FRm/S78'AO Fisheries Syhosis o. 78SAST - smelt - 1,23(04),003,O1(Distribution restricted)
SY LJ' OF BIOLOGICAL DATA ON SMELTOsmerus epe?kus (Linnaeus) 1758
19i'cpard by
T. 1. BeI nina
FFOOD AND AGRICULTURE ORGANIZATION OF THE UNITED NATIONSROME, 1969
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FAO Fieheries Synopsis No078(Distribution restricted) SAST - Smelt l,23(04),003,O1
SYNOPSIS OF BIOLOGICAL DATA ON SMELT
Osmerus perlanua (Linnaeus) 1758
Prepared by
T.L BELYANINL
isltuta of EvolutionaryMopho1ogy and Ecology of Animals
Academy of SoncoaMocow, U.S,S.R
FOOD AN]) AOl JIWLUlDii O)GA'lZATION OF TUE UNITED NATIONSìmiu9 Thoombor 1969
FRm/878
The preparation of this Synopsis was promoted in view of the need to review theabundant und much scattered information on the biology and exploitation of Osmeruseperlanus 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 thecourse of personal research work on the species and also, on information received fromvarious sources, most of which are listed in the Bibliography.
The author and the editor are grateful to Dr. D,I. Williamson, Marine BiologicalStation, Port Erin, Isle of Man, U.K., for his editorial work on the synopsis.
Distribution
FAO Department of FisheriesFAO Regional Fisheries OfficersRegional Fisheries Councils andCommissionsSelector SMAuthor
PREPARATION OF THIS SYNOPSIS
"Current Bibliography" entry
Belyanina, T.N. (1969) 16-6B228FAO Fish.Synops., (78)zpag.var.Synopsis of biological data on smeltOsmerus eperlanus (Linnaeus) 1758
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 andmanagement. Pond culture.
23 Determinan
22i Spawn, larvae and juveniles222 Adults
s of dis
24 Hybridization
24i Hybrids; frequency of hybridization; species with uhiohhybridization occurs; othods of hybridization
2q42 Influence cf natural h1r1dization in ecology and morphology*
3 BIONONICS AND LIFE HISTORY
31 Reproduction I
3l1 Sexualiiy I
312 Maturity313 Mating i
3q14 Fertilization j
3l5 Gonads j
316 Spawnirtg 33q17 Spawn 3
32 Proadult hase 9
321 Embryonic phase3q22 Larval phase3q23 Adolescent phane
33 Adult phase (mature fish)
3q31 Longevity3q32 Hardineec333 Competitors334 Predators3o35 Parasites, diseases, injuries and abnormalities
ribution 3han.es
CONTENTS
FRm s78 Smelt iii
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 31032 Cytomorpho logy 71q33 Protein epocificity 9
2 DISTRIBUTION 2:1
21 Total area 1
22 Differential distributic-, I
99
11
3:13
13131313
13
II
2
2
3:1
1V FRm/S78 3mejt
Page
3.4 Nutrition and growth 3:13
3.41 Feeding3.42 Food3,43 Growth rate3,44 Metabolism
3.5 Behaviour 20
3.51 Migrations and local movements 203.52 Schooling 203.53 Responses to stimuli 20
4 POPULATION 4: 1
4.1 Structure i
13171720
4.11 Sex ratio4.12 Ago composition4.13 Size composition
4,2 Abundance and density (of population) i
4,21 Average abundance i
4.22 Changes in abundance i
4,23 Average density*4.24 Changes in density 2
4,3 Natality and recruitment 2
'1.31 Reproduction rates 24.32 Factors affecting reproduotion 24.33 Recruitment 2
404 Mortality and morbidity 2
4.41 Mortality rates 24.42 Factors causing or affecting mortality 34.43 Factors affecting morbidity 34044 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 distribution5,22 Geographic ranges5,23 Depth ranges5.24 Conditions of the grounds*
5.3 Fishing seasons
5,31 General pattern of season(s)5.32 Dates of beginning, peak and end of season(s)5.33 Variation in date or duration of season
i
i
i
2
FErn 578 Smelt
* As no information was available to the author, those items have been omitted from the text0
V
Pe ge
651 Maintenance stockjng*6,52 Transplantation, introduction
7 POND FISH CULTUPE 7:1
71 Procurement of etocks
7,2 Genetic selection of stock*
7°3 Spawning (artificial; induced; natural)*
7,4 Holding of stocke
sto0 )*7,5 Pond mama ement fertilization' acuatic .lant control;
7,6 Foods; foeding
7,7 Disease and parasite eontrol
7,8 Harvest*
7,9 Transport
8 REFERENCES 8i
54 Fishinc' oserations and results 5:2
54l Effort and intensity 25,42 Selectivity 25,43 Catches 2
6 PROTECTION AND MANAGEMENT 6:1
6 I Regulatory (legislative) measures
6,11 Limitation or eduotion of total catch6,12 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
FRm/578 Smell; i :1
1. IDENTITY
I I Nomenclature
1.11 Valid name
Osmerus eperlanus (Limnaous, 1758).Original combination: Salmo planus
Linnaeue, 1758, Syst. Nat, (ed. IO), p, 310
t,12 Objective synonymy
Salmo eerlano-marinus Bloch, 1782,Naturgeech. d.. Fische Deutschlands 1, p. 128,
P1,28, Fig.1.
Osmerus operlanusz LacpMe, 1804, Hist,nat,des Poissons, 5, p, 229,
Salmo (Osmerue) Pallas, 1814,Zoogr.Ross.Ae., 3, p, 386
1.2 Taonomye
1,21 Affinities
Supragenoric
Supeiphylum-ChordataPhylum-VertebrataSubphylum-.Crani at a
Suporciass-UnathostomataSeries-PincenClass-TeleostomiSubclass-Act inoptnrygiiOrder-C lupo iformeSuborder-Salmomo de iFamily-OcmeridaeSubfami ly-Osmerinae
Generic
Osrnrus Linnus, 1758 Syst.Nat. (ed, io)
p, 310. Typo specen Salmo er1anuLinnaous, 1758e /L-nmaeua iiI) adopted foursub-divisions of his gstus Sa1rno one of whichhe named :Osmori:, The go'er?o nane OsmerusLinnaeus, 758, wih ype-s»ecies Salmoeno Linnacuc, vaiidct.od in Direction69 o! the Intor onJ Commiosion on Zoologi--cal Wononc) aturo,,j$flfl lnro canine n t1ier cido ci' vomer,somotimeui nocop od by smaller ones, Pala-
tine teeth exilared aneriorly. t'lazilla.ry ox-
tondo be obout posterior border of orbit. Noobrino on opcvole or ovboporole, Dictanocfrom onoub lo oroal oviim ohortor than fromdorsal oigm o orwsl bacs, A midlateralrido, bu no anal shelf or e1onga'ed midlate-rol coalos in moie. Gill-rakern 25 to 37.Pylorie cocoa 3 be 8 Stomach itti blind sao.
