most of which are listed in the Bibliography.

The author and the editor are grateful to Dr. D,I. Williamson, Marine Biological Station, Port Erin, Isle of Man, U.K., for his editorial work on the synopsis.

Distribution "Current Bibliography" entry

FAO Department of Fisheries Belyanina, T.N. (1969) 16-6B228 FAO Regional Fisheries Officers FAO Fish.Synops., (78)zpag.var. Regional Fisheries Councils and Synopsis of biological data on smelt Commissions Osmerus eperlanus (Linnaeus) 1758 Selector SM Author IN. AN. P. White Sea, North Sea. Baltio, North America. Canada. Northern Europe. USSR. Taxonomy. Morphology. Distribution. Hybridization. Reproduction, Nutrition and growth. Behaviour. Population - structure, density, dynamics. Exploitation. Protection and management. Pond culture. FRms78Smelt iii CONTENTS

Page

I IDENTITY 1:1

101 Nomenclature

1011 Valid flame I

112 Objective synonymy I

1,2 Taxonom i

l21 AfÍinitie I 1q22 Taxonomic status 1

1q23 Subspecies i 124 Standard common names and vernacular names 3

13 Morphology 3

1q31 External morphology 3 1032 Cytomorpho logy 7 1q33Protein epocificity 9

2 DISTRIBUTION 2:1

21 Total area 1

22 Differential distributic-, I

22i Spawn, larvae and juveniles I 222Adults I 23 Determinans of disribution 3han.es

24 Hybridization 2

24i Hybrids; frequency of hybridization; species with uhioh 2 hybridization occurs; othods of hybridization 2q42 Influence cf natural h1r1dization in ecology and morphology*

3 BIONONICS AND LIFE HISTORY 3:1

31 Reproduction I

3l1 Sexualiiy I 312Maturity 313Mating i 3q14Fertilization j 3l5 Gonads j 316Spawnirtg 3 3q17Spawn 3

32Proadult hase 9

321 Embryonic phase 9 3q22 Larval phase 9 3q23Adolescent phane 11

33Adult phase (mature fish) 3:13 3q31 Longevity 3q32 13 Hardineec 13 333 Competitors 13 334 Predators 3o35 13 Parasites, diseases, injuriesand abnormalities 13 1V FRm/S78 3mejt

Page 3.4 Nutrition and growth 3:13

3.41 Feeding 13 3.42 Food 17 3,43Growth rate 17 3,44Metabolism 20

3.5 Behaviour 20

3.51 Migrations and local movements 20 3.52 Schooling 20 3.53 Responses to stimuli 20 POPULATION 4 4: 1

4.1 Structure i

4.11 Sex ratio 4.12 Ago composition 4.13Size composition

4,2 Abundance and density (of population) i

4,21 Average abundance i

4.22Changes in abundance i 4,23Average density* 4.24 Changes in density 2

4,3 Natality and recruitment 2

'1.31 Reproduction rates 2 4.32Factors affecting reproduotion 2 4.33 Recruitment 2

404 Mortality and morbidity 2

4.41 Mortality rates 2 4.42 Factors causing or affecting mortality 3 4.43 Factors affecting morbidity 3 4044 Relation of morbidity to mortality rates*

405 Dynamics of population (as a whole) 3

406 The population in the community and. the eoosystem 3

5 RXPLOITATIOV 5: 1 5.1 Fishing equipsent

5.11 Gears i

5,12 Boats i

5.2 Fishing areas

5,21 General geographic distribution 5,22 Geographic ranges i 5,23 Depth ranges 5.24 Conditions of the grounds* i

Fishing seasons 5.3 i 5,31 General pattern of season(s) 5.32 Dates of beginning, peak and end of season(s) 5.33Variation in date or duration of season 2 FErn 578 Smelt V

Pe ge 54 Fishinc' oserations and results 5:2

54l Effort and intensity 2 5,42 Selectivity 2 5,43 Catches 2

6 PROTECTION AND MANAGEMENT 6:1

6 I Regulatory (legislative) measures

6,11 Limitation or eduotion of total catch 6,12 Protection of portions of population

6,2 Control or alteration of physical features of the environment

621 Regulation of flow* 622 Contrel of water levols* 623 Control of erosion and silting* 6,24 Fishways at artificial and natural obstructions* 6,25 Fish screens* 626 Improvement of spawning grounds* 6,27 Habitat improvement* 6,3 Control or alteration of chemical features of the environrnente

6,4 Contro' or alteration of the biological features of the enviroriment*

6,5 Artificial stocking

651 Maintenance stockjng* 6,52 Transplantation, introduction

7 POND FISH CULTUPE 7:1 71 Procurement of etocks 7,2 Genetic selection of stock*

7°3Spawning (artificial; induced; natural)* 7,4 Holding of stocke 7,5 Pond mama ementfertilization' acuatic .lant control;sto0 )* 7,6 Foods; foeding 7,7 Disease and parasite eontrol 7,8Harvest*

7,9Transport

8 REFERENCES 8i

* As no information was available to the author, those items have been omitted from the text0

FRm/578 Smell; i :1

1. IDENTITY Lower jaw spatulate, Anal rays 11 to14. Midlateral scales 58 to 72. Lateral line in-

I I Nomenclature complete. Adiposo baso about two-thirds of orbit diameter (rarely in large specimens equal 1.11 Valid name to orbit). Orbit diameter two-thirds or loss of caudal ped.unole depth, Proothmoids double. Osmerus eperlanus (Limnaous, 1758). Mosothmoid simple, without other ossifications. Original combination: Salmo planus Parietale completely soparatod by supraoccipi- Linnaeue, 1758, Syst. Nat, (ed. IO), p,310 tal. Pterosphenoide not reaching parasphenoid anteriorly. Slit present between hyomandibu t,12 Objective synonymy lar. Four simple actinonts", (McAllister, 1963). Salmo eerlano-marinus Bloch, 1782, Naturgeech. d.. Fische Deutschlands 1,p.128, Anadromous or landlocked forms in the P1,28, Fig.1. Polar, Pacific and Atlantic Oceans and their drainages. Osmerus operlanusz LacpMe, 1804, Hist, nat,des Poissons, 5, p, 229, Subjective generic synonyms Salmo (Osmerue) Pallas, 1814, Zoogr.Ross.Ae.,3, p, 386 E2lanus Gaimard, 1850-52, Voy. on Islande et Groenlanci, Paris, Atlas, pl,XVIII, fig.2 1.2 Taonomye (type-ope oies: Eperlanus vulgarie Gaimarci).

1,21 Affinities Allosmerus Hubbo, 1925, Proc.,Biol,Soc,Wash., Supragenoric 38,p,53. type-species: Osmerus attenuatus Hubbs). Supeiphylum-Chordata Phylum-Vertebrata Specific Subphylum-.Crani at a Suporciass-Unathostomata Three forms of the boreal smelt have gen- Series-Pincen orally been recognised; O. of the Class-Teleostomi No'th andbiticens, O. dentex of bho Paoific Subclass-Act inoptnrygii nortk into the Arctic and west bo the White Sea, Order-C lupo iforme O, ordax of the Western Atlantic. Following Suborder-Salmomo de i detailed studies of their populations Berg Family-Ocmeridae (1948),.driashov(1954)and other Russian Subfami ly-Osmerinae wers'.o reduced dentex to a subspecies of eperLanus, Some american authors (Kendall, Generic 1927;McKenzie,1958,and others) consider the american smelt ne a separate species, but Osrnrus Linnus, 1758 Syst.Nat. (ed, io) BigeÏow et al.(1963)and MoAllistor(1963), p, 310. Typo specen Salmo er1anu who made a rsirnionO.'the Osmeridao, considoind Linnaous,1758e /L-nmaeua iiI)adopted four that the snuo Osmerus includes only one spoces, sub-divisions of his gstus Sa1rnoone of which O, esrlanus (L). he named :Osmori:, The go'er?o nane Osmerus Linnaeus, 758, wih ype-s»ecies Salmo 1,,22 Ta,xonomio status eno Linnacuc, vaiidct.od in Direction 69 o! the Intor onJ Commiosion on Zoologi-- t r a norpho.apooioo, a polytypio opooioo cal Wononc) aturo,,j with may goog:aphioal cn1000logicalformo. $flfl lnro canine n t1ier cido ci' vomer, somotimeui nocop od by smaller ones, Pala- 1,23 Subspecies tine teeth exilared aneriorly. t'lazilla.ry ox- tondo be obout posterior border of orbit. No Keyo nubopn&eo of Osmerus eperlanue obrino on opcvole or ovboporole, Dictanoc (from 1oAilis1or 1963). from onoub lo oroal oviim ohortor than from dorsal oigm o orwsl bacs, A midlateral 1(2) Po-od acabo in the lateral lino usually rido, buno anal shelf or e1onga'ed midlate- 14-'28 (roroly 13.-30). Ucotorn Atlontie, Pacif- rol coalos in moie. Gill-rakern 25 to37. ic c-nd koobio uot to thehito Soc.c-nd. their Pylorie cocoa3be 8 Stomach itti blind sao. ctoJaçon , , . n , n . oQo .Q . . . , ' .oQ Q O n , o e e e . e o Orirnorun2?!)US.. ïU!2 (Miioh1b)

*After Berg (1948)and McAllister (1963) i 2 FPm/578 Smelt

2(1)Pored scales in the lateral line usually Medium sized canines on dontary, enlarging 6-13(rarely 0--16). Baltic and North seas and posteriorly. 5mal]. pointed teeth on premaxil- tbir drainages and Upper Volga River iary and maxillary. Canines on anterior end 0vris eperlanus eperlanue (Linnasue) of tongue enlarged. Pelvic origin anterior to dorsal origin. Ductus pneumaticus attached to Osmerus rlanus mordax (Mitchill)1814* anterior end of gas bladder, Latera]. line in-. complote, ceasing about a head's length along Atherina mordax Mitohill,1814, Po 15, the body. Ìou York (Speirs1951,indicated that this was tho type description)0 Peritoneum light with dark speckles, more intenso dorsally. Ventral portion of body Osmerus viridescens Le Sueur,1818, p. 230, pale; dorsum speckled with black, more intense- coast of Maine, ]y around exposed borders of scales, Chin and top of head evenly speckled. A dark medio- Osmerus Norris,1868, p. 93, lateral bar without sharply delimited border. Sohuylki],l River, New Jersey (described but not In life sides an iridescent silver, the back an named by Norris,1868, p, 58). olive green. Total length to324mm according to Berg(1948), A larger subspecies than O.e. (Osmorus soronti auctoruni). nus.

Osmerus speotrum Cope 1870, p.490, Oernerus eerlanus eporlamus(Linnaeus) Wilton Pond, Kennobec County, Maine, (Fig. 1). Osmerus dontex Steindachner,1870, p.429, Salmo eperlanus Linnaeus,1758, p. 310. Dekastris Bay, USSR. European seas and rivers, Osmerus dvinonsis Smiti,1882, p, 32, Eporlanus Schonfoldii Rutty,1772, p. 358, Northern Dvina River, Russia. Ireland,

ho following natios named by Petrox, Salmo e erlano-marinus Bloch,1782, p. 229, 1925 p, 82, 108,have no status under the rules Table28,Fig. 1, northern and Baltic, seas and of the Int,Comm,Zool,Nomencl Osmerus e, dentex into rivers. roao kaninensis, Choshskaya Gulf; O,e, dento.x nabo nisseensis, Enisey River at Tyurin and Salmo eperlanus var, marinus Walbaum, some other localitie7. 1792, p. 57. Diagnosis Salmo spirinchus Pallas,1814, p, 387, lakes and rivers. Germany, European Russia, Distinguished from other osmerid species (Boloczero, Chudsk, and other localities). by the two large canines, one on either sido of the vomer opercular striae; the maxillary ex- ¿?Jl'lanua Rcndeletii Willughby,1789, tending past the pupil. 4 to8pyloric oaeca, p. 202,Table6, Fig. 4,Anadromous in Thames Phis subspecies is distinguished from O.e, oper- (unavailable since it is a reprint of a pre- lanus by the greater number of pored lateral Linnaean edition originally published in1686v line scales, usually 14 to28, Salmo eperlanue marinus: Walbaum,1792, Description p. 57(ex. Bloch), D8.-io(ii);C 19;A (12)13-.15(16);V 8; porlanus fluviatilis Gesner, in the syn- P 11-13(14); LL(13,14)1528(29,3o); midlat-. onymy for Salmo eporlanus of Donovan,1804, eral scales(62)63-69(72);vertebrae(58,59) p. 189(not available, a pre-Linnaean name cited 60-66(67,68,70); gill--rakers8-11 + 18-24 in a mynonymy7, (26) 27-..36(37); branohiostegals(6)7-8;pylorio caoca 4--8, Standard length3.8 to 4,4 Eperlanus vulgaris Gaimard,1851, p. 207, times head length,4,7to7.4timas depth. p, 18, Fig. 2,Iceland (this species not known Pectorals reaching from one-half to two-thirds in Iceland before or since this time according the distance to pelvic insertion; pelvic, a to Saemundsson,1949). little more than half way to anal origin, Adipose base short, about two-thirds of eye Osmerus e. eperlanus natio ladogensis diameter. Berg,1932, p, 281,Lake Ladoga (infrasubspeoif- io nanee have no status under the Int.Rulee Zool,Nomeno].7 ODiagnosis, description of subspecies end synonymy by McAllister(1963:15-21), FRm/578 Smelt 1:3