*After Berg (1948) and McAllister (1963)
Lower jaw spatulate, Anal rays 11 to 14.Midlateral scales 58 to 72. Lateral line in-complete. Adiposo baso about two-thirds oforbit diameter (rarely in large specimens equalto orbit). Orbit diameter two-thirds or lossof caudal ped.unole depth, Proothmoids double.Mosothmoid simple, without other ossifications.Parietale completely soparatod by supraoccipi-tal. Pterosphenoide not reaching parasphenoidanteriorly. Slit present between hyomandibular. Four simple actinonts", (McAllister,1963).
Anadromous or landlocked forms in thePolar, Pacific and Atlantic Oceans and theirdrainages.
Subjective generic synonyms
E2lanus Gaimard, 1850-52, Voy. on Islandeet Groenlanci, Paris, Atlas, pl,XVIII, fig.2(type-ope oies: Eperlanus vulgarie Gaimarci).
Allosmerus Hubbo, 1925, Proc.,Biol,Soc,Wash.,38, p, 53. type-species: Osmerus attenuatusHubbs).
Specific
Three forms of the boreal smelt have gen-orally been recognised; O. of theNo'th and bitic ens, O. dentex of bho Paoificnortk into the Arctic and west bo the White Sea,O, ordax of the Western Atlantic. Followingdetailed studies of their populations Berg(1948), .driashov (1954) and other Russianwers'.o reduced dentex to a subspecies ofeperLanus, Some american authors (Kendall,
1927; McKenzie, 1958, and others) consider theamerican smelt ne a separate species, butBigeÏow et al. (1963) and MoAllistor (1963),who made a rsirnion O.' the Osmeridao, considoindthat the snuo Osmerus includes only one spoces,O, esrlanus (L).
1,,22 Ta,xonomio status
t r a norpho.apooioo, a polytypio opooioowith may goog:aphioal cn1 000logical formo.
1,23 Subspecies
Key o nubopn&eo of Osmerus eperlanue(from 1oAilis1or 1963).
1(2) Po-od acabo in the lateral lino usually14-'28 (roroly 13.-30). Ucotorn Atlontie, Pacif-ic c-nd koobio uot to the hito Soc. c-nd. their
ctoJaçon , , . n , n . o Q o . Q . . . , ' . o Q Q O n , o e e e . e oOrirnorun 2?!)US.. ïU!2 (Miioh1b)
i 2
2(1) Pored scales in the lateral line usually6-13 (rarely 0--16). Baltic and North seas andtbir drainages and Upper Volga River0vris eperlanus eperlanue (Linnasue)
Osmerus rlanus mordax (Mitchill) 1814*
Atherina mordax Mitohill, 1814, Po 15,Ìou York (Speirs 1951, indicated that this wastho type description)0
Osmerus viridescens Le Sueur, 1818, p. 230,coast of Maine,
Osmerus Norris, 1868, p. 93,Sohuylki],l River, New Jersey (described but notnamed by Norris, 1868, p, 58).
(Osmorus soronti auctoruni).
Osmerus speotrum Cope 1870, p.Wilton Pond, Kennobec County, Maine,
Osmerus dontex Steindachner, 1870, p.Dekastris Bay, USSR.
Osmerus dvinonsis Smiti, 1882, p, 32,Northern Dvina River, Russia.
ho following natios named by Petrox,1925 p, 82, 108, have no status under the rulesof the Int,Comm,Zool,Nomencl Osmerus e, dentexroao kaninensis, Choshskaya Gulf; O,e, dento.xnabo nisseensis, Enisey River at Tyurin andsome other localitie7.
Diagnosis
Distinguished from other osmerid speciesby the two large canines, one on either sido ofthe vomer opercular striae; the maxillary ex-tending past the pupil. 4 to 8 pyloric oaeca,Phis subspecies is distinguished from O.e, oper-lanus by the greater number of pored lateralline scales, usually 14 to 28,
Description
D 8.-io(ii); C 19; A (12)13-.15(16); V 8;P 11-13(14); LL (13,14)1528(29,3o); midlat-.eral scales (62)63-69(72); vertebrae (58,59)60-66(67,68,70); gill--rakers 8-11 + 18-24(26) 27-..36(37); branohiostegals (6)7-8; pylor-io caoca 4--8, Standard length 3.8 to 4,4times head length, 4,7 to 7.4 timas depth.Pectorals reaching from one-half to two-thirdsthe distance to pelvic insertion; pelvic, alittle more than half way to anal origin,Adipose base short, about two-thirds of eyediameter.
O Diagnosis, description of subspecies endsynonymy by McAllister (1963:15-21),
Medium sized canines on dontary, enlargingposteriorly. 5mal]. pointed teeth on premaxil-iary and maxillary. Canines on anterior endof tongue enlarged. Pelvic origin anterior todorsal origin. Ductus pneumaticus attached toanterior end of gas bladder, Latera]. line in-.complote, ceasing about a head's length alongthe body.
Peritoneum light with dark speckles, moreintenso dorsally. Ventral portion of bodypale; dorsum speckled with black, more intense-]y around exposed borders of scales, Chin andtop of head evenly speckled. A dark medio-lateral bar without sharply delimited border.In life sides an iridescent silver, the back anolive green. Total length to 324 mm accordingto Berg (1948), A larger subspecies than O.e.
nus.
490, Oernerus eerlanus eporlamus(Fig. 1).
FPm/578 Smelt
(Linnaeus)
429, Salmo eperlanus Linnaeus, 1758, p. 310.European seas and rivers,
Eporlanus Schonfoldii Rutty, 1772, p. 358,Ireland,
Salmo e erlano-marinus Bloch, 1782, p. 229,Table 28, Fig. 1, northern and Baltic, seas andinto rivers.
Salmo eperlanus var, marinus Walbaum,1792, p. 57.
Salmo spirinchus Pallas, 1814, p, 387,lakes and rivers. Germany, European Russia,(Boloczero, Chudsk, and other localities).
¿?Jl'lanua Rcndeletii Willughby, 1789,p. 202, Table 6, Fig. 4, Anadromous in Thames(unavailable since it is a reprint of a pre-Linnaean edition originally published in 1686v
Salmo eperlanue marinus: Walbaum, 1792,p. 57 (ex. Bloch),
porlanus fluviatilis Gesner, in the syn-onymy for Salmo eporlanus of Donovan, 1804,p. 189 (not available, a pre-Linnaean name citedin a mynonymy7,
Eperlanus vulgaris Gaimard, 1851, p. 207,p, 18, Fig. 2, Iceland (this species not knownin Iceland before or since this time accordingto Saemundsson, 1949).
Osmerus e. eperlanus natio ladogensisBerg, 1932, p, 281, Lake Ladoga (infrasubspeoif-io nanee have no status under the Int.RuleeZool,Nomeno].7
FRm/578 Smelt
Diagnosis
Thin subspecies is distinguished from 0.e.mordax by the lower number of pored lateral linescales, 4 to 13.
Description:
D 7-9; C 19; A (11)12-13 (14); V 8;P 11-12(13); LL (0)4-13(16); midiateral scales
(58)61-69; vertebrae (55,56)57-61(62,63);gill-rakers 25-37; branchiostegls 7(8), py-bric caaca 3-7. Standard length about 4 to 5times head length, 5.6 to 6,1 times depth;pectorals extend one-half to two-thirds thedistance to the pelvic insertion; pelvics abouthalf-way to anal origin. Adipose base short,about two-thirds the orbit diameter.
Medium sised canines on dentary, enlargingposteriorly. Small pointed teeth on premaxil-lary and maxillary, Ductus pneumaticus attach-ed to anterior end of the air bladder, Lateralline incomplete, ceasing less than a head'slength along the body.
Peritoneum light with dark speckles. Backgreenish, sìdes silver in life. Total lengthto 307 mm according to .rg (1948). A smallerform than O,e. mordax,
1,24 Standard common names andvernacular names See Table I.
TABLE IStandard common names and vernacular
names of Onmerue erlanus (Le)
Country
Englandfr an ce
NetherlandsFinlandSwedenNorwayCanadaUSAUSSR
Japan
Standard commonname
smelteperlanemaeltkuorenorsmorssmelt, éperlan
eltkor iushka
kyuri-uwo
Vernacularname
rainbow smeltzubatka, ogu-reohnik, korehasnetok, tint,norms, ealaka,
1.3 Mphobog'
1,31 External morphology
McAllister (1963), who studied the geo-graphical variability of smelt, came to the
1:3
conci-) on Sat ho only difference betweendentcz nn mo'cisc in hc vertebral number
YÍ ho"ovor io may be because oflower -n-n' ornra turc during the developmentof dcnty ogn T1o&lUs;or therefore con-siderod TonL synonyr of morciax. Mordaxarid nper nu di. ífer coaderab1y (especiallynumber c' pornrl i ntercl line scales - Table VI),Using }i) cbnr-' c-ter oloris over 90 percent ofall bs cpnciox'n or ho-i-b forms can be separatathSinco bn i-o forma oro olbopatric and sinosvertebral and snal ray differences all ovorlapconsiderably it is necessary to consider thetwo forms as aubapecifically different, Thedifference is nonolinal (McAllister, 1963).
Data on plastic characters of differentsmelt populations of urope and Asia are givenin Tables III-V, oil rieristic characters inTable VI,
Plactic characters of the smelt vary con-siderab]y with ago (especially height of finsand their situations, ddth of forehead, eyediane ter ) , J( rp cbui)or ( 935 ) found thatheight cT fino decronoen lTiih grouih corrola-tion coe:í'iccnj hrnn,, boight of A andstandard Jongth (L) thc TTh ta Sea smelt were-0,40 anc - 045 (for venous samples), betweenheight of D and L ucrn -'0o3ß, -'0.42, -0.73;botucu P and L oro -0/5 cud -0.47, botwoon Vand L S-0,54. iIyO diO!1Otfl' PicO chews a nega-.1ve oorrolmtoa 0o79 sud -'075. Snout
longih sd foreman udih theroaso uith age:tha corrototini oocff: cionn unro i 0,66 and+00 75. VoIries o f--V P-iY, D- also increase,although tho corvo) eion 000ff'i cint is ratherlow (ua:oJJy lesF íhr -;J.50)0
The smelt scale is very specific (Figs. 2,3), t'The smaller yr-.-n.old smelt has scalesui-th rolo tJ.rciy L-Lrgo focus and completecircul i. i ri ho Oj (;oo nth Çi,o i rhoroas rica
oC ihn .1 orger fT n) boro n rina31 or focun rind. oneto several completed ci roui t0 floih typos how-erar exhibit clearly the diriuon io features ofa grañuoi rhon ii of tSr horncahoe-shaped ini noomplo tr ci ncn ï on Grouth ni ors ïn lato sea-son and a distinct cutting over the first cir-culun of the second growing season. Later an-nubi are easily recognizable. A growth fieldcontains n of complote circuli thatgradually 5QHl .s growth slows. The firstcirculus that mpears in the following growingseason cuts a so these shortened oirouli°(Baybey, 1964) The year anriuli form as scarsoutside tli' o of incomplete circuli of thelast growilg'Hon (Figs.2, 3), Annulusformation otao-i near Juno. Circulus numberin 'the successive years of growth in differentsmelt populations is shown in Table VII.
14
Fig0 i Osmerue sperlanus eperlanue (L) (from Berg, 1948)
Fige 2 Scale cf 2 yearoldsmelt. White Sea.MEgnif. 6 x 3,
/s78 Smelt
Fige 3 Scale of 5 yearoldsmelt. White Sea.Magnif. 6 x 3.
FRm/878 Smelt i s,
Locality
Mordai iperlanu1
1419 6-13(13-19) (o6)
60-63 58-61(58-65) (55-63)
23-4,4 23-26
13-.i5 12-13
9-10 79
(1112i3 11i2(13)
63-65 61-69
28-35 25-37
4-7 3o7
L O L0
(min -mc, )
Tbightof D
Characters
White Sea 127-67 7.0-9.8Chesha B 149-167 8,4Pechora River 161-170 10,8.