Diagnosis conci-) on Sat ho only difference between dentcznn mo'cisc in hc vertebral number Thin subspecies is distinguished from 0.e. YÍ ho"ovor io may be because of mordax by the lower number of pored lateral line lower -n-n' ornra turc during the development scales,4 to 13. of dcnty ogn T1o&lUs;or therefore con- siderod TonL synonyr of morciax. Mordax Description: arid nper nu di. ífer coaderab1y (especially number c' pornrli ntercl line scales - Table VI), D7-9; C 19; A (11)12-13(14); V 8; Using }i) cbnr-' c-ter oloris over 90 percent of P 11-12(13);LL (0)4-13(16); midiateral scales all bs cpnciox'n or ho-i-b forms can be separatath (58)61-69;vertebrae(55,56)57-61(62,63); Sinco bn i-oforma oro olbopatric and sinos gill-rakers 25-37; branchiostegls 7(8),py- vertebral and snal ray differences all ovorlap bric caaca3-7. Standard length about4to5 considerably it is necessary to consider the times head length,5.6to 6,1 times depth; two forms as aubapecifically different, The pectorals extend one-half to two-thirds the difference is nonolinal (McAllister,1963). distance to the pelvic insertion; pelvics about half-way to anal origin. Adipose base short, Data on plastic characters of different about two-thirds the orbit diameter. smelt populations ofurope and Asia are given in Tables III-V, oil rieristic characters in Medium sised canines on dentary, enlarging Table VI, posteriorly. Small pointed teeth on premaxil- lary and maxillary, Ductus pneumaticus attach- Plactic characters of the smelt vary con- ed to anterior end of the air bladder, Lateral siderab]y with ago (especially height of fins line incomplete, ceasing less than a head's and their situations, ddth of forehead, eye length along the body. diane ter ) , J( rp cbui)or (935 )found that height cT fino decronoen lTiih grouih corrola- Peritoneum light with dark speckles. Back tion coe:í'iccnj hrnn,, boight of A and greenish, sìdes silver in life. Total length standard Jongth (L) thc TThta Sea smelt were to307mm according to .rg (1948). A smaller -0,40 anc - 045 (for venous samples), between form than O,e. mordax, height of D and L ucrn -'0o3ß,-'0.42,-0.73; botucu P and L oro-0/5 cud -0.47,botwoon V 1,24 Standard common names and and L S-0,54. iIyO diO!1Otfl'PicOchews a nega-. vernacular names See Table I. 1ve oorrolmtoa 0o79sud -'075. Snout longih sd foreman udih theroaso uith age: TABLE I tha corrototini oocff: cionn unro i0,66and Standard common names and vernacular +00 75. VoIries o f--V P-iY, D- also increase, names of Onmerue erlanus(Le) although tho corvo) eion000ff'icint is rather low (ua:oJJy lesF íhr-;J.50)0

The smelt scale is very specific (Figs. 2, Country Standard common Vernacular 3), t'The smaller yr-.-n.old smelt has scales name name ui-th rolo tJ.rciyL-Lrgo focus and complete circuli.i ri hoOj (;oo nth Çi,o i rhoroas rica oC ihn.1 orger fT n)boro n rina31 or focun rind. one England smelt to several completed ci rouit0 floih typos how- fr an ce eperlan erar exhibit clearly the diriuon io features of Netherlandsemaelt a grañuoi rhon ii of tSr horncahoe-shaped in Finland kuore i noomplotrci ncn ï on Grouth ni ors ïn lato sea- Sweden nors son and a distinct cutting over the first cir- Norway mors culun of the second growing season. Later an- Canada smelt, éperlan nubi are easily recognizable. A growth field USA elt rainbow smelt contains n of complote circuli that USSR kor iushka zubatka, ogu- gradually5QHl.s growth slows. The first reohnik, koreha circulus that mpears in the following growing snetok, tint, season cuts a so these shortened oirouli° norms, ealaka, (Baybey, 1964) The year anriuli form as scars Japan kyuri-uwo outside tli' o of incomplete circuli of the last growilg'Hon (Figs.2, 3), Annulus formation otao-i near Juno. Circulus number 1.3Mphobog' in 'the successive years of growth in different smelt populations is shown in Table VII. 1,31 External morphology McAllister(1963),who studied the geo- graphical variability of smelt, came to the 14 /s78Smelt

Fig0 i Osmerue sperlanus eperlanue (L) (from Berg,1948)

Fige 2 Scale cf 2 yearold Fige3 Scale of5yearold smelt. White Sea. smelt. White Sea. MEgnif. 6 x 3, Magnif. 6 x 3. FRm/878 Smelt i s,

TABLE II Comparison o±' different forms of Osmerum (McAllister, 1963)

Characters Dentex Mordai iperlanu1

LO L0 15-20 1419 6-13 (min -mc, ) (13-3o) (13-19) (o6)

vertebrae 63-66 60-63 58-61 (snir0omaco) (597o) (58-65) (55-63) Head L/anal Iie1ht 26-32 23-4,4 23-26

Modal anal rays 13-14 13-.i5 12-13

Dorsal rays 8-10 9-10 79

Pectoral rays 12-13 (1112i3 11i2(13)

idiateral scales 63-72 63-65 61-69

Gill rakers 26-35 28-35 25-37

Pylorio casca 4-7 3o7

TABLE III

Data on fins ( 5 o body ] ) (after Kirpiohnikov, 1935;Tircìckf,1956,and others

Tbight Eo Ln3th Lnth Locality of D of P cf V of D baco of JI baso

White Sea 127-67 7.0-9.8 2o412o 9 82-ß6 119-1209 Chesha B 149-167 8,4 130 8092 123-132 Pechora River 161-170 10,8. 78-03 1303 Pekov-Leke 16.3 9,8 805 124 White L&o 16,7 bol 132 Finnish Gulf 15,0 9,5 29 0nega-L 169 10.3 138 82 132 Lados Y ':e 124-14,5 9,3-10,3 131 707 14 3 Neya River 157 9,9 130 G 8 7 123 Far Et 12,2-14,9 69-80 123139 109-12.9 Ycmimoy River 128-13,7 73.44 3GO-1 3 2 86 14.3 i33 124

tThJO oro ovoraorj o.d pies, not iin0niiz, vr1 uon of chorutoo o head (after Kirpichnikov, 1935; Data on nout length(as of hLo SeaLecaliy 21, 2-2 38 standard length *Theee are averages of dif Looality Eye diameter 5,7-7,0 length)(as % ofhead Head length Nikoisky, Forehead1956, and others) Lorer jaw Upper ja etciard(z % of li % length)widthof head (as standardlength)(as % of (as % of length)standard PechoraUhitePaitovChosha Lake RiveLakeBay 19,9-21,321. 7-224 20,921,0 409-5066.0-6.9 409407 16,6-20,9 21.921,219,021,1 20,4-24,8 24.920.4 - 10,9-11,013,1-13,812,5-14,3 11,811.4 7,8-8,28,9-9,37,6-9,6 8,3 NeyaLaciogaOnegaFinnish River Laits LakeGulf 28-22,5len 22,322082205 702700 20.6-24,8 19,622,327.1 20,5-21,4 23,724.525.319,1 13.011.3 - 11,2 8,59,2 - FarYe,iseyNo r East Rire 212136-21,7 4-2 1,5 223 erent samples, not min.-max. values of characters.7.1-7,5 6,9 19,9-20,619,0-20,6 19,9 22,8-25,322.8-25,3 25,1 8,69,8 12,813.014.8 (a±'er Kirpiohnilzov,Piasio oharace 1935; of suielt (as % of standard length) TABLE V Nikoisky, 1956, and others) A-D A-V P-V V-A Preanallength D-d peduncle lengthCaudal Pskovsko-ChudPechoraChesha1hite BayRiver Sea 1 121-4611312,7-16.0 0-1632 14,6 5.1-5.64.8-5148.5.4 45,8-47,8 5,0 4),45,7-46,6 3-46. 9 445-46.946,4-48,545.3-49,7 23,24,5-27,625.1-27.3 5-27, 4 24,0-25,522.1-23321.2-23,4 70.7 18,8-21,719.6-20.319.8-21.8 115-13.9 12.311,0 LadogaOrtegaFinntGhUhite LaiteLakeLaize Gulf 15.6-16.3 130315,6145 5,05,45.5 49,947.647,447,2 49,147.946,746.1 27,9-28,1 26,4250927.42401 24.222.222422.923,7 18,519.918.618,0 11.712,210,611,6 UorueyYeFar nice ]ast isvay River 1414.8-16.5 3-14.4 15,9 4,7-524.7-5,0 45.8-46.24903-4908 5.15.2503 47. 2-47. 8 48.848,6 46.9-47.7 49.3 25.9-28.126.1-27.1 27.226,6 22.22, 7-23.01-22.2 23.022.6 68.5-70,3 70,271.1704 19.719,9 - 12.1-12.711.6-11.911,3-12.5 11.2123 FRm/578 Smelt 17

TABLE VI retio obraotare of c'ron' ro! populations (after IirpichnU:ov91935 iko:Iay9 56 Lillelund, 1961)

tebrae Pored RzyaiíD Rays in A scale,Uobrenoheci Branohod Umb nched Branched in LL White Soa 29-38 60-67 15-27 1-3 5-10 2-4 12-16 Chesha Bay 26-34 62-67 16-24 Pechora River 31-35 58-63 7-8 2-3 7-0 3-4 12:14 Pakovso-Chudskoye I 32-35 58-60 - 2 8-9 3 12-14 White Lake 32-37 57-60 - 2-3 7-9 3 11-13 Finnish Gulf 32-36 59-62 8-10 2-3 8-9 3-4 11-13 0nea Lake 33-35 - - 2-3 - Ladoga Lake 4-11 2-3 10-11 3 12:14 Nava River - 4-11 2-3 8-9 - Far East 66 18-28 2 9 3 12:14 Yenisey River 66-67 14-26 2-3 9 3 13 1oray 59-62 - - Balic Sea 59-62 10-13 Chukotka - i420 West Germany 55-60 4-H

TAiLE VII

Ciroulus number in the suo sive years of gro h in different amai populations

Locality Yaarof grorth Authority i 2 3 4

Rivers of East 4 18 9-10 flacierman 1913 England

Miramiohi River 10 15 70 annie1958 (min.-max0 ) Ol9 102O4-o12

White Sea 2-3 9 9 5 4-5 Original (mino -max0 ) 0-6 712 8-134-7

i32 (;tor:ology(Chromosome may oo di lly in the short arm of one of co) tbos3 3moc pirs The rod shaped ohromoso»s y in length, Thus the longest Svbrdson (1945) found that "the smelt h rod-shaped pair at any rate in certain early short and slender chromosomes9 considerably motaphaoce may bo four tines ao long as the smaller than in the other species0 Their nhoreot0 Onenfthe rocI-ohaped pairs has number is the iouest amow tho Siodimh rep- proximal conetrotion0 The naioois taa not ntatiea of the Salmonida9 fanui3y, their studied"0 cliploid number being merely5t As arule, ho7e,rer, bo th Vhaped and rod-ehaped chromo- Otieiannil:ov(1885), Cunnincham (1886 - conos occur0 There aro five paire V-ehaped oiod by Tillelunci, 961) and Lillolund (1961) chromosomes0 Two of them c. app:;:- have denc:'ihod rpo000ytsoo' the amcl, o.tely oç'unl lantiì ha'eai ot]r threc p:i their nonbenos ond mioropyie0 Lons(1904) hao one markcdly long c'n0 Constriotioai nudtod vitolîogonae!e of smelt0 ICuenoteov l8 PRm 578 Smelt

(1964) gave a conparxtiv.ncJyois of oogonesis (Fig. 4);these vacuoles do not contain oil, of the migrant sselor tLic Ne-a river and non- During the phase of initial acounulation of migrant smalL-ci;od onolt (enotok) of the Pekov vitellin and oil the oocyte diameter is0- lake. He found $ha'i.the oocytes of the Nova 160»., the nuoleun is oval but in section its river smelt are largarho.n those of the Pekov margin ifestoonsd, At the periphery of the lake at the sarce phauos cf oogenoeia. Oogene- nucleus there are 6-12 nucleoli. The cyto- eis has aleo been studied in the White Sea plasm is vacuo].ated (Fig.5);the peripheral smelt. Acoordiig to Moyen (1939, 1944) the vacuoles contain oil, the oircumnuolear ones period of protoplasmatic growth of oocytes in- oontain vitellin, The oocyte membrane is a orn- aludes the juvenile phase and the one-layer layer zona radiata. Durinr the phase of in- follicle phase, During the juvenile phase the tensive tra hoplasmatic_2utb (Fig. 6the oocyte size in the Uhis Sca smelt is about - oocyte diameter measures from 320 to550,u. l3.5-36p, the nuclear diameter is ll3-27p. The oocyte is round or slightly ovol, The The cellular membraao 5e tb5n and. hae no visible long nucleus is in the middle, i.ts margia io structures. The nytopicom Jo griular There heavily featooned. The nu«i ' diameter is are 7-10 nucleoli ki thoivolsue (in a section). about 100,&. Thera are fom to 12nuolsoli During the ono-lryer fol] lolo pheo the oocyte at the periphery st the nucleus(in asection). size io Iron 45 to il2/ ihe nuclear diameter The cytoplasm is liosv5ly vacuolated and the is from 36 to 7605,u. f'.t the periphery of the vacuoles contain more oil (Fig0 7). Granules nucleus there are 20-40 nucleoli. The oocyte of vitellin, previously in the vacuoles, now is rounded, the cellular membrane is thin. appear in the cytoplasm. The oocyte membrane There is a more intensely coloured oiroumnuolearis thick and consists of two layers;zona zone at sorno distance from the nucleus whioli radiata externa and zona radiata interna.. The disappears after vacuolization has started first is more oloarly striated (Figo 6).Both (Fig.4) membranes'orm a Uttlo bo3.1.o7 at -thu Zorma-tion thu mioropy]o. Th&o thcphc'oo st' ripe- The pori OU (ir trOphop]CF1T1rl(r groït.h of floun the oocyte 0.rthu UI) to Soc' orno t roachoc smelt 000ytoo may bu divided into4phucoc. its definitive oio (chaut 800-. 900,u). The During the phuse of marginal voouolizatior the nuoleer cliamotou iabout 120/u0 The muoiono oocyte diemotor io uboot l44/bthe nuolear .a oval, its marginc feo tooed in section, diameter is abou8i1u !suries cf little J.t the boginniriof thu phase there ere vacuoloc appear at the periphery o: the oocyte nucleoli et -the nuclearoripbery but then

fl41 'Ii ' i'