Pekov-Leke 16.3 9,8White L&o 16,7 bolFinnish Gulf 15,0 9,50nega-L 169 10.3Lados Y ':e 124-14,5 9,3-10,3Neya River 157 9,9Far E t 12,2-14,9 69-80Ycmimoy River 128-13,7 73.44
14.3
tThJO oro ovoraorj o. diin0niiz, vr1 uon of chorutoo o
Comparison o±' different formsof Osmerum (McAllister, 1963)
pies, not
TABLE II
Dentex
15-20(1 3-3o)
TABLE III
Data on fins ( 5 o body ] )(after Kirpiohnikov, 1935; Tircìckf, 1956, and others
Eo Ln3thof P cf V of D baco
2o412o 9130
138131130 G
1231393GO-1 3 2i33
Lnthof JI baso
119-1209123-132
1303
12413229
13214 3123
109-12.9
124
vertebrae 63-66(snir0omaco) (597o)
Head L/anal Iie1ht 26-32
Modal anal rays 13-14
Dorsal rays 8-10
Pectoral rays 12-13
idiateral scales 63-72
Gill rakers 26-35
Pylorio casca
82-ß6809278-03
805
827078 7
86
Looality
Lecaliy
hLo Sea
Dat
a on
head
(after Kirpichnikov, 1935;
21, 2-2 38
*Theee are averages of dif
nout length
(as
of
standard
length
5,7-7,0
Eye diameter
(as % of
head
length)
16,6-20,9
20,4-24,8
12,5-14,3
7,6-9,6
erent samples, not min.-max. values of characters.
TABLE V
Piasio oharace
of suielt (as % of standard length)
(a±'er Kirpiohnilzov, 1935;
Nikoisky, 1956, and others)
Caudal
D-d
peduncle
length
1hite Sea
12,7-16.0
48.5.4 45,8-47,8
45.3-49,7
25.1-27.3
21.2-23,4
70.7
18,8-21,7
115-13.9
Chesha Bay
13 0-1632
4.8-51
45,7-46,6
46,4-48,5
24,5-27,6
22.1-233
19.8-21.8
11,0
Pechora River
121-461
5.1-5.6
4), 3-46. 9
445-46.9
23, 5-27, 4
24,0-25,5
19.6-20.3
12.3
Pskovsko-Chud 1
14,6
5,0
47,2
46.1
2509
23,7
18,5
11,6
Uhite Laite
145
5.5
47,4
46,7
2401
22.9
18,0
10,6
FinntGh Gulf
15,6
5,4
47.6
47.9
26,4
224
18.6
12,2
Ortega Lake
1303
5,0
49,9
27.4
22.2
19.9
11.7
Ladoga Laize
15.6-16.3
4.7-5,0 4903-4908
49,1
27,9-28,1
24.2
704
19,9
11,3-12.5
isva River
15,9
503
48,6
26,6
22.6
19.7
123
Far ]ast
14.8-16.5
5.2
47. 2-47. 8
26.1-27.1
22, 1-22.2
70,2
-11.6-11.9
Ye nice y
14 3-14.4
4,7-52 45.8-46.2
46.9-47.7
25.9-28.1
22. 7-23.0
68.5-70,3
-12.1-12.7
Uoruey
5.1
48.8
49.3
27.2
23.0
71.1
-11.2
Chosha Bay
21. 7-224
6.0-6.9
21,1
20.4
13,1-13,8
8,9-9,3
Pechora Rive
19,9-21,3
409-506
19,0
-10,9-11,0
7,8-8,2
Paitov Lake
20,9
407
21,2
24.9
11.4
8,3
Uhite Lake
21,0
409
21.9
25.3
11,8
8,5
Finnish Gulf
2205
700
27.1
19,1
11.3
9,2
Onega Laits
2208
22,3
24.5
--
Lacioga Lake
28-22,5
702
20.6-24,8
20,5-21,4
13.0
11,2
Neya River
22,3
19,6
23,7
--
No r
223
19,9
25,1
9,8
14.8
Ye,isey Rire
2136-21,7
6,9
19,0-20,6
22.8-25,3
8,6
13.0
Far East
21 4-2 1,5
7.1-7,5
19,9-20,6
22,8-25,3
8,6
12,8
Nikoisky, 1956, and others)
Forehead
Lorer jaw
Upper ja
width (as
(as % of
(as % of
% of head
standard
standard
length)
length)
length)
Preanal
A-D
A-V
P-V
V-A
length
li
Head length
(z % of
etciard
len
White SoaChesha BayPechora RiverPakovso-Chudskoye IWhite LakeFinnish Gulf0nea LakeLadoga LakeNava RiverFar EastYenisey River1orayBalic SeaChukotkaWest Germany
TABLE VI
retio obraotare of c'ron' ro! populations(after IirpichnU:ov9 1935 iko:Iay9 56 Lillelund, 1961)
29-3826-3431-3532-3532-3732-3633-35
60-67 15-27 1-362-67 16-2458-63 7-8 2-358-60 - 2
57-60 - 2-359-62 8-10 2-3- - 2-3
4-11 2-3- 4-11 2-366 18-28 2
66-67 14-26 2-359-62 - -59-62 10-13
- i42055-60 4-H
tebrae
i32 (;tor:ology (Chromosomeco)
Svbrdson (1945) found that "the smelt hshort and slender chromosomes9 considerablysmaller than in the other species0 Theirnumber is the iouest amow tho Siodimh rep-
ntatiea of the Salmonida9 fanui3y, theircliploid number being merely 5t As a rule,ho7e,rer, bo th Vhaped and rod-ehaped chromo-conos occur0 There aro five paire V-ehapedchromosomes0 Two of them c. app:;:-o.tely oç'unl lantiì ha'eai ot]r threc p:ihao one markcdly long c'n0 Constriotioai
Pored Rzya ií Dscale, Uobrenoheci Branohod Umbin LL
TAiLE VII
Ciroulus number in the suogro h in different amai
5-10 2-4
7-0 3-48-9 37-9 38-9 3-4-
10-11 38-9 -
9 39 3
sive years ofpopulations
Locality
Rivers of EastEngland
Miramiohi River(min.-max0 )
White Sea(mino -max0 )
Yaar of grorthi 2 3 4
4 18 9-10
10 15 70Ol9 102O 4-o12
2-3 9 9 5 4-50-6 712 8-13 4-7
Authority
flacierman 1913
annie 1958
Original
Rays in Anched Branched
12-16
12:1412-1411-1311-13
12:14
12:1413
may oo di lly in the short arm of one oftbos3 3moc pirs The rod shapedohromoso»s y in length, Thus the longestrod-shaped pair at any rate in certain earlymotaphaoce may bo four tines ao long as thenhoreot0 One nf the rocI-ohaped pairs hasproximal conetrotion0 The naioois taa notstudied"0
Otieiannil:ov (1885), Cunnincham (1886 -oiod by Tillelunci, 961) and Lillolund (1961)have denc:'ihod rpo 000ytso o' the amcl,their nonbenos ond mioropyie0 Lons (1904)nudtod vitolîogonae!e of smelt0 ICuenoteov
FRm/578 Smelt 17
Fig., 4 Smelt oocytes during the juvenilephase (right) and the end of thephase of one-layer follicle,M*nif. 10 c 1XL Stuin ironhaomatox. (U',.ioAtheth)
l8 PRm 578 Smelt
(1964) gave a conparxtiv. ncJyois of oogonesisof the migrant ssel or tLic Ne-a river and non-migrant smalL-ci;od onolt (enotok) of the Pekovlake. He found $ha'i. the oocytes of the Novariver smelt are largar ho.n those of the Pekovlake at the sarce phauos cf oogenoeia. Oogene-eis has aleo been studied in the White Seasmelt. Acoordiig to Moyen (1939, 1944) theperiod of protoplasmatic growth of oocytes in-aludes the juvenile phase and the one-layerfollicle phase, During the juvenile phase theoocyte size in the Uhis Sca smelt is about -
l3.5-36p, the nuclear diameter is ll3-27p.The cellular membraao 5e tb5n and. hae no visiblestructures. The nytopicom Jo griular Thereare 7-10 nucleoli ki tho ivolsue (in a section).During the ono-lryer fol] lolo pheo the oocytesize io Iron 45 to il2/ ihe nuclear diameteris from 36 to 7605,u. f'.t the periphery of thenucleus there are 20-40 nucleoli. The oocyteis rounded, the cellular membrane is thin.There is a more intensely coloured oiroumnuolearzone at sorno distance from the nucleus whiolidisappears after vacuolization has started(Fig. 4)
The pori OU (ir trOphop]CF1T1rl(r groït.h ofsmelt 000ytoo may bu divided into 4 phucoc.During the phuse of marginal voouolizatior theoocyte diemotor io uboot l44/b the nuoleardiameter is abou 8i1u ! suries cf littlevacuoloc appear at the periphery o: the oocyte
'Ii '
fl41
i'
(Fig. 4); these vacuoles do not contain oil,During the phase of initial acounulation ofvitellin and oil the oocyte diameter is 0-160»., the nuoleun is oval but in section itsmargin i festoonsd, At the periphery of thenucleus there are 6-12 nucleoli. The cyto-plasm is vacuo].ated (Fig. 5); the peripheralvacuoles contain oil, the oircumnuolear onesoontain vitellin, The oocyte membrane is a orn-layer zona radiata. Durinr the phase of in-tensive tra hoplasmatic_2utb (Fig. 6 theoocyte diameter measures from 320 to 550,u.The oocyte is round or slightly ovol, Thelong nucleus is in the middle, i.ts margia ioheavily featooned. The nu«i ' diameter isabout 100,&. Thera are fom to 12 nuolsoliat the periphery st the nucleus (in a section).The cytoplasm is liosv5ly vacuolated and thevacuoles contain more oil (Fig0 7). Granulesof vitellin, previously in the vacuoles, nowappear in the cytoplasm. The oocyte membraneis thick and consists of two layers; zonaradiata externa and zona radiata interna.. Thefirst is more oloarly striated (Figo 6). Bothmembranes 'orm a Uttlo bo3.1.o7 at -thu Zorma-tion
thu mioropy]o. Th&o thc phc'oo st' ripe-floun the oocyte 0.r thu UI) to Soc' orno t roachocits definitive oio (chaut 800-. 900,u). Thenuoleer cliamotou i about 120/u0 The muoiono.a oval, its margin c feo tooed in section,J.t the boginniri of thu phase there erenucleoli et -the nuclear oripbery but then
Fig. 5 Smelt oocytes during the phaseof initial accumulation ofvitollin and oil.Manif. 10 x 8. Stain; Mallory
.4'
FRm/S 78 Sine it 1 9
they disappear. Vitollin granules and oilglobules of various sizes fill in the oocyte.A very thin layer of fine-grained cytoplasm ad-joins the oocyte membrane from within. Theoocyte membrane consists of two layers of zonaradiata; the thickness of zona radiata externais about 1103,t19 the thickness of zona radiatainterna is about 13.5/n. The micropyle isdefinitively formed (Fig0 8). The animal and.the vegetative poles of the egg aro defined..The nucleus moves from the middle of the eggtowards the mioropyle. The layer of f me-grained cytoplasm is thickest at the mioropyle
: r
Fig. 6 Oocytes in the phase ofintensive trophoplasmaticgrowth. Magnif. 10 x 40.Stain: Mallory
-
. .. .....
(Fig. 8). Vitellin and oil form a homogeneousmass at thin vegetative pole. After ovulationeggs come into the body cavity and are t}enshed. Generally growth and developmont ofoocytes in the White Sea smelt last 3-4 yinars,the period cf protoplasmatie growth being thelongest.
1.33 Portein spooifioity
Studies on this question are not yet pub-lished, but Rupp and. collaborators are workingon it (Rupp and Reclmorvi, 1966).
Fig, 7 Oocytes in the phase ofintensive trophoplasmaticgrowth. Magnif. 10 x 8.Stain: Sudan-Ill
Fig. 8 Part of an oocyte in thephase of ripeness. Magnif,10 x 20, Stain: Mallory
FRm/S78 Smelt
2. DISTRIBUTION
2,1 Total area
Anadromous and landlocked fishes in thePacific, Arctic and Atlantic oceans and theirbaine (Fig. 9), According to McAllister(1963), "the simplest explanation of the un-usual distribution of the two subspecies(particularly the Arctic hiatus of O.eperlanuamordax) involves an origin of the species inthe North Pacific or adjacent Arotio, Spreadthen took place westward along Arctic Russiato the White Sea. From the White Sea theywere either "sluiced-up" in proglacial lakesor dammed in front of advancing pleistoceneglaciers aoci carried into the Baltio, Or theymay have migrated around Scandinavia to theBaltic while isotherme wore depressed southwardduring glaciation. In either ease, followingdeglaoiation, the populations were left strand-.ed in lakes and cooler portions on the Balticto slowly undergo adaptation to higher tempera-turo and lowered salinities, the interveningpopulations along the outer Norvegian coastdisappearing. During the resultant isolationthe present !erlanus type evolved. Theorigin of eastern North American mordax fromwestern North American populations of the sub-.species can be ascribed to a migration acrossArctio Canada during a more recent interglacialperiod or during the warm postglacial hypsi-thermal period. Cooling them resulted in thepresent separation of populations. If themigration took place during an interglacialperiod, one may assume that intermixing tookplaco during the hypsithermal thus preventingdifferentiation of east and west populations.Thus the two subspecies, one with disjunct dis-tribution, can be readily derived, Range ofmorciax South to Barkley Sound, Vancouver Island,British Columbia, in the east Pacific; mouth toWonsan, Korea, in the West Pacific; in the Arc--tic east to Cape Bathux'st, Northwest Territories,west to the White Sea, The northernmost pointis on the southern island of Novaya Zemlya. Inthe Western Atlantic it is known from Pike RunCovo, Lake Melville, Labrador south to DelawareRiver, Pensylvania, and ueetionably to Virginia.Range of eperlanus: the Baltic and. North Seasand their drainages, southward, inoludingEngland to the mouth of the Loire River, France;upper Volga system" (McAllister, 1963), Duringthe last 10-15 years small smelt have spreaddown the Volga River system to the Gorkov andKuybishev waterbodios, (Kuznotzov, 1951;Kojevnikov, 1958).