.4'

Fig., 4Smelt oocytes during the juvenile Fig. 5Smelt oocytes during the phase phase (right) and the end of the of initial accumulation of phase of one-layer follicle, vitollin and oil. M*nif. 10 c 1XL Stuiniron Manif. 10 x 8.Stain; Mallory haomatox. (U',.ioAtheth) FRm/S78Sine it 1 9

they disappear. Vitollin granules and oil (Fig. 8). Vitellin and oil form a homogeneous globules of various sizes fill in the oocyte. mass at thin vegetative pole. After ovulation A very thin layer of fine-grained cytoplasm ad- eggs come into the body cavity and are t}en joins the oocyte membrane from within. The shed. Generally growth and developmont of oocyte membrane consists of two layers of zona oocytes in the White Sea smelt last 3-4 yinars, radiata; the thickness of zona radiata externa the period cf protoplasmatie growth being the is about 1103,t19 the thickness of zona radiata longest. interna is about 13.5/n. The micropyle is definitively formed(Fig0 8). The animal and. 1.33 Portein spooifioity the vegetative poles of the egg aro defined.. The nucleus moves from the middle of the egg Studies on this question are not yet pub- towards the mioropyle. The layer of f me- lished, but Rupp and. collaborators are working grained cytoplasm is thickest at the mioropyle on it (Rupp and Reclmorvi, 1966).

: r

Fig. 6 Oocytes in the phase of Fig,7 Oocytes in the phase of intensive trophoplasmatic intensive trophoplasmatic growth. Magnif. 10 x 40. growth. Magnif. 10 x 8. Stain: Mallory Stain: Sudan-Ill

...... Fig. 8Part of an oocyte in the phase of ripeness. Magnif, 10 x 20, Stain: Mallory

FRm/S78 Smelt 21

2. DISTRIBUTION 1920), with high oxygen saturation, high pII (1.8-8,6 and more), high degree of mineraliza- 2,1 Total area tion and Jew oxidability (Petrov, 1940) and rich development of zooplankton (about 300- Anadromous and landlocked fishes in the 1300 mg/rn3), Kojevnikov (1955) found that the Pacific, Arctic and Atlantic oceans and their Finnish Gulf smelt migrates and spawns in rivers baine (Fig.9), According to McAllister and estuaries at water temperatures from +2°C (1963), "the simplest explanation of the un- to 12-13°C, pH from 7,0 to 7.5, oxidability usual distribution of the two subspecies from 8,94 to 15.13 mg/i, concentrations of bi- (particularly the Arctic hiatus of O.eperlanua carbonates from 19.23 to 27,28 mg/i, oxygen mordax) involves an origin of the species in about 100 percent saturated, In summer the the North Pacific or adjacent Arotio, Spread smelt stays in open waters of the Finnish Gulf, then took place westward along Arctic Russia below the epilimnion layer at temperatures to the White Sea. From the White Sea they about 5-7°C. In autumn the smelt stays at were either "sluiced-up" in proglacial lakes depth of at least 20 in,at water temperatures or dammed in front of advancing pleistocene of 5-8°C, pH of 7,0-7,4, salinity of 340/00. glaciers aoci carried into the Baltio, Or they In winter the smelt stays at depths of 20-30 ni, may have migrated around Scandinavia to the temperatures of + 20 and salinity of 5-60/00. Baltic while isotherme wore depressed southward Generally the Finnish Gulf smelt spawns at during glaciation. In either ease, following temperatures from 5-6°C to 10-12°C, feeds at deglaoiation, the populations were left strand-. 5-10°C, spends the winter at +2°C. The White ed in lakes and cooler portions on the Baltic Sea smelt lives under more severo climatic con- to slowly undergo adaptation to higher tempera- ditions, The Sea covers with ice during 6-7 turo and lowered salinities, the intervening months of the year, the water temperature drops populations along the outer Norvegian coast below zero. As the ice breaks the sea begins disappearing. During the resultant isolation to warm and in summer the temperature of the the present !erlanus type evolved. The surface layer rises to 16-18°C (and higher), origin of eastern North American mordax from The average salinity of the surface layers of western North American populations of the sub-. the White Sea is 25-26 o/Oo, of deeper layers species can be ascribed to a migration across 30-31 o/oo; it ini lower near the river mouths Arctio Canada during a more recent interglacial (riega, North Dvina and others). Subarctic period or during the warm postglacial hypsi- populations of smelt inhabit waters with aver e thermal period. Cooling them resulted in the zooplankton biomass of 50-400 mg/rn3 and avore present separation of populations. If the biomass of bentho of 4-30 g/m2 (excluding migration took place during an interglacial Mollusca and Echinodermata which the smelt does period, one may assume that intermixing took not feed on), placo during the hypsithermal thus preventing differentiation of east and west populations. 2.2 Differential distribution Thus the two subspecies, one with disjunct distribution, can be readily derived, Range of 2,21 Spawn, larvae and juveniles morciax South to Barkley Sound, Vancouver Island, British Columbia, in the east Pacific; mouth to The smelt spawns in rivers or near their Wonsan, Korea, in the West Pacific; in the Arc-- mouths, Eggs are deniersal, adhering to the tic east to Cape Bathux'st, Northwest Territories,substratum. When hatched, smelt larvae drift west to the White Sea, The northernmost point downstream where they aro carried back and forth is on the southern island of Novaya Zemlya. In under th influence cf the tide, Fingerlings the Western Atlantic it is known from Pike Run stay offshore, feeding in pelagio waters in the Covo, Lake Melville, Labrador south to Delaware same regions as adults,, River, Pensylvania, andueetionably to Virginia. Range of eperlanus: the Baltic and. North Seas 2,22 Adults and their drainages, southward, inoluding England to the mouth of the Loire River, France; The smelt stays near the shores for moot upper Volga system" (McAllister, 1963), During of the year. It oom.s to the deeper waters of the last 10-15 years small smelt have spread seas (or lakes) only in mummer when shore waters down the Volga River system to the Gorkov and become too warm Smolt of all es spend the Kuybishev waterbodios, (Kuznotzov, 1951; winter in estuaries and mature fishes migrate Kojevnikov,1958). upstream in spring.

Over its large range the smelt lives under 2.3 Determinants of distribution oha. es various ecological conditionst salinity varies from oceanic to freshwater, temperatures vary o 2,1,3.16, 3.32 from below 0°C (in winter) to + 20°C and higher (in summer). Land-locked forms of the smelt inhabit pure lakes, some over 30niin depth, others shallow, with running water (Arnold, 2*2 FRm S3Smelt

24 Hybridization fertilization and breeding of eggs from fast growing coastal smelt and slow growing smelt 2.41 Hybrida; frequency of hybridi- from fresh waters (inland, lakes) have shown zation; species with which hy- that it is possible to hybridize these two mor- bridization occurs; methods of phological races, which have been isolated since hybridization the Yoldia geological period. Evidently the data relate to migrant and landlocked smelt of Interspecific hybridization is mot known. aerma). Lillelund(1961)writes that experimonts on

PPROxIUA1IOcALE 2000 3000 MILES o loon 200 3O0 41oo KIL0M00005

Fig.9Distribution of Osmorus eperlanus mordax-oirole, and O, e. eperlanustrianglea. FRm S78 Smelt 3:1

3 BIONOMICS AND LIFE HISTORY Intensive growth during the first year of life is oorrelated with earlier maturity (lake 31 Reproduction populations of Europe and North America; populations which grow more slowly reach maturity 3.11 Sexuality later (Table VIII)0 Fast growing year-classes reach maturity earlier than those growing more The smelt is normally heterosexual. slowly in the same population (Morosova, 1960; Lillelund, 1961; original data on White Sea According to Hoffmeister (1939) about 3.7 smelt). peroent of adult individuals of the Elbe river smelt are hermaphrodites (these individuals Size and weight at sexual maturity have male and female elements in thesame go- See Table IX. nads). According to Lillelund(1961),the figure forthe same river is about 2 percent. 3.13 Mating These individuals do not lose the ability to reproduce. Hermaphrodite individuals occur Each female usually spawns in the company more often among young smelt: Lillelund (1961) of one male. Thefemale extrudes her eggs, reports that 14 percent of the O-group smelt of then leaves the spawning ground. According to the Elbe river are hermaphrodites. Probably Hoover (1936) and Lillolund. (1961) an individus]. some of the hermaphrodites become males in the female may continue to spawn for severaldays, future, Hormaphroditism occurs more often in but the process Is completed in several hours migrant smelt populations than in landlocked in White Sea smelt (original data). Males ones of West Europe (Hoffmeieter, 1939). Herma- continue spawning with other females. Males phrodites occur very rarely in the smelt popula- spawn for a longer time and stay at the spawn- tions of eastern Europe. ing grounds longer than females.

Sexual dimorphism is described by McA11I 3.14 Fertilization (1963) who states, "paired and anal fins slight- iy larger in males, large tuberoles on scales; Fertilization is external. small ones on head and leading rays of dorsal, anal, caudal and paired fins in males. Tuber- 3,15Gonads oies reduced in females, There is a tendency in the males for a lateral muscular ridge to According to Kendall (1927)"... the go- form, to be darker and to have a punctate oper- nade of the smelt are unsymmetrical organs, one oulum", on each side of the abdominal cavity. The organ of the left sido is very much larger than 3q12Maturity that of the right aM. they are situated one behind the other. The leftgonad is much larger The age at, which sexual maturity im rea..d in both sexes, the right being quite small and differs widely in different populations (Table not far behind the outlet". VIII), Most cmelt populations reach maturity at 2-3 years. Theageof maturity gradually The correlation coefficiente for different decreases on advancing from the northern part years between egg number and body weight in the of East Europe eastwards to East Siberian shoram. White Sea smelt were +0.89 and. +0,95, betwoen Some workers noto that males reach maturity a egg number and body length 0,85 and *0.91 year earlier than females, but others do not (Beiyanina, 1966a). The egg number produoed note such a difference. by a female inoreasom with age; as a rule older fishes produce more eggs than younger f ishes Both sea and freshwaer smelt populations of the same mizo (Pable X). Data on fecundity are often biologically heterogeneous; there of various smelt populations according to siso are ecological forms differing in size and rato and ago are given in Table XI and Table XII, of growth, age and size at maturity, period and Land-locked freshwater forms and Siberian popu- placo of spawning and other peculiarities. lat ions have the lowest fecundity;nigrant This holds true for the smelt populations of forms (White Sea, ltio, Elbe river) have the the Baltic Sea baya (Marre, 1931* Kojevnikov, highest fecundity. 1955), lakes of East Europe (Arkhiptzova, 1956; Stefanovskaya, 1957 and many others), and North Variability of relative weight of ripe American populations (Kendall, 1927 and others). ovaries (to body voiglit) in different smelt The smaller and earlier maturing form is always populations is rather smalis fron 18 to 22 per- non-migratory, the larger and later maturing cent (Table XIII). Weight of ripe teti is one usually migrates for more or leas oonsxder-. about5percent. able distances from feeding to spawning grounds. Existence of those ecological forms promotes According to Lillelund (1961) for the the more complete use of food supply andspawn- Elbe river smelts ing grounds. y 2.737x - 0.28637 where "y" fecundity,'z" body length VL5CLZQ iLkc LUt ateb Pyc.arQOnz ct-cli cza irIicn Enf Lc rc.d3 tLb llavc Rivar 4o 5c97,2I O TichîgaLBaain iratch. RirezCzga. 3zy Seas 1. t 9'î 12,9 Lene. Rivnr akh & 2 8.2 10.7

8,5 11,0 87 9,4 to,' ta,93 1,B 12,7 15.6 20,4tO. 6 21,2I 36r'3 414.7 903 53.2 1609 10.1 54.5 19.2 1100 b'V Et flZ Ç!E VC 4.7 12.9 18,8 22.6 988,1 14.112.5 17,615.2 18,3 t 2.2 2355,8 8 2..4 2 22,017,6 6.3 tTe 6 4-63-41-3 2 2 2-3 2

2-4 3-4 6__J gr o up 8 3-4 4-5 5t3 51.8 123 I o5 li'o c'e ot' populatione turatioz feo tl1 4_5 12.9 t6 14,4 56 18,3 24,059.4

25,023.217,71901 2035903 27.4 21,5 27.0 20,5 20,2 223 24.6 22.2 20,923,9 27.4 19.6