Over its large range the smelt lives undervarious ecological conditionst salinity variesfrom oceanic to freshwater, temperatures varyfrom below 0°C (in winter) to + 20°C and higher(in summer). Land-locked forms of the smeltinhabit pure lakes, some over 30 ni in depth,others shallow, with running water (Arnold,
1920), with high oxygen saturation, high pII(1.8-8,6 and more), high degree of mineraliza-tion and Jew oxidability (Petrov, 1940) andrich development of zooplankton (about 300-1300 mg/rn3), Kojevnikov (1955) found that theFinnish Gulf smelt migrates and spawns in riversand estuaries at water temperatures from +2°Cto 12-13°C, pH from 7,0 to 7.5, oxidabilityfrom 8,94 to 15.13 mg/i, concentrations of bi-carbonates from 19.23 to 27,28 mg/i, oxygenabout 100 percent saturated, In summer thesmelt stays in open waters of the Finnish Gulf,below the epilimnion layer at temperaturesabout 5-7°C. In autumn the smelt stays atdepth of at least 20 in, at water temperaturesof 5-8°C, pH of 7,0-7,4, salinity of 34 0/00.In winter the smelt stays at depths of 20-30 ni,temperatures of + 20 and salinity of 5-6 0/00.Generally the Finnish Gulf smelt spawns attemperatures from 5-6°C to 10-12°C, feeds at5-10°C, spends the winter at +2°C. The WhiteSea smelt lives under more severo climatic con-ditions, The Sea covers with ice during 6-7months of the year, the water temperature dropsbelow zero. As the ice breaks the sea beginsto warm and in summer the temperature of thesurface layer rises to 16-18°C (and higher),The average salinity of the surface layers ofthe White Sea is 25-26 o/Oo, of deeper layers30-31 o/oo; it ini lower near the river mouths(riega, North Dvina and others). Subarcticpopulations of smelt inhabit waters with aver ezooplankton biomass of 50-400 mg/rn3 and avorebiomass of bentho of 4-30 g/m2 (excludingMollusca and Echinodermata which the smelt doesnot feed on),
2.2 Differential distribution
2,21 Spawn, larvae and juveniles
The smelt spawns in rivers or near theirmouths, Eggs are deniersal, adhering to thesubstratum. When hatched, smelt larvae driftdownstream where they aro carried back and forthunder th influence cf the tide, Fingerlingsstay offshore, feeding in pelagio waters in thesame regions as adults,,
2,22 Adults
The smelt stays near the shores for mootof the year. It oom.s to the deeper waters ofseas (or lakes) only in mummer when shore watersbecome too warm Smolt of all es spend thewinter in estuaries and mature fishes migrateupstream in spring.
2.3 Determinants of distribution oha. es
o 2,1, 3.16, 3.32
21
2*2 FRm S 3 Smelt
24 Hybridization
2.41 Hybrida; frequency of hybridi-zation; species with which hy-bridization occurs; methods ofhybridization
Interspecific hybridization is mot known.Lillelund (1961) writes that experimonts on
o loon
fertilization and breeding of eggs from fastgrowing coastal smelt and slow growing smeltfrom fresh waters (inland, lakes) have shownthat it is possible to hybridize these two mor-phological races, which have been isolated sincethe Yoldia geological period. Evidently thedata relate to migrant and landlocked smelt ofaerma).
PPROxIUA1I OcALE
2000 3000 MILES200 3O0 41oo KIL0M00005
Fig. 9 Distribution of Osmorus eperlanus mordax-oirole,and O, e. eperlanustrianglea.
FRm S78 Smelt 3:1
3 BIONOMICS AND LIFE HISTORY
31 Reproduction
3.11 Sexuality
The smelt is normally heterosexual.
According to Hoffmeister (1939) about 3.7peroent of adult individuals of the Elbe riversmelt are hermaphrodites (these individualshave male and female elements in the same go-nads). According to Lillelund (1961), thefigure for the same river is about 2 percent.These individuals do not lose the ability toreproduce. Hermaphrodite individuals occurmore often among young smelt: Lillelund (1961)reports that 14 percent of the O-group smelt ofthe Elbe river are hermaphrodites. Probablysome of the hermaphrodites become males in thefuture, Hormaphroditism occurs more often inmigrant smelt populations than in landlockedones of West Europe (Hoffmeieter, 1939). Herma-
phrodites occur very rarely in the smelt popula-tions of eastern Europe.
Sexual dimorphism is described by McA11I(1963) who states, "paired and anal fins slight-iy larger in males, large tuberoles on scales;small ones on head and leading rays of dorsal,anal, caudal and paired fins in males. Tuber-oies reduced in females, There is a tendencyin the males for a lateral muscular ridge toform, to be darker and to have a punctate oper-oulum",
3q12 Maturity
The age at, which sexual maturity im rea..ddiffers widely in different populations (TableVIII), Most cmelt populations reach maturityat 2-3 years. The age of maturity graduallydecreases on advancing from the northern partof East Europe eastwards to East Siberian shoram.Some workers noto that males reach maturity ayear earlier than females, but others do notnote such a difference.
Both sea and freshwaer smelt populationsare often biologically heterogeneous; thereare ecological forms differing in size and ratoof growth, age and size at maturity, period andplaco of spawning and other peculiarities.This holds true for the smelt populations ofthe Baltic Sea baya (Marre, 1931* Kojevnikov,1955), lakes of East Europe (Arkhiptzova, 1956;Stefanovskaya, 1957 and many others), and NorthAmerican populations (Kendall, 1927 and others).The smaller and earlier maturing form is alwaysnon-migratory, the larger and later maturingone usually migrates for more or leas oonsxder-.able distances from feeding to spawning grounds.Existence of those ecological forms promotesthe more complete use of food supply and spawn-ing grounds.
Intensive growth during the first year oflife is oorrelated with earlier maturity (lakepopulations of Europe and North America; popu-lations which grow more slowly reach maturitylater (Table VIII)0 Fast growing year-classesreach maturity earlier than those growing moreslowly in the same population (Morosova, 1960;Lillelund, 1961; original data on White Seasmelt).
Size and weight at sexual maturitySee Table IX.
3.13 Mating
Each female usually spawns in the companyof one male. The female extrudes her eggs,then leaves the spawning ground. According toHoover (1936) and Lillolund. (1961) an individus].female may continue to spawn for several days,but the process Is completed in several hoursin White Sea smelt (original data). Malescontinue spawning with other females. Malesspawn for a longer time and stay at the spawn-ing grounds longer than females.
3.14 Fertilization
Fertilization is external.
3,15 Gonads
According to Kendall (1927) "... the go-nade of the smelt are unsymmetrical organs, oneon each side of the abdominal cavity. The or-gan of the left sido is very much larger thanthat of the right aM. they are situated one be-hind the other. The left gonad is much largerin both sexes, the right being quite small andnot far behind the outlet".
The correlation coefficiente for differentyears between egg number and body weight in theWhite Sea smelt were +0.89 and. +0,95, betwoenegg number and body length 0,85 and *0.91(Beiyanina, 1966a). The egg number produoedby a female inoreasom with age; as a rule old-er fishes produce more eggs than younger f ishesof the same mizo (Pable X). Data on fecundityof various smelt populations according to sisoand ago are given in Table XI and Table XII,Land-locked freshwater forms and Siberian popu-lat ions have the lowest fecundity; nigrantforms (White Sea, ltio, Elbe river) have thehighest fecundity.
Variability of relative weight of ripeovaries (to body voiglit) in different smeltpopulations is rather smalis fron 18 to 22 per-cent (Table XIII). Weight of ripe teti is
about 5 percent.
According to Lillelund (1961) for theElbe river smelts
y 2.737x - 0.28637where "y" fecundity, 'z" body length
VL5
CLZ
QL
kcLU
t
iat
ebL
cir
Iicn
Enf
cza
rc.d
3O
nzL
cPy
c.ar
Q c
t-cl
itL
blla
vc R
ivar
Tic
hîga
L
irat
ch. R
irez
Baa
inSe
asC
zga.
3zy
akh
Len
e. R
ivnr
7,2
4o I O
5c9
1. t
& 2
9'î 12,9
8.2 10.7
8,5 11,0
87 9,4
to,'
ta,
93 1
,B 1
2,7
15.6
20,4
tO. 6
r'3
I 36
21,2
4.7
10.1
54.5
19.
241
903
53.
2 16
094.7 12.9 18,8 22.6
8,1 12.5 15.2
18,3
98 14.1 17,6
1100
t 2.2
55,8
23 8
tTe 6
6.3
17,6
22,0
b'V
Et
VC
flZ
Ç!E
22..4
4-6
1-3
2
3-4
2 2 22-3
2-4
3-4
3-4
4-5
4_5
56
gr o up
6__J
8
5t3 51.8 123
12.9 t6 14,4
18,3
I
o5 li'o
c'e ot'
turatioz
feo
tl1populatione
59.4
24,0
23.2
1901
25,0
17,7
5903
203
27.4
21,5
27.0
20,5
20,2
223
24.6
22.2
20,9
23,9
19.6
27.4
22.2
244
23.4
28.1
25.5
26.3 27.6 28.8 30.1
OU
Din
spa
nin
s10ok
iuthoL'ity
1-2
Fod.orova, 1953
l-3
Domraoh
and.
Pravdin, 1926
1.-3
Fed.orova, 1953
1-3
Lap
in,
1955-56
1.-3
Ui].lor, 1926
2.-5
Will
or,
1926
1-2
Marre, 1931
2-4
tar,
1931
2-8
Ste
f an
ov&
zeja
, 1957
4-12
Iely
ante
v,1946
2-6
Ark
hipt
zeva
,1956
2-5
Li].].elund, 1961
3-9
Koj
evni
kov,
1956
2-3
Creaser, 1929
2-4
Balthiin, 1948
2-5
IleK
eazi
e,1958,19(4
2-5
Rup
p,1959
2-7
Bal
agur
ova,
1957
3-9
Kirpiohnikov,1 935
3-5
Original
4.-7
Kirpichnikov, 1935
4-8
tiB]a
v8ky
1959
5-9
Tyurin, 1924;
Neiman, 1957
7-12
Piro
jnik
or,
1950
3EI:
E B
aycz
1f o
Ob
3o5
Tht
IiC
7 fl
iVC
'4,
6
56, 5 90'4
so. 6
10.7
T7.
313
.6 S
55.5
15.6
17,2
8,0
9.5
Tot
13.
47o
8 Sl
ot9.
2 15
.7t 3.7
130 T
12.2
6.3
10,5
133
3a
a
jJ w e 4
1Rm 578 Sm
Locality Length Weight
(cmi
Rybinak waterboclyWhite Lakelimen LakeLazmiaden LakeKurishes HaffLadoga LakeOnega LakeEibe RiverWhite SeaOb RiverYenimey HiverLena RiverUpper Lake
Length(orn)
Size anc woigiit at maturity Çfirs spawseason) in lifforont smelt popu1tion
TABLE IX
6.0-9,0 1.9-6,26,0-8.5 1.5-3.54,7-8.2 1.1-4.39.3-11.8 5.1-8.5
6,3 1,48,o-.11.o 3.3-8,78,8-10,6 3.7-6,116,5-18,2 abt.1718,8-22,6 41.7-75.518,3-19,3 42.5-48,520.3-22.3 51.2-68.019.6-23,4 53.0-94,014.0-16,0 17,0-25,5
Number of eggs per i g of body weight ofone female is given in Table XIII. Siberiansmelt populations have the lowest value ofthis character, the Pekov lake smelt have thehighest.
The smelt spawns once a year, so the numberof eggs produced by a female during a year maybe found in Table XII. Small smelt (Pakov-lake, limen lake oto.) spawn once only (Lapins1960; Kuznetsov, 1964). Migrant sea popula-tions have a longer life oyole; most White Seasmelt epawn once or twice during their livesanO, morne spawn three times (Belyanina, 1966a),Siberian smelt apparently may spawn more often,but there is sorno evidence that ouch fish donot spawn every year (Kravohuk, 1958). Thenumber of eggs produced by an individual duringa lifetime may be calculated from Table XII.Belyanina (1966b) reported that White Sea smeltcontained less fat and had lighter gonads, lower
TABLE X
Fecundity (thousands of eggs) of females of the samesizes but different ages (White Sea smelt, original data)
ig
Lapin, 1955-56Schotinina, 1954Domrachev & Pravdin, 1926W:Iller, 1926Marre, 1931Arkhiptzeva, 1956Stefanovskaya, 1957 and otheLillelund, 1961OriginalAmetislavaky, 1959Tyurin, 1924 and othersPirojni.kov, 1950Bayley, 1964
Age roupc3 4 5
Average min.-max, Averado min,-max, Aver e mth.-max.