22.2 244 28.1 23.4

OUD iuthoL'ity in spanin s10 l-31-2 ok DomraohFod.orova, 1953 and. 1.-31-3 Fed.orova,Pravdin,Ui].lor, 1926 19261953 1955-56 25.5 2-42.-51-2 Marre,Willor,Lapin, 1931 1931 1926 56,so. 5 6 90'4 2-52-64-122-8 Arkhiptzeva,Li].].elund,Ielyantev,tar,Ste f anov&zeja, 1961 19561946 19561957 2-32-52-43-9 IleKeazie,Kojevnikov,Creaser,Rupp,Balthiin, 1929 1948 1959 1958,19(4 10.7 4.-73-53-92-7 Kirpiohnikov,1Kirpichnikov,OriginalBalagurova, 935 1935 1957 ThtIiC7cz1f o fliVC' 3EI:E Bay Ob 4,63o5 26.3 27.6 28.8 30.1 7-125-94-8 Neiman,Pirojnikor,Tyurin, 1957 1924; tiB]av8ky 1950 1959 13.655.5T7. 3 S 8,0 9.5 15.6 Tot 13.4 17,2 7o8 Slot jJ 9.2 15.7 t 3.7 6.3 10,5 3 133130 T a a 12.2

w e 4 1Rm 578 Sm 3g 3

TABLE IX

Size ancwoigiit at maturity Çfirs spawig season) in lifforont smelt popu1tion

Locality Length Weight Authority (cmi

Rybinak waterbocly 6.0-9,0 1.9-6,2 Lapin, 1955-56 White Lake 6,0-8.5 1.5-3.5 Schotinina, 1954 limen Lake 4,7-8.2 1.1-4.3 Domrachev & Pravdin, 1926 Lazmiaden Lake 9.3-11.8 5.1-8.5 W:Iller, 1926 Kurishes Haff 6,3 1,4 Marre, 1931 Ladoga Lake 8,o-.11.o 3.3-8,7 Arkhiptzeva, 1956 Onega Lake 8,8-10,6 3.7-6,1 Stefanovskaya, 1957 and othe Eibe River 16,5-18,2 abt.17 Lillelund, 1961 White Sea 18,8-22,6 41.7-75.5 Original Ob River 18,3-19,3 42.5-48,5Ametislavaky, 1959 Yenimey Hiver 20.3-22.351.2-68.0 Tyurin, 1924 and others Lena River 19.6-23,4 53.0-94,0 Pirojni.kov, 1950 Upper Lake 14.0-16,0 17,0-25,5 Bayley, 1964

TABLE X

Fecundity (thousands of eggs) of females of thesame sizes but different ages (White Sea smelt, original data)

Length Age roupc (orn) 3 4 5 Average min.-max, Averadomin,-max, Aver e mth.-max.

19,0-19,9 37.6 25.6-44,5 42,0 37.1-50.0 20.0-20.9 53.0 45.5-61,0 53,2 40.7-66.3 21.0-21,9 52.8 - 62,1 43.9-83,8 a4.o-24. 9 88,6 87.7-90,0 95,5 88,8-102,3

Number of eggs per i g of body weight of fecundity and smaller eggs after the heavy one female is given in Table XIII. Siberian winter of 1960-61 than after the good winter of smelt populations have the lowest value of 1961-.-62 (Table xlv). this character, the Pekov lake smelt have the highest. According to LilleluncJ. (1961) the yield of different year-classes in the Elbe river shows The smelt spawns once a year, so the number a positive oorrelation with the water level and of eggs produced by a female during a year may is affeotect by temperature and food conditions be found in Table XII. Small smelt (Pakov- during the four weeks after hatching. Some lake, limen lake oto.) spawn once only (Lapins workers note that the yield tends to increase 1960; Kuznetsov, 1964). Migrant sea popula- in oolder years. tions have a longer life oyole; most White Sea smelt epawn once or twice during their lives 3.16 Spawning anO, morne spawn three times (Belyanina, 1966a), Siberian smelt apparently may spawn more often, The smelt spawns onoe a year am a rule but but there is sorno evidence that ouch fish do Kravohuk (1958) noted that sorne Siberi9n smelt not spawn every year (Kravohuk, 1958). The do not spawn every year. number of eggs produced by an individual during a lifetime may be calculated from Table XII. Sometimes smelt spawn under ice. In most Belyanina (1966b) reported that White Sea smelt areas spawning oontinues for about one month, contained less fat and had lighter gonads, lower but peak spawning usually lasts only ?-4 ds.yo; Peoundity of different smelt populationn (thoueawis of eggs) Data grouped by TA.BI XI h groups Pskocko-.Looality 10 11 12 13 14 1 Length groupe(on) 16 1 18 1' 20 21 22 23 26 2 28 Authority UchinokoycOnud.r31oyo 1. vaterbody White lake Rybinok iaorbody Kurishoo Iaff (-31 Pyoo sere

Onega Lake

Ladoga Lake

Kuriahea HaIT (sea-. migrant) Neya River

Finnleh Ou

Elbe River

White SeasKandalak- elia Bay 2.1 Gulf of ObOnega Bay

Te nia eyRiver 1,7

2,8 1.4

2.6 1,7 3,2 2,6

2,9 4,65.57,7 2,9

4,53,7 5,5 5.3 5,6

5.3

2,9 6,4 13.5 7,58,6 5.8 10.9 6.59.87.8

11.2 9,5 12,2 12,7 8.0

11,8 6.4 9.1

16.1 13.6 16.7 12,0

15,1 12.7 19,0

17.0 16,1 19,5 14,5

20,1 16,9 22.4

19.7 16,1 23,6 18,0

24,8 18.6

24.4 25.4 31.9 22.5

30.1 27,9 25.1

26,130.8

28.032,933.6 45,6 32,9 15.5 31.3

41.931.335,0 35.0 48,5 35,816,3 34.5

36.7 33,9

38.0

49,6 44.5 35.9 22,8

41.5

61.2 45.0 49.525.061.4

41.9 45,2 45.5 54.572,0

55.037.480,5 65.483.5 68.5 48,7

114 47.3 L.pin'o unpub.. Fedorova,SpannScheGrygorah,ihe. toay datcmina, 195319G3 Melyantsev,Marre, 19461954 1931 93.5 Arkhiptzeva,Markun,Alezandrova, 19561963 1926; 54.1 124 137 140 161 Kojevnikov,Marre,Lapin'a unpub.- 1949 1931 lished data OriginalLilieafter lund, Balagu- data 1361 Neiman,Tyurin,Amatimlavaky,raya, 1959 1957 19571924; (data grouped by age-groups and given in thousands of eggs Fecundity of different smelt populations TABLE XII Locality 2 3 4 5 Ago groups 6 7 8 9 10 ii 12 Lapin's data Authority Melyantzev,Fedorova,Schetinina, 1953 1946 1954

Alexandrova, 1963

1iare, 1931

Lapin'sLillelund, data 1961 WhitePskovskoChudskoye Lake L e Amatislaveky,OriginalSaiagurova, data 1959 1957

Rybinek waterbody Tyurin, 1924, Neimarx,1957

Pyaozero (Karelia)

Omega Lake

Kurishea Haf f

Finnish Gulf

Elbe River

White Seas Kandalakaha Bay Gulf of Ob Oriega Bay 206

Yenisey River 1.7

2,6

3.22,7 800 5,7

9.6 21,0 6,7

1306 4,6 - 19.9 2,9 10,6 1304

3705

35.6 22,9 24,733.1 2.93.7 12.5

69,639.5 41.9 505 15,9 25,0 126,1 60.1 5.9 20,9 8.6 6,3 133.6 25,5 3105 - 18,0 45.6 48.5 17400 3704 48.7 54.1 3f FRm/878 Smelt

TABLE XIII

Egg number per g of body weight, size of ripe eggs and relative weight of gonads (as % of bodyweight)th different. smelt populations

Eg number Average Relative Locality p'r g of diameter gonad Authority hor3 woihi of eg mm weiht

Pekoveko-Chudekoyc La'' 1050 0.55-0.75 18.4 Lapin's data Kuznotzov, 1964 White Lake 580 0.85-0,90 - Fedorova, 1953 Schetinina, 1954 Hybinek waterhody 710 0.10-0.90 20,7 Lapin's data Sohetinina, 1954 Pyaozero ((ars1ia) 560 0,79 13,5 Melyantzev, 1946

Ladoga lake 610 0.80 18.2 Markun, 1926 Arkhiptzeva, 1956 Kurishes Haff 665 0,82 - Marre, 1931

Finnish (iulf 630-700 0,75-0.95 18.1 Lapin's data Kojavnikov, 1949 Elbe River 650 0.70-0,90 18-19 Lillelund, 1961

White Sea: Kandalakeha Bay 700 0.86-0.94 22 Original data Onega By 710 0.77 22 Balagurova, 1957

Ob River 350 1.06 18-20 Amstislvsky, 1959

Yenisey River 370 0.96 20 Tyurin, 1924

Suyfun (Far East) 530 - 18.4 Dulkeit, 1937

TABLE XIV

Comparison of fecundity of females of the same weight in 1961 and 1962 (White Sea, original data)

Weight without internal or Number Characters (g) of 30 4 -60- - 10 fishes

1961 Fecundity (thousandsofeggs) 33.5 42.5 66.1 74,9 86.6 Egg number per g of body 884 862 947 927 910 weight 100 Average weight of gonads(g) 10.1 11.3 22,8 23.1 27.3 Average diameter of an egg 0.83 0.79 0,87 0,87 0.85 (mm)

Fatness (% of wet weight) 1,7-2,1

1962 Fecundity (thousands of eggs) 36.5 48,6 68.6 78.0 99.2 Egg number per g of body 893 913 1008 980 1016 weight 204 Average weight of gonacls(g) 11.2 15.7 21.9 25,1 33.4 Average diameter of an egg 0,91 0,92 0,92 0,92 0,93 (mm)

Fatness (% of wet weight) 2,9-3,0 FRm/S78Snielt 3:7

TABlE XV

Spawningseasons of smelt in different areas

Area Spawning season

West Europe March Lillelund, 1961 Baltio and drainage April - May Probatov, 1927 and other Upper Volga system April - May Fedorova, 1953 and others White Sea, Karelia May, early June Nelyantzev, 1946; Balagurova, 1957, and many others Siberia, Far East Juno, early July Agapov, 1941; Amstislavmky, 1963; Ivanova, 1955; Kuznetzova, 1962 Atlantic oats of From February MoKenie, 1964 North America (the south bound- ary of the area) to the beginning of June (the Mir- amichi River tn1 northwards)

according to McKenzie (1964) in the Mirarnichi begins earlier in warm years with high water river it lasts 5-10 days. There may be several level, The smelt enters rivers and spawns at spawning peaks, depending on weather conditions high tide. Sometimes the spawning run is de- and population heterogeneity (subpopulations). layed because of ioe movements,

As a rule smelt spawn at night (Domrachev Esox lucius, Leuciscus idus, Rutilus and Pravdin, 1926; Kendall, 1927, Hoover, 1936; rutilue,, Misgurnus fosailia, Acerina oernua Liovenso,1954,Rupp, 1959, McKenmie 1964, and spawnelmost at the amotime as smolt in original data). freshwatersof easternEurope (Meshkorand Sorokin, 1952). The majority of workers have noted a decrease in the size of smelt during the spawaing As a rulo smelt ascend rivers to spawn run: the older and larger individuals spawn (see 3.51), but some freshwater smelt popula- first, Changos in the sex-ratio on thspawn- tions spawn near river mouths and do not as- ing grounds aro discussed below. oend. therivers (Melyantzev, 1946; Fedorova, 1953; Sohetinina, 1954,etc.). Manyauthors According to Rupp (1959) the character of note that water depths at smelt spawning the spawning run depends on ice conditions: grounds varyfromseveral centimetres to soy- the spawning run of smelt in Maine, U.S.A. (in oral metres, According to Stofanowskaya 80 percent of casos) begins within the first 10 (1957) smelt eggs occur at the depths of 17 m days after the ice has broken. The spawning in some Karelian lakes.. The eggs are deposit- run breaks off when the weather is stormy or ed on stones, pbblos, water plants, submerged the moon is very bright. Colour of water does parts of bushes, grass and other things, They not influence the spawning run. do not occur on muddy bottoms, The current velocity in the spawning grounds of the White All over its great range(see Table xv) the Sea smelt is between 0.3 and 2 rn/sec (original smelt begins to spawn when the water temperatun data), is about400(sometimes 120 lower or higher). The spawning peak oocurs at water temperatures Spawning areas are larger in the years of 6.-9°C (Chumayevmkaya-Svetovidova, 1945; with high water level. Rupp and Redmond Marcotte, 1946; Marcotte and Tremblay, l943; (1966) noted that smelt taken from spawning Dryagin, 1949; Schetinina, 1954; Ivanova, grounds in rivers and introduced into some 1955; Stofanovekaja, 1957; McKenzie, 1964, lakes of Maine did not enter other rivers but etc.). Spawning breaks off at a sharp de- spawned nearthe banks ofthe lakes, crease in water temperature. It lasts longer in cold years than in 'warm ones, The spawn- A numberofauthors, including Kendall ing smelt usually avoids temperatures lower (1927), Baldwin(1948),Lillelund (1961), than 4°C and higher than 12°C. Spawning Amstislavsky(1963), and unpublished datare-