Authority
3g 3
fecundity and smaller eggs after the heavywinter of 1960-61 than after the good winter of1961-.-62 (Table xlv).
According to LilleluncJ. (1961) the yield ofdifferent year-classes in the Elbe river showsa positive oorrelation with the water level andis affeotect by temperature and food conditionsduring the four weeks after hatching. Someworkers note that the yield tends to increasein oolder years.
3.16 Spawning
The smelt spawns onoe a year am a rule butKravohuk (1958) noted that sorne Siberi9n smeltdo not spawn every year.
Sometimes smelt spawn under ice. In mostareas spawning oontinues for about one month,but peak spawning usually lasts only ?-4 ds.yo;
19,0-19,9
20.0-20.921.0-21,9a4.o-24. 9
37.653.052.8
25.6-44,545.5-61,0
-
42,053,262,188,6
37.1-50.0
40.7-66.343.9-83,887.7-90,0 95,5 88,8-102,3
TA.BI
XI
Peoundity of different smelt populationn (thoueawis of eggs)
Data grouped by
h groups
Looality
Length groupe(on)
1011
1213
141
161
181'
2021
2223
262
28Authority
Pskocko-.
Onud.r31oyo
1.U
chin
okoy
cvaterbody
White lake
Rybinok
iaorbody
Kurishoo
Iaff (-31
Pyoo sere
Onega Lake
Ladoga Lake
Kuriahea
HaIT (sea-.
migrant)
Neya River
Finnleh Ou
Elbe River
White Seas
Kandalak-
elia Bay
Onega Bay
Gulf of Ob
Te nia ey
River
2.1
1,7
1.4
2,8
2.6
1,7
3,2
2,6
2,9
2,9
7,7
4,6
5.5
3,7
4,5
5.3
5,5
5,6
5.3
6,4
2,9
8,6
7,5
7.8
10.9 9.8
6.5
13.5 5.8
9,5
11.2
12,2
12,7 8.0
9.1
6.4
11,8
16.1
13.6
16.7
12,0
12.7
15,1
19,0
17.0
16,1
19,5
14,5
16,9
20,1
22.4
19.7
16,1
23,6
18,0
18.6
24,8
24.4
25.4
31.9
22.5
30.1
25.1
27,9
26,1
30.8
33.6
28.0
32,9
32,9
15.5
45,6
31.3
31.3
35,0
41.9
35.0
35,8
16,3
48,5
34.5
36.7
33,9
38.0
49,6
35.9
22,8
44.5
41.5
45.0
61.4
49.5
25.0
61.2
41.9
45,2
72,0
54.5
45.5
80,5
55.0
37.4
83.5
65.4
48,7
68.5
114
47.3
54.1
93.5
124
137
140
161
L.p
in'o
unp
ub..
ihe.
dat
cS
pann
oay
Grygorah,
19G
3Fedorova,
1953
Sche t mina,
1954
Marre,
1931
Melyantsev,
1946
Alezandrova,
1963
Markun,
1926
;Arkhiptzeva,
1956
Marre,
1931
Kojevnikov,
1949
Lapin'a unpub.-
lished data
Lilie lund,
1361
Ori
gina
l dat
aafter Balagu-
raya,
1957
Amatimlavaky,
1959
Tyurin,
1924
;Neiman,
1957
Locality
TABLE XII
Fecundity of different smelt populations
(data grouped by age-groups and given in thousands of eggs
Ago groups
23
45
67
89
10
ii
12
Authority
Lapin's data
Fedorova, 1953
Schetinina, 1954
Melyantzev, 1946
Alexandrova, 1963
1iare, 1931
Lapin's data
Lillelund, 1961
Original data
Saiagurova,
1957
Amatislaveky, 1959
Tyu
rin,
192
4,Neimarx,1957
PskovskoChudskoye L
e
White Lake
Rybinek waterbody
Pyaozero (Karelia)
Omega Lake
Kurishea Haf f
Finnish Gulf
Elbe River
White Seas
Kandalakaha Bay
Oriega Bay
Gulf of Ob
Yenisey River
206
1.7
2,6
800
2,7
3.2
5,7
9.6
6,7
21,0
10,6
-
4,6
1306 2,9
19.9
1304
3705
35.6
22,9
12.5
2.9
3.7
33.1
24,7
69,6
39.5
15,9
505
41.9
25,0
126,1
60.1
20,9
5.9
8.6
133.6
25,5 -
6,3
3105
3704
18,0
1740
0
48.7
54.1
45.6
48.5
3f FRm/878 Smelt
TABLE XIII
Egg number per g of body weight, size of ripe eggs and relative weightof gonads (as % of body weight) th different. smelt populations
TABLE XIV
Comparison of fecundity of females of the sameweight in 1961 and 1962 (White Sea, original data)
LocalityEg numberp'r g ofhor3 woihi
Averagediameter
of eg mm
Relativegonadweiht
Authority
Pekoveko-Chudekoyc La'' 1050 0.55-0.75 18.4 Lapin's dataKuznotzov, 1964
White Lake 580 0.85-0,90 - Fedorova, 1953Schetinina, 1954
Hybinek waterhody 710 0.10-0.90 20,7 Lapin's dataSohetinina, 1954
Pyaozero ((ars1ia) 560 0,79 13,5 Melyantzev, 1946
Ladoga lake 610 0.80 18.2 Markun, 1926Arkhiptzeva, 1956
Kurishes Haff 665 0,82 - Marre, 1931
Finnish (iulf 630-700 0,75-0.95 18.1 Lapin's dataKojavnikov, 1949
Elbe River 650 0.70-0,90 18-19 Lillelund, 1961
White Sea:Kandalakeha Bay 700 0.86-0.94 22 Original dataOnega 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
CharactersWeight
30
without internal or(g)
4 -60- - 10
Numberof
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 27.3Average diameter of an egg
(mm)0.83 0.79 0,87 0,87 0.85
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 204Average weight of gonacls(g) 11.2 15.7 21.9 25,1 33.4Average diameter of an egg
(mm)
0,91 0,92 0,92 0,92 0,93
Fatness (% of wet weight) 2,9-3,0
FRm/S78Snielt 3:7
Area
Spawning seasons of smelt