FRm/578 Smelt 3:9 over according to Unanyan and Soin (1963) only Egg predators includo insects and fishes dead egga of the White Sea emelt become de- (etickiehack, smelt itself and, others).Eggs tached from the aubetratum may become parasitized by Saprolegnia. The vitellum io round or a little oval in Nany authors (including Schneherger, 1937; chape. A thin layer of cytoplasm ourrouncio Neehknv and Sorokin, 1952; Unanyan and Soin, it. The vitelline membrane le thin, and flume- 1963) have remarked on the high rate of survival of smelt eggs during incubation. Accord- roue oil1obules are dieperood all over the ing to LiJlelund, (1961) the rate of mortality vitellus (see 1.32). inoreasos only at the end of the incubation 32 Pre-adult phae period, when about 86 percent of eggs of the Elbe river smelt die, However Rothschild 32IEmbryonic phase (1961) haoLated that lose of smelt eggs in the Dean-Brook river, Maine, U.S.A., is 99.5 The embryonic phase is clesorihed by Unanyan percent. The main causes are drying after t} and Soin (1963). After extrusion into water, water ìvel ham dropped, overwarmning and pre- the egg immediately swells. Its diameter in- dation (see above). Influence of these fac- creases about 0.2 mm, perivitelline apace forma tors is least in cold. years with high water between the vitellus and the membranes. In 4 h level. (temperature of 805°C) the blastodimo forme at the animai isole of the egg (Fig. 10.1). In 8 h Rothschild (1961) and McKenzie (1964) after fertilization 2 blaFitomeres are present found that the number of larvae produced per (Fig. 10.2) in 13 h 30 mm, 8 blastomeree are unit area morcases up to egg densities of present (Fig. 10.3), In 2,5 days after fert- about 11-13 per and declines when the den- ilization (12°c) the biastula has formed (Fig. sity of eggs is. higher. The relativo larval 10.6). Formation of the embryonic plates be- production is highest (hatching rate 3.6%) gins 2 days 14 h after fertilization (Fig. 10.7).whom the average density of eggs is 0.5 per cm2 In 3 days gastrul atiois complote (Fig. 10.8). and declines to 0,03% at an egg density of 194 In 4 days 5 h (10.5°c) the embryo has eyecups PST cm2. (Fig. 11.34);in days 5 h after fertilization (io°c5 segmentation of the mesodermn begins, Before hatching, the embryo begins to move the main parte of the brain form, auditory veo- very energetically, the caudal part moving es- toles and c:rystallins lonsea in the eyes are pecially sharply. The membrane bursts and the present. The embryo has a ohorda, 30-32 rnyo- tail comas out first, Moving energetically tornee, and the caudal part begins to separate the embryo frees itself, the egg-case romaine from the yolk sac (Fig. 11.5,6,7). In 7 days on the substratum or is torn off during hatch- 10 h the embryo ham a long tail separated from ing. the yolk sac, the eyes have developed black pigment;the embryo showm twitohing movemento 3.22Larval phase within the egg (Fig, 11.8). In 9 days and 16h after fertilization (9.5°C) hatching glands are The period of incubation varios in differ- present all over the body surface, particularly ent smelt populations (Table XVI). Generally on the head and i.eo±oral fins;the parts of small forme have a shorter period of incubation the brain are well formed;t}1e black pigment and the new:ly hatched larvae are smaller. in the eyes become stronger and yellow pigment Naturally the sum of day-degrees from fertili- appears. The yolk sac is egg-shaped. The zation until ha'tohing is not constant, but in- embryo han olfactory vesicles;oemiolocular oreases exponentially with decreasing water canales are preoent in thauditory vesiclem. temperature. The head is not distinct from the yolk sac. The mouth iat the lower side of the head. A newly batched pro-larva has the yolk sao Four rudimentary gill slits and, a rudimontary about 0.7 mm in length. There is an oil glob- operculurn are present. Melonophores are found ule in it. The globule resolves after the on the ventral aide of the caudal myotomos, yolk. Grib (1946) described 4 stages in the under the ai isentary canal, and on the yolk sac. larval development of the Nova river smelt: The heart pulsates regularly;it consists of two ohamhe:ra. The blood has no corpuscles. 1. Pro-larva with yolk-sac. This etage There are 42 pre-anal and. 17 poet-anal myotomes laste 7-9 days. The dorsal fin fold begins at (Fig. 12_i) the second segment. The pro-anal fin is well developed. Pectoral fins aro present. Pig- Thring embryonic development there are no mentation is on the ventral part only, The differences between migrant and small freshwater body is transparent. The blood has no cor-S smelt except the body size and number of myo- puedes. The mouth becomes terminal at the tornee (Orib, 1946). end of 'the stage.. The pro-larva begins feeding while the oil globule is still presont. 'l'ho 'o. loi"vmeasures to 7 mm in lngth (c"ig.12 Fie. 10Early stages of development of eggs of the White Sea smelt (to the end of gastrulation) (from Unanyari and Soin, 1963).

Fig0 11 Late stages of development of eggs of the White Sea smelt (to hatching) (from Unanyan and Soin, 1963) .FRJS78 Smelt

TABLE XVI Duration of period of incubation and body length at hatohing in different smelt populations

Locality Sum of Body length at Authority day-degrees hatching (mm)

Pskov-lake 80 3,8 Mohkov and Sorokiri, 1952 Valday-lake 110 6,2 Chumayovskaya-Sve tovi.dova,1945 Rybinmk-waterbody 138 4.2-4,9 Sohetinina, 1954 Elbe river 60-110 5.0-6.0 Lilleluni,1961 eva river 140-180 5.4-6,0 Grib,1946 Thite Sea aht.170 6,0 Unanyan and Soin,1963 Ob estuary 132 - Amatisiaveky,1959 irasichi river 174 5.0 Ì4oKenzie, 1964

Larva without yolk-mao, 6.8-13.4 mrs. Smelt oggo and larvae arc fed on by the Blood oorpusolem are present. The fin fold three-spine stickleback, gobiido, herring, is reducod, dorsal and anal rays are developing.perch and, other fishes (including the r"elt Pigmentation is similar to that of the pro- itself) and larvae of Inseota. One o ha larva;a pigment spot of two melanophores le main causes of egg mortality is SaJ: present under the anas, The alimentary canal is tubular. A rudimentary gas-bladder can be Unanyan and Soin(1963)sta;;:d .t lr.ae seen at the end, of the stage. This stage r" tho White Sea ornait begin food within lasts from 10-14 days after hatching (Fig.13), 6days of hat chirig and aro on -.r l.- 4cpmndeni on external feeding9 days pf c Larvae of 13-13 mm. The body is trame-. According to Gri(1946) the '''L1rcr parent. Thdorsal, anal and adipose fins begins to food7-9days after ) ch'c The have formed, Bays are present in the unpaired corresponding time for Pekov-laka "rich i" 7-8 fins, The caudal fin is rounded. Tho pro- days (Meshkov and Sorokin,1952), anal fold begins from the eighth pre-anal nag- ment. Lilelunrl(1961)stated that a cro-larva with yolk-sac may live without fc ri.::.r. 'or 10- Larvae of more than 18 mrs (28 mm in 14days at a temperature of 10-12 average), Such. larvae appear in the Nova river in July-August. The caudal fin is in- Larval smelt feed ort email pinetonic dented, The final number of raye are present forms:young stages of Copepcda hotatoria, in the dorsal arid anal fins. The embryonic larvae of Gastropoda and Lainoliibr:cichiata, fin fold can be distinguished on the rentra.]. aide in fish of up to 30 mm. The ventral fins 3.23Adolescent phase are situated more posteriorly than earlier as a result of growth of the gas bladäier. Upper According to Grib (1946) the adoJeecen and lower vertebral arches are present. The phase in the Neya river smelt bogr-j n operouluni covers the gills completely, Pig- September when the first scaler ,'rr'e (c4 cm), ment is concentrated mainly on tho ventral side, Ehrenbaum (1909) gave the eate dc for.hs E]be Melanophores appear on the operculum, on the river smelt. However McKemz'a (9') e'aect top of snout, under the ayes, along the chorda, that first scales in the Mircrr o r - - cr rmait on the body sides and on the dorsal part cf th form at a length of 20-25 mrs (g 2 'c) By tail to the end of the stage, Larvae and this time the pigmentation of1r do'sal par adults have similar nieristic and plastic cha- becomes moro intenso. racters. The number of myotomes does not change during the larval period. The main mortality occurs rhr rg the'irr h months of life, Lillolund, (t3) otmchrI, According to Raes (1949) in general the the mortality coefficient is ghoefrrrs smelt larvae belong to the "herring type" hav- May-August, its value decrcrsmr;r' flnoPec rci ing a relative short caudal part, isometric then it is of little iwporteucc pigmentation, low disposition of the pectoral fins, homocerca]. tail and so on. 3x12

1mm o

o.- ,,g e,

Fig. 12 1. Embryo of White Sea smelt9 eggmembranes removed

2,3, Prolarvae of White Sea smelt (from Unanyan and. Soin, 1963)

1mm

Fig013 Larvae of White Sea smelt after resorbtion of yolk (from Unanyan and Soin, 1963) PRm/S75 Smelt 3r13

The young smelt feeds on planktonic Orum- Cyprinidae (Rutilus rutilus, Alburnus alburnus, tacoans (copepoàa, Cladocera) and, other groupe Abranis brama, Leuciscus idus and others) and of zooplunkton. perch (Perca fluviatilis) (Kojevriikov, 1955). The young of the Elbe river smelt compete for 3,3 Adult phase (mature fish) food with the young of Clupea finta from May to August (Lillelund,1961). The fresh water 3.31 Longevity smelt of the north-western part of USSR occupies the food niche of a plankton-feeder in open Longevity varies greatly in various popula- waters and competes for zooplankton with Core- tians of Osmerus eperlanus (Table VIII). Gen- albula, Abramia ballerus, Alburnus albur- erally small freshwater smelt do not eurviv nus, young of Perca fluviatilis, sometimes with for more than 3 years. Individuals of 12 and, Rutilus rutilue and Acerina cernua (Petrov, oven 15 years old may be found in the smelt 1940; Meehicov and Sorokin, 1952, and others). populations of estuaries of the Yenisey, Lena, Anadyr and Khatanga rivers (Agapov, 1941; 3,34 Predators Mikhin, 1941; Lukyanohikov, 1964). The max- 1mal age of malos is usually 2 years less than Sea mammals, birds and fishes (cod in that of females. particular) food on migrant spawning smelt in subarctic waters. Ezox lucius, Lucioperca 3.32Hardiness luo1operca and large individuals of Perca f lu- viatilis feed on smelt in European fresh and The environment of smelt populations is brackish waters, Salvelinus fontinalie, discussed, in section 2,1. Cristiwomer nanaycush, gnue clupeaformis, Stizorstodion vitreum, Perca flavescens, Anguilla According to LilloluncI (1961) experiments rostrata, Ì4icropterus dolomleu, Norone americana made in Hamburg harbour showed that eggs develop and. Lota ,yiaoulosa feed on smelt in North American in spite of sewage water. Only in the last wato'"rKendall, 1927). phase of development where there is very little current is there a great mortality of eggs, There is sorne evidence that the specific caused by the accumulation of detritus, proto- smell of ornelt stock scares away other fishes. zoa and algae on the eggs. Eggs of smelt are fertilo up to a salinity of16o/oo. The sua- 3.35 Parasites, diseases, injuries oeptihi.lity of eggs to salinity during the arid abnormalities period of incubation depends on the stage of embryological development. A salinity below According to Petrow(1940) 28.5-36.6%of 10 0/00 15 neither injurious to eggs nor to the Pokey-lake and 20% of the White laico smelt larv-ae, The upper limit of temperature toler- are parasitized. Parasitism io less intonsiv ance of early eggs lies in the range 17,7-20.7°C, in other lakes of West Europe. Shulman and for later stages it is 21-24°C. Comparable Shulman-Aihova (1953) list 16 species of para- data for adult smelt are not available. sites for the White Sea smelt,

3.33 Competitors The list of smelt parasites recorded from waters of the USSR is given in Table XVII The main food competitors of Siberian smelt are Coregonidae and Acerina cernua feed- 3,4 Nutrition and growth ing on Chironomidas larvae and nektobenthic Oruetacea (Pirojnikov, 1950, 1955; Amst1elavçy, 3.41 Peoding 1963, and others). Anstiolavoky and ussynina (1963) stated Young of the White Sea smelt compete for that the Ob river smelt feeds during day and zooplankton with herring, adults compete for might but with different intensity: maximum Amphipocla, Mysidacea and some Cumacea with feeding takes place at 13 and 21 h corresponding herring in April-May and. September-October to vertical migration of zooplankton; minimum (Epstein, 1957). There is great similarity feeding is at5h. in diet (especially on the basis of Polychaeta) between the White Sea smelt and Eleginus navaga, Migrant smelt feeds in coastal and offshore yoxocephalus M. uadricornis (Russa- waters of seas and in river estuaries; land- nova1 1963)Thasterosteus aculeatusTimakova, locked forms feed in open waters of lakes. 1957), Pleuronectos flesus and Liopsetta glacialim (Kudereky and Russanova, 1963). The smelt searches in the water taking food According to Abdel-Malek(1963)there is no organisms selectively (Table xviii), food competition between adults of smelt and three-opined stickleback. In the early summer According to Timakova (i 957) the White Sea the Finnish Gulf smelt competes for planktonic smelt feeds on fish (herring) and Crustaoea cruataceans with young and partly adult during the post-spawning period. Intensity of 3214 FRín/S78 Suieli

TABLE XVII Parasites of smelt (for territory of USSR)!!

Class or Stage of para- Order Species site infeotirig smelt

Miorosporidia Glugea herwigi* Adult usoles, gills, Baltic, White Weissenberg, me menteries and Bareniz 1921 etc. Seas and their basins, lakes of the Upper Volga, Amur. Cestoidoa TriaenophoruB Larva (plouro- Muscles and Proceroolds Area of Esox crassus Forel, cercoids) internal in body oavitylucius and 1880 organs of Copepoda, Esci reiohorti adult is a parasite of p ike Triaenophorus * Larva (enoys-Liver and Any Copepods, All over USSR nodulosus ted pleuro- other organs the final (Pallas, 1781) cerco ida) hosts aro pike, perch and other predators Eubothrium Adult Intestine PleurocercoidsWithin the ar crassum Bio oh, in intestine oarea of Sal- 1779 perch, pro car-moflida coids in any Cyclops .h liobothrjumLarva (pleu- Musolea, liverThe final host-North-We stern atumL.,175; rocercoids) and other mammals USSR, Volga organs Yenisey, Ob Rivers, basins DiphyllobothriumEncys ted Tissues of Final hosts Not recorded norveicum Vik, Pleuro cerco idsintestine and mammals and 1957 stomach birds Proteocephalus Adult Intet me North-Wes tern longicoll is USSR, Siberia ÇLeder, 1800) Kajnchatka, Ainur River

Trematoda Leicithaster sp.Adult Intestine Not recorded Orientophorus Adult Pylorio oaeoa Far East, trovi Kainchatka (Layman, 1930) Podocotyle Adult Intestine Basins of the reflexa White, Barentz, (Creplin, Japan Seas 1925)

JJTable based on "A key for identification of parasites offreshwater fishes of USSR", edited by B. Byhhcveky (1962) FRm/578 Sm1t 3x15

TABlE XVII (continued)

Class or St of parai- Infected Other hosts Order Spe oie s site infecting organs and stages Are a smelt in them

Trematoda Buoejhalue Adut Gill tissues, Baltic and White 01 mor hum under skin Seas basins, Baer, 1 27 Siberian and Far East Rivers Cotylurum rile- Cyets Tissues of Final hosts- Baltic and White atus (Run,, 1802) gas bladder birds Sea, Volga and other Siberian Rivers internai Rybinek waterbor, organs, body Kare lia cavity Totracotyle Larva He art Adult unkno Not recorded intermedia Hughes, 1928 Diplostomum**31 Larva Crystalline Final hosts- ot recorded spathaoeum lens of oyes birds (Rud., 1819)

Nematoda Contracaeown Adult St mach, ceso- North-Wee tern aciuncum phagus, pylo- USSR, Ob and Amur (Rud., 1802) rio oaeoa Rivers, Japan Liver, n. and Okhotsk Seas. ry, muscles basi mci Porro cae own Adult, ep eri ani larva Dorsal muscles North-Western (Linstow, 1019) under skin USSR Cystidicol Adult, Gas bladder Baltic, White, far io ni s larva Barentz, ring Fisher, 1798) Seas basins Camalianus Acilt, Intestine Evsryuhere, e- lacustris iorra copia the Amur (Zoega, 1776) River basin Camailanuçj Intestine 'ytihere, ex- truncatus i rva the Amur 1iidT7il 4) Philomotra Adult, Body cavity, recorded manguinea larva the skin bot- Rud,, 1917) ween fin yo 3:16 FRrn/78 Smelt

TABLE XVII (continued)

Class or Stage of para- Infected Other hosts Order Spe oies sito infecting organs and stages Are a smelt in then

AcantocephalaEchinorhynohue Adult Intestine Not shown Not recorded adi Mil1or, i 77

Pseudoo chi- Adult lutes tine Intermediate Everywhere chua ho s ts-Gamînarus clavula (Bu- pulox and. jardin, 1845) Fontoporeia af f luis

Mete ohinorhyn- Adult Intestine Intermediate chue salmenis host-Ponto- TiiTier, 1780) porela affinis

Ponrnhorhyuchuu Adult Intestino, laovie livor Gammars pule North-Western (MUller) USSR, Volga

Co ryno s orna. Larva Body cavity Final hosts North and East semermo sea marinais Seas of USSR (Forsoell,1904) rarely birds Coryn000m Larva Body o'iy, Final honte -North and East U t3.'IUUO CUIR loulicles, lu- esa luamnale Se of USSR (Ruth, ThO2) terial orUano rarely brclo

Copepoda iraoiJio. Adult GiLls 2a1tio,Siberia, ant Markouitsh, Amur River 932

Ergasilis sio*Adult Gills Baltic, Japan Sea boldi Nordmann, Siberia 1832

Adult Gills, skin ar East Gussov 1951

Banchiura Ax' lus :í:o *.du1t Gills and Europe, Siberia cous L.,1(56) skin

* May causo mass mortality of fish Causes loss of weight ** Couses blincthess and mortality FRm/S78 Smelt 3:17

TABLE XVIII

Selection of food organisms by the Lena river smelt (after Firojnikov,1955). Fishes were 6-11 years old and measured161-317mm

Components share (in %) Share in the in plankton in nekton diet of smelt

Copepoda 51.2 O Ciado cera 32,9 o Mys idao 24.2 67.3 Amphipoda 11,9 11,4 Isopoda 1 .7 o Cumacea 700 o

feeding increases until July. Polychaeta(45%) (Cyclops5go,Diaptomus sp. etc.). The maim and herring eggs(27%)aro the main food cornpo- food groups of adults of different populations nents. In summer older individuals feed on are Nysidacea, Amphipoda and young fish. The Polychaeta, Decapoda and fishes, in autumn they smelt of the Atlantic coasts of North America feed on Polychaeta and Amphipoda. Intensity of feeds on pelagic Isopoda (Kendall, 1927). The feeding decreases until December. lu winter it diet of the White Sea smelt consists of60com- is low, and food consists of nektobenthic Crus- ponents (Kudersky and Russanova,1963), tacea, Amstislavsky and Brussynina(1963) Polychaeta playing the main rolo (mainly Nereida noted that the Ob river smelt feeds on Amphipoda which do not occur in smelt diet in other areas). in winter. Thiring the vegetative season Mysi- dacea, Copepoda, Cladocera and young smelt are 3043 Growth rato important in smelt diet. The Lena river smelt has the same diet (Pirojnikov,1950,1955), The According to :lyanina(1968)all smelt Elbe river smelt foods on Eurytemora from April populations may be divided into 2 groups based to October and Gammaridae from November to March on character of the growth rate during ontogeiys (Ladiges, 1935). Planktonic Crustacea dominato (i) populations whose individuals grow most in the diet of small freshwater smelt. Their intensively during their first year of life; importance in the diet depends on seasonal then the growth rate decreases comparatively abundance. sharply (smelt populations of West and East European basins, oxoludin the extreme North, According to Tiniakova(1957)intensity of and North-American waters); feeding of ritales and females of the same age (2) populations whose individuals grow com- varies with the seamon after spawning females paratively slowly during their first year while feed much more intensively than males; in July their rate of growth increases more or less and August maies feed more intensively, starply during their second year. Then it Females oonsumo more fish than males. decreases gradually (smelt populations of the White Sea, the Cheshskaya Bay and Siberian The assortment of food components, their coasts). Sizes of individualrj of different sizes and quantity change with growth. Young smelt populations in successive years of life of all populations feed on zoopiankton. Fishes are given in Table VIII. Valuen of the linear aged. 2 years eat Nysidas, Amphipoda, Inseota growth (in % to maximal length) of some popula- larvae and fishes, The importance of fishes tions of both groups are shown in Fig. 14. increases with :e. Condition factor (Ponderal index) In winter the smelt feeds very little. As a rule smelt abstain from feeding during the The well-known length-weight relation in spawning and pro-spawning period (Nikolsky, fish may be expressed by the following equatn 1956,and others). Q Al', where Q weight, i . length, 3.42 Food A const., n - near3

Young and adults of small freshwater smelt Values of "n'e for different smelt populations feed on planktonio CruBtaoea: Cladocera are given in Table XIX. (Bosmina sp., Daphnia sp. etc.) and Copepoda 3:18 FRmJ78 Smelt

TABLE XIX

Values of ??I for different smelt populatione

Looality n Authority

Landlocked White lake 2.70 Sohetinina,1954 Rybinck waterbody 2.91 Schetinina,1954 Kuriahea Haff 2.87 Marro,1931 Great lakee 2,82 Lillelund,1961

Sea,.-migrant Elbe river 3.28 Lilleiund,1961 F!tramioh! river 3.20 McKenzie, 1958f Lillelund,1961 hito Sec Ki1uIciha Bay 3.48 Original data Drina Buy 3.36 Kirpiohnikov,1935 lonisoy rIver 3.15 Tyurin,1924and otberm

Ryhinek watorbody Finnish Gulf

(50 (50

IIQ L'o

20 20

1 2

A. Populations of the 1group

Yenisey River

r;Q

1 2 4 5 (5 1 2 4 5 6 7 8 yesro ei. life years of life

B. Populations of the 2group

Fig, 14 Linear growth of fish in successivo years of life (as % of maximal length) of different smelt populations (from Belyanlna,1968). FRs/S73 Sm I 3 19

ThBLTI XX

Rate of growth in different smelt populationa. (from data in Table VIII)

Locality Length increase in successive years cf life 2 3 4 5 6 7 8 9 10 11 12

Frochwaters

Pskov-1 alce '(,2 2.93.2 limen lake 5.23.02.5 Valday lake 5034.61.3 White lake 6.02052.5 Ryb1nk wtb 5.92,80,7 Kurishee Haf f 6.34.2 Darley lake 7.1 3.61,51.5 Lazniiaclon lake 8.21.12.5loi2,9 Lad.oga lake 8.0 1,51,22092.22.5 Onega lake 6.32.50,61,20040.30,50.5 Michigan 9.26.51.4 Upper lake 6,58.34.22.31.22.31.2

Sea-migrant Elbe river 7.16.33,93.92.6 Finnish Gulf 7.83,32.5204 1,6

White Seas Onega Bay 4075,44.44074,0 4,2 Dvina Bay 4.15.23093,7 2,2 2,4 3.12.80.7 Kanciaiaksha Bay 4,7 8,2 5,93.82.42,0 Chesha Bay 3.54.64.42,72.52081.7 Yenisey river 4,55,34,33.52.7200 1060,51.1 Lena river 3Q8 2.9 1.2 1.3 Amur river 505 6.0305

The populations of the first group live of growth increases during the second year of under temperate climatic conditions0 Their life and greatly exceeds that of the first year season of growth lasts6-7months. They live (Fig. 14). This is evidently a result of in waters with a good supply of small zoopiank- changes inooding habits; the smelt begins to ton whioh the fry and young feed. on te feed. on lrc C:custaoeans (Amphipoda, Mysiciacea). plankton biomasa is about300-1300mg/rn , as The Kandalakaha Day smelt (white Sea) is remark- may be concluded from various literature able for iba most rapid growth (linear and sources0 The rate of growth of freshwater weight) th'cin the socond (and. third.) year of landJocked populations decreases sharply during life (Tablao VUI,tX, xIx), This population ontogeny (Belyanina, 1968 Tablos VIII, IX, lives under conditions of higher salinity (25-. XIX, XX Fig014). PoDulations of the second 28 O/oo)than do ali other smelt pop'lations, group live under severo sub-arctic conditions. The food resources of the White Sea are richer Their season of groui,h lasts5-6months (White }1r'those of estuarios of Siberian rivera; the Sea smelt) or only iaonths (East Siberian average biomasa, of benthos of the White Sea is populations)0 The average zooplankton biornass about10-30 g/rn2(excluding Mollusoa and Eohino- is about50-300 mg/n3dulng the season0 The derynete, which smelt does not feed on), while latei' spawning period (May-June and even July) this valuo is only4-5 g/m2for the estuarios compared to that of the European and American of the Siberian rivers (Oreas, 1957, and others), populations (March-April), the lower tempera- Also the Kandalaksha Bay smelt feeds intensively ture during the season, the earlier cold In on heteronsreid. stages of Polychaeta, mainly autumn, the poorer food resources * all theme Nerelu virons (Abdel-.Malek), 1963; Kudorsky and factors c1.00rmine the slow growth-rate of the ilusoonova, 1963) which is an eaiiy accessible young smelt of these regions0 However, rate and very abundant food in this region. 32O FRni373 niei&

The length of the growth season is differ- 3.5 Behaviour ent for different age groups young f leh begin to grow earlier and finish the season later 3.51 Migrations and local movements than older individuals (Lillelund, 1961 and. others). This may be related to spawning in During most of the year smelt stay near spring or in the beginning of summer. shores. After spawning the fish migrate to deeper water, evidently to avoid high tempera- Smelt grow faster in warm years and slower tures, In autumn fish of all age-groupe keep in cold ones (Kirpiohnikov, 1935; Meshkov and together. Smelt tend to form local popula- Sorokin, 1952, original data on the White Sea tions. Feeding migrations of smelt are short. smelt). The effect of temperature on growth They spend the winter at river mouths. In is seen more clearly in younger fish. sprin, they migrate to spawning areas Upstream. Extent of spawning migrations is given in Generally strong year-classes grow more Table XXIV, season of migration in Table XV. slowly than weak ones (Lillelund, 1961; After spawning adult smelt actively move down- Morosova, 1960, and original data). According river, prolarvae drift passively. to Abrosov and Agapov (1957) the rate of growth of the small smelt of the Jijitzkoye lake de- Differences in the migratory behaviour of creases with increasing population density. different populations are mentioned in Section 3.12, 3.44Metabolism 3,52Schooling The metaboljm of smelt has not been studied. Evidently smelt form schools during all stages of the life cycle (Kendall, 1927 and Leviyova (1952), Kleimenov (1962) and morne others). By our observations only the largest others give data on the chemical composition of and oldest examples of White Sea smelt (6-8 yr smelt (Table XXI), Data on fat are shown in old) do not form schools but search for fish Table XXII, Siberian smelt have the most fat alone. Spawning and wintering concentrations in the body and reproductive products. The are rather dense, and feeding schools are of small smelt of the Upper Volga system have t'ne i:ms densìty. In autumn and winter, smelt lowest fat content. During the spawning period concentrations consist of fish of both sexes the fat content of the body and internal organs and different age groups. In spring, mature of males (except gonads) is higher than that of individuals form pawning schools, grouping by females. Chechenkin (1952) reported that the size or age. fat of the Pakov lake smelt ccnsiets mainly of non-saturated fatty acids, with oleic acid Gohiids, flatfishes and other species dominant. sometimes occur in fishing note together with the White Sea smelt; herring, stickleback, Fatness chances during life and by seasons sprat nd others with the Finnish Gulf smelt, (Belyanina, l966b). Localization of fat in smelt (in por cent to total fat) is shown in Larvae and young smelt feeding on plankton Table XXIII. perform vertical movements (Amstislavsky, 1963; McKenzie, 1964). Ripening of the female gonads results in considerable changes in the localization of fat The behaviour of spawning schools is dis- and its breakdown, The fat of the intestine oumsed in Section 3.16. and mosenteries is used up almost entirely. This process is much less intensive in males. 3.53 Responses to stimuli Acoordíng to Belyanina and Ì4akaroira (1965) a female smelt uses up about 65% of initial fat Kendall (1927), Baldwin (1948), Dryagin during epawning, 53% being removed as fat of (1948) and others have noted that adult smelt extruded eggs and 12% being used in activity. avoid strong light. Strong light destroys A male uses up about 47% of its fat, fat in developing eggs. Hatching larvae react extruded reproductive products being 6-10%; positively to light (Unanyan and 5oin 1963). a great part of the fat is used in activity during spaming. Unanyan and Soin (1963) noted that seawater of 26 O/<, salinity has a harmful effect Sudocrine systems and hormones and osmotic on the reproductive products of smelt, prevent- relat,ionc of the smelt have not been studied. ing fertilization. Seawater of salinity above FRrn/s Sri t 3:21

TABLE XXI

Chemical composition of smelt (from Levlyeva,1952and Kleimenov,1962)

Organ Month Chemical composition (%) Calorific Water Fat Total ProteinExtractiveTotal protein Ash value N N N ..N6.2

Nova river* males May 75 4 5.2 2.67 1.97 0.70 16.69 2056 (entire) June 79.5 3,3 2.39 1,96 0.43 14. 94 2,3085.2-89.7

Fern ale s May 76.4 3,9 2.77 2.13 0.64 17.31 2,52 (entire) June 81.4 2.1 2,27 i . 84 0.43 14, 19 2.31

bodies June 81.4 1.4 2,49 1.94 0.55 15,56 1.84 heads May 78,4 2.9 2.20 1,43 0.77 13.75 4 45 June 80.6 2.9 2,05 1,48 0,57 12,81 4.41 livers May 74.9 7.6 June 79.8 3,9 internal May 59.727.3 organs with- June 78.3 8.3 out reproduo- tive produots

ar East

e nt ire May 77-784 4- 16,1-17.0 1 O- 110.6 5.0 1.1

eggs 62 12,0 23,0 2.0

te st is 66 7,0 24,0 3.0

Contente inmg/bog

IC Ca Fe 1MG 388 83 07

Aminoaoid oontent as % of protein

Isolouoino Leuoine Lysine MethioninoPhenilalanine Treonino Va].ino

4.8 .8 11,1 2.8 4.0 4.3 5.4

There ae 2340 international units oi viosin D in i; of omolt livor fat and 170-200i.u. in i g o:':'r of uitomc'lognnr 3:22 FRm/578Smelt

PABLE XXII

Fat content of opaming smelt from different populations

Locality Fa conten in % of we subs tance Authority body eggs testis

White Sea 1.7-3.9 8-9 5 Bolyanina,1966b Nova river 2.5-3.5 - Leviyeva,1952 Siberia 4.7 - 5.3 12 7 Kyzevetter,1949 Uoh inskoye 0.5 - 2.4 48 - 6 Original data waterbody North Germany 10.2 5.1 orawa,1956

TABLE XXIII

Localization of fat in smelt (as % of total fat)

Sex Organ Season AjilMa31st0ctobe White Sea Females body 44-53 59-65 72 gonads 45-57 2-3 7 other internal organs to4 33-38 21 ales body 78-88 63-79 gonads 6-12 4-6 other internal organs to16 20-33

Uchinskoye Females body 36 77 waterbody gonads 64 8 and other internal organs very little 15 Rybinolz waterbody Malos body 61 78 gonads 7 7 other internal organs 32 15 FRm/s78Srnel t 3:23

TABLE XXIV

Extent of upriver migrations in different smelt populations

Extent of Locality migration Authority km

Yenisey river 1000 Tyurth,1924,and others Lena river l80-200 Pirojnikov,1950 Amur river 270 Kuznetzova,1962 Suyfun river and other rivers of the Far East 16-.18 Ivanova,1955 Elbe river 120 Lillelund,1961 Rivers of the White Sea 2-3 Balagurova, 1957 and others basïn Saint lawrence river 300 Magnin and alieu,1965 Miramiohi river 50 McKenzie,1964

130/00 results in pathological changes in by migrating downstream0 Froehwater has a developing eggs: flowing out of yolk, deforma- bad effect on the survival of smelt larvae in tien of body of embryo, absence of orystallino later stages of development (Unanyan and Soin, lens in eyes, destruction of brain9 delay in 1963). Lillelund (1961) stated that salmi- hatching, sometimes death of embryos. Earlier ties below 10 0/00 have no harmful effect on stages of development (till complote gastrula- developing eggs and larvae, The upper limits tion) aro mora sensitivo to salinity than those of thermal tolerance during early stes of later (after the embryo body has formed). development of eggs lie within the range17.7- Hatching prolarvae react positively to salinity 20.7°C,at later stages within the rango 21- 24°C0

FRrn/578Smelt 421

4 POPULAfPION 4,13 Size composition

4.,I Structure ])ata relating length to age and maturity in different stoc.ks are given in Table VIII. 4,11Sex ratio The length-weight relationship of differ- According to Kirpichnikov(1935)males ent stocks is given in Table XIX. predominate in the majority of smelt populations. The number of maies decreases in older 4.2 Abundance and donsit (of population) ago-groups because of their shorter life cycle (Table xxv). A decrease in the relative 4,21 Average abundance number of males in each age-group may he seen on comparing the sex composition of spawning According to McKenzie(1964)the smelt and feeding concentrations, This may be stock of the Mirainiohi river is estimated about ossod by the higher total mortality rate of 375million fish; Lillelund(1961)estimated males during spawning (Tremblay,1946). the stock of the Elbe river of60-85million fish, The sex ratio on spawning grounds is discussed in Section 3.16.. 422 Changes in abundance 4.12 Age composition Dornrachev and Pravdin(1926),Petrov (1940, 1947),Dryagin(1948)and others have The age composition of different spawning noted aharp changes in abundance in short-lived populations is given in Table XXVI, Catches smelt populations. Lapin(1960)gave an analy- of feeding and, wintering concentrations in- sis of causes of that phenomenon, Sharp de- cludo some younger immature fishes, creases in abundance are related to poor densir of one or two year classes. Usually the popu- For age at first capture, age at maturity lation density recovers in several years, and maximum age see Table VIII, Most of the Sometimos a fall in density is a result of an spawning (and fishing) stocks ars formed by epidemic (Van Oosten,1947; Patterson,1948, fish of minimal spawning age - See Table XXVI. o' others). In most areas fishing has no

TABLE XXV

Sex ratio in different age-groups of White Sea smelt (original data)

Year and season Sex Ago groups Number of sampling 2 3 4 5 6 7-8 of fishes

1961 Spawning Females - 415 37,0. 100 100 100 184 concentrations Males - 58,5 63,0 270 (spring)

Feeding Females 52,1 5405 75.0 75.0 100 100 234 concentrations Males 47.9 45.5 25,0 25,0 192 (summer)

1962 Spawning Females 66,0 50,0 72,7 66.7 100 335 concentrations Males - 34.0 50.0 27,3 3303 - 282 (spring)

Feeding Female s 52,1 56,7 70,5 100 - 152 concentra oms Malos 470.9 43.3 29,5 127 (summer) FR/S78 Sme t

TABLE XXVi

Ago composition (%) of epawning stocks in different smelt populatione

Locality Age groups Authority 1 2 3 4 5 6 7 89 10 1112 13 14 15 Pskovoko-Chudskoyo lake 90 IO Moshkov & Sorokin, 1952 Rybinek watorbody 17 77 6 Sohetinina,1954 Dactey lake 74 178 1 Wilier,1926 Lazmiaden lake 6 37 38 154 Wilier,1926 Ladoga lake 6 24 32 25 ii 2 Arkhiptzova,1956 and others Onega lake 4284020 7 I Alevandrova,1963 irumiohi river 66 304 McKenzie,1964 uron lake 54 388 Baldwin,1948 Upper lake 324917 I 1 Be.yley,1964 Neve, river 1284224 3 1 Kojevnikov,1949 White Sea* Onega Bay 7 32 41 17 3 Balagurova,1957 Karidalaksha Bay 304623 1 Original de.te. Yenisey river 8 20 43 23 6 Kravohuk,1958 Lena river 6 148 39 28 5 Pirojnikov,1950 Chat ariga river 3 10 10 19 28 254 Luicyanchikov,1964 Anadyr river 17 21 33 13 7 4 3 1 1 Agapov,1941 serious effect on population density, but Moro- 4033Recruitment soya(1960)and Amatisiavsky(1963)have stated that intensification of the fishery has led to Slowly growing year classes mature and a sharp decrease of catohee in the White lake join fishable stocks later than those which (Vologcta district) and the Ob estuary. grow faster. Unpublished data chow that the 1958year-class of White Sea smelt grew slowly 4.24 Changes in density during the first year of life and reached maturity only at the fourth year (only20% Annual varìations in landingi per unit of matured at the third year of life). The1959 fishing effort for the Miramichi river, Canada, and1960year classes grew faster and matured are given in Fig.11. during the third year of life. Similar data are given by Morosova(1960)and Lillolund Smelt occur in reilatively shallow waters. (1961). Seasonal variations in the available stock Recruits enter the epawning stock in are mentioned briefly in Section3.52 spring. The relative number varies from year to year because of differences in the strength 4.3 Natality and recruitment of year-classes and in the rate of maturing (see above and4,22). 4.31 Reproduotion ratos 4.4Mortality and morbidity Rupp(1959) found that the survival rato of eggs of the smelt of Maine is about6%. 4,41 Mortality ratos According to Kojevnikov (1949)survival coefficient (from eggs to adult) in the Nova According to Lillelund (1961) average river smelt is about0.02%. fishing morta].il;y coefficient in the Elbe river smelt is 0,17; natural mortality coeffioient 4.32 Factors affecting reproduction for different age groups is as follows s

See Sections3,16, 3,21, 3.22, 3.23 Age 1 2 3 4 Mortalityco- 0.90 1,08 1,30 1,59 efficient FRm/S78 Smelt

McKenzie (1964) stated that average annual The epidemic cas apparently caused by a virus fishing mortality, coefficient in the Miramichi or a bacterium. river smelt is about 3-5% of the stock. Physical factors causing morbidity of adult 4.42 Factors causing or affecting smelt are not known. mortality 4.5 Dynamics of populations (as a whole) Predators: See Sections 3.15, 3,22, 3.34. Lillelund (1961) gave a model of dynamics Food of larvae and post-larvae and its of year-classes in the smelt population of the availability: See Section 3,22. Elbe river in 1953-60. Lapin (1960) has worked out a scheme of dynamics of age-groups in the Physical factors: See Sections 3,15, 3,17. population of thsmall smelt of the Rybinak Sometimes stocks of smelt become frozen in ice waterhody (Fig. is). This model is made on the during the spawning migration in Siberian assumption that the Rybinak smelt spawns once rivers. only before death, The model demonstrates that different rates of growth and maturing may cause Direct effects of fishing: See Section changea in the age composition of the stock. 4.41. 4,6 The population in the community amd Indirect effect of fishing: the ecosystem

Sa:nsonov (1910) showed that great mortal- Pirojnikov (1955) included the Siberian ity of eggs of freshwater smelt is caused by smelt in the Arctic freshwater complex of fish fishing gear. (Huol-jo hucho, Salvelinus fontinalis, Stenocius leuciohthys, Coregonide). Ponornareva (1949) 4.43Factors affecting morbidity noted that smelt larvae in the Kara estuary occur together with larvae of Coregonus Parasites aoci diseasez: See Table XVII autumnalis, Salvelinus fontinalis, Myoxocephalus scoxyius, Liopsettalaoialis and Eleginus It is well known (Van Oosten, 1947, and n avaga. others) that mass morbidity and mortality of smelt occurred in the Great Lakes in 1942/43. See also Sections 3,33, 3.34, 3.42

- mature fish of one-year old

- mature fish of two-year old

m'il» - maturo fish of three-year old

- immature fish

Fig. 15 chemo of dynamics of year-classes in populations of small smelt (fromLapin, 1960), Seasons aro shown on the horizontal axis, body lengthon the vertical. Arrows show that an age-group (or part of it) han reached maturityand spawned in ,pring (circles signify laid eggs), The dotted line on the level of 60 mm shows that smaller fishes remain immature, Two cases (A and B) with different initial age structure of population are considered,

FRm/S78 Smelt

5 EXPLOITATION Smelt is frightened by light, and effec- tive methods of attracting the fish are not 5.1Fishing equipment known.

5.11 Gears 5.12 Boats

In the Eibe river they catch smelt in In Europe smelt are usually fished from river-stow-nets, which lie on either side of motor boats of 12-14 m long and 4.5 n wide of a cutter, or the nets may be anchored to the 40-75 hp (Lillelund, 1961). In North America bottom of the river. Each cutter has one or catamarans and man-propelled soows of 6-9 m two river-stow-nets (Lillelund, 1961). In long aro frequently used (McKenzie, 1964). the Groat Lakes they use nylon gill nets of 25-38 mm mesh or smaller (extension measure); 5.2 Fishing areas these are 75 m long. Otter trawis are used also; these are semiballoon type, with head- 5.21 General geographic distribution rope lengths of 945-1250 cm and mesh cizes of 62 mm in the wings and body and 12 mm in the Within the area of distribution (see Fig. cod end. Shore seines are used during the 9) smelt are fished in coastal waters of seas, spawning run; these are of 12 mm-mesh cotton estuaries of rivers and lakes. During the twine, 180 cm deep and 76 m long. Smelt spawning period smelt are fished in rivers. from rivers are captured at sea-lamprey bar- riers In weir traps of 120 x 180 x 180 ein 5.22 Geographio ranges constructed of 6 mm-mesh (bar measurement5 hardware oloth over wooden frames, Spawning See Section 5.21 run smelt are captured in commercial pound nets 210 cm deep made of cotton twine of 30 mm Smelt are moet abundant in the north- mesh, shrunk to about 24 mm mesh through treat- western part of U.S.S.R. and the Arnur river ment with prebervatives. The boom shocker basin, Atlantic coasts of Canada and U.S.A. consists of two electrical wire paddies sus- pended from booms extending ahead of a 16-ft New rangos of smelt fishery aree Great outboard-powered boat. Power for the shocker lakes, where the smelt was introduced in the in supplied from a portable 2.5-kw, 220-volt, first quarter of the twentieth century 3-phase, AC generator (Bayley, 1964). (Speirs, 1951); estuaries of Siberian rivers, whore the smelt fishery did not develop until According to McKenzie (1964), In the after 1935; artificial waterbodies of the Miramiohi River "open-water fishing gear is Upper and Middle Volga system. used at the beginning of the season unless the ice is strong enough to set winter fishing 5.23 Depth ranges gear. Open-water gear consists mostly of bag nets set on stubs or pickets near shores pro- Smelt occur In midwater or near the bot- tected from the full force of storms, The tom in relatively shallow places. The young gear is fished from catamarans and man-propelled stages carry out vertical migrations in pur- scows 6-9 m long". suit of plankton.

In Russia, smelt is fished by special 5.3 FIshing seasons traps ("mereja") made of email-mesh cotton twine stretched on 4-6 hoops of wood with two 5.31 General pattern of season(s) wings. Traps are fixed by anchors at mouths of spawning rivers. Shore seines of 80-170 m Smelt are caught mainly in spring, during long with mesh sizes of 32-30-28-26-20-18-16 mm spawning. The fishing season Is rather short in wings and 16-12-8 inn (inwards) in the cod (see Sections 3.16, 5.33). In autumn the end (Tyurin, 1939). Large river-fixed-nets fishery is based on less denso concontrations and trawls are used also. than in spring. In winter angling is usod (McKenzie, 1964, and others). Angling for smelt ocours also in some regions (Canada, north-western part of USSR). 5.32 Dates of beginning, peak and Baits are earth worms and young fishes, end of season(s)

During the development of the fishery The beginning of the fishing season coin- there has been a tendency to change from simple oldes with the formation of offshore feedin traps and cotton gill nets to nylon gill nets, concentrations In autumn and continues In trawling and electrical fishing. winter. The peak of the fishing season is based on spawning concentrations (seo Table Echosounders and fish detectors aro not xv). The season onde after spawning when used by smelt fishermen. smelt have migrated to deeper waters. 5z2 PR/73S

5,33 Variation in late or duration 5.42 Selectivity of season Bayloy (1964) reported that "the gill The fishery for spawning smelt is influ- nets from 25-125 cm mesh captured almost ex- encod. by water temperature and other factors clusively fish of age group II or older. The which control spawning (see Section3.16). shore seine, weir trap and pound-nets, fished on].y in the spring, sampled solely the spawn- According to McKenzie (1964) the fishing ing population. Since segregation by maturity season in the Miramichi river basin from1931- precedes the spawning migrations, immaturo ape- 1963varied irregularly fron77to136days. cimens (all fish of ago-group I and part of This variation in length of season appears to açe-group II) were absent. The otter trawls have little effect on the final total catch. and boom shocker fished at times other than the spawning period, appeared to be the least sel- The highest proportion of large smelt is ective gears. Trawl collections usually in- caught early in the spawning season. Size oluded smelt of all sizes and age groups. The of fish declines as the season progresses, and boom shocker fished only in shallow water seem- the price of fish decreases. ed, to sample adequately in the waters fished". Fishing operations and results 5,4 According to McKenzie (1964) "Increasing the mesh size from 29-32amincreased the gill- 541 Effort and intensity ing 5times arid, in the38mm not7times. Most smelt under 10 cm escape the commercial nets. Lillelund and McKenzj,e (1964) have (1961) Sise selection by the nets is one of the fac- given statistics of the smelt fishery in the tors causing diminution in the size of smelt Elbe river and the Miramichi river for many caught as the commercial season progresses", years, including data on catches per unit of fishing effort, Data on Miramiohi annual 5,43 Catches commercial smelt landings,1931/32-1962/63, and catch per licensed net per season are given Total annual yields for various countries in Figs.16, 17. are given in Table XXVII and average yields for different f ishing grounds in Table XXVIII. thousands of owt 2

211

11 0

3 2

r ye a years 9 9-60

Fig. 16 1iranichi annual commercial smelt Fig. 17 Miramiohi smelt oatoh per licensed landings,1931-32 to 1962-63. net per season (from McKenzie, 196L1). (from McKenzie,1964). FRm S8Smelt

TABlE XXVII

Total annual yields of smelt in various countries (thousands of metric tons) (Yearbook of Fishery Statistics FAO,1966)

Country Years 1958 1961 1962 1963 1964 1965 1966 Total 25,0 27,0 25.0 25.0 29,0 28.0 28,0 U.S.S.R. 14.5 15.1 15,3 15.5 16.1 16,7 16.6 Canada 4.3 7,3 1.5 1.6 2.1 2.1 2.1 U.S.A. 6.0 2.5 2.2 2.3 2.4 1.8 1.4 Finland 0.1 1.1 0.6 0.4 0.7 0,8 0.9 France - 0.3 0.2 0.1 0.1 FederalRepubilo of (hrmany 0.4 0.2 0,2 0,2 0,4 0.3 0.3 Netherlands 0.2 0.1 0,2 0.1 0,2 0,3 0,3 Norway - 0.9 - 0,1 0,6 0,2 0.2

TABLE XXVIII

Average annual yields from different fishing grounds

Fishing grounds Average yield Years Authority (tons)

Pskovsko-Chudskoye 3600 1931-1963 Shlrkova & Pihu,1966 lake limen lake 363 1946-1958After Kudereky,1962 White lake 261 1945-1958After Kudersky,1962 Onega lake 623 1945-1959After Kudereky,1962 Ladoga lake 1374 1946-1954 After Kuderaky,1962 Neya river 400 1932-1948After Kojevnikov,1949 Finnish Gulf 640 1932-1948 After Kojevnikov,1949 Kurishes Haff 983 1948-1957 After Noskov,1959 Elbe, Eider, Weser 311.5 1955-1959Lillelund,1961 White Sea 400 Yenisey 150.7 1946-1955 Podiosnyy,1958 Ob estuary may. 1540 1960 Amstislavsky,1963 Amur 1000 1949-1958 Kuznetzova,1962 Miramiohi river 726 1931-1963 McKenzie,1964 Great lakes 7264 1960 Bayley,1964

FRm/S7U Smelt 6l

6 PROTECTION AND MANAGEMENT break the swift smooth flow, The bottom of 6,1 Regulatory (legislative) measures the culvert at the outfall (at least when built) should be level with, or preferably low- 6,11 Limitation or reduction of er than the stream bed. Action to facilitate total catoh the passage of smelt through highway oulverts to their spawning grounds would be of consider- In many oases catches of smelt may be in- able importance to the smelt fishery in New creased, as only a email part of the popula- Brunswick, Smelt often escape the strong cur- tion is removed by fishing. However, in some rents in streams during spring froshets by cases it is necessary to limit catches going out over banks and spawning far from the (Morosova, 1960; Ametielaveky,1963). regular stream bed, In some locations it seems practical to prevent loss of both eggs 6.12 Protection of portions of and parents from stranding by constructing population dykes. In the larger Miramiohi branches and tributaries flood control by means of dams The size of caught smelt is limited by would no doubt increase the larval production". the market8 for example in the Elbe river they do not catch smelt snaller than 14 cm 6.5Artificial stocking (Lillolund,1961). 6,52Transplantation, introduction 6.2 Control or alteration of physical features of the environment Transplantation of' smelt from the Green lake (Maine) into the Crystall lake (Michigan) Physical features of the environment may was made in 1912 and in following years and be altered to Improve spawning grounds by has given excellent results (Van Oosten,1931; clearing such impassible obstacles as brush Creamer,1926, 1929,and others). Smelt has jams, old dams, steep rapids or falls, sluice- spread all over the Great lakes and formed a ways, pulpwood logs as well as road oulverts. good fishery population there (Table XXVIII). McKenzie (1964noted that "at moderate cost In Russia smelt was repeatedly transplanted much can be done to overcome these situations, into some lakes of the Leningrad and Novgorod Streams full of brush and debris from lumbering Districts (Píkhiy,1941). These operations operations oan be cleared and laws enforced have given good results in42%of oases for to prevent recurrences. Old dams can be re- small smelt but for long-lived smelt only17% moved or if in use can In some cases be opened have been successful. As a rule unrepeatod during the smelt spawning season. "Smeltways" transplantation has rarely given good results. can be built also to permit passage over the Snielt wan introduced into some Ural lakes also dams. Grades through culvorts should be made (Karabak,1930), Some introduction of smelt to conform to the slope of the original stream into the waters of Karelia were proposed by bed with addition of baffles, if necessary, to Stefanovskaya (i957).

FRs 'S Smelt 7:1

7 POND FISH CULTURE during5-6days i iena than 5% (lcarabak,1930). Good survival of transported smelt eggs and 7.9Transport larvae was noted by Richardson and lknap (1934),fishes of 6-10 cm length being the moat Smelt has not been cultured in ponds. hardy. Early larvae are too de].icate and they /ccording to Karabak (1930) and Suvorov (1939) should not be transported. Smelt eggs aro smelt eggs may be incubated and transported in gathered on speoal i:creens during the spawn- a cool wet atmosphere. 4ortality of eggs ing period.

FRrn/S78 Smelt 8i

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SYNOPSIS OF FISHERIES BIOLOGICAL DATA

This is one of a series of documents issued by FAO, CSIRO and USFWS concerning species and stocks of aquatic organisms of present or potential economic interest. The primary pur- pose of this series is to make existing information readily available tofishery scientistsac- cording to a standard pattern, and by so doing also to draw attention to gaps in knowledge. ltis hoped that synopses in this series will be usefulto other scientistsinitiatinginvestiga- tions of the species concerned or of related ones, as a means of exchange of knowledge among those already working on the species, and asthebasisforcomparative studyof fisheries resources. They will be brought up to date from time to time as further information becomes available either as revisions of the entire document or their specific chapters.

The relevant series of documents are:

RAO Fisheries Synopsis No. FR/S replacing, as from 1.1.63 FAO Fisheries Biology Synopsis No. FB/S

CSIRO Fisheries Synopsis No. and DFO/S

USFWS FAO Fisheries Synopsis No. BOF/S

Synopses in these series are compiled according to a standard outline described in Fib/Si Rev. 1 (1965).

FAO, CSIRO and USFWS are working to secure the cooperation of other organizations and of individual scientistsindrafting synopses on species about which they have knowledge, and welcome offers of help inthis task. Additions andcorrectionstosynopses alreadyissued will also be most welcome. Comments including suggestions for the expansion of the outline and requests for information should be addressedtothe coordinators and editorsofthe issuing organizations.

FAO: W. Fischer Fishery Resources and Exploitation Division Marine Biology and Environment Branch Food and Agriculture Organization of the United Nations Viale delle Terme di Caracalla 00100 Rome, Italy

USFWS: CSIRO: L.W. Scattergood, Chief, Branch of Reports Maureen A. Wright, Scientific Editor U.S. Department of the Interior CSIRO Division of Fisheries and Oceanography Fish and Wildlife Service Box 21 Bureau of Commercial Fisheries Cronulla, N.S.W. Washington, D.C. 20240, U.S.A. 2230 Australia

Consolidated lists of species or groups covered by synopses issued to date or in preparation will be issued from time to time. Requests for copies of synopses should be addressed to the issuing organization.

The following synopses in this series have been issued since January 1969:

BCF/S40 Synopsis of biological data on the Pacific mackerel Scomber ja pon/cus Houttuyn (Northeast Pacific) February 1969 FRi/S30 Synopsis of biological data on the Pike Esox lucius (Linnaeus) Rev. i 1758 May 1969 FRm/S43 Synopsis of biological data on the anchoveta Caten grau/ls mysti- cetus Günther, 1866 October 1969 DFO/S3 Synopsis of biological data on the Tiger Prawn Penaeus escu- lentus Haswell, 1879 July 1969 FRh/S35 Synopsis of biological data on West African Croakers Pseudo- Rev, i to//thus typus, P. senegalensis and P. elongatus November 1969 FRm/S78 Synopsis of biological data on smelt Osmerus eperlanus (Lin- naeus) 1758 December 1969 MI/96024/3.70/E/1 /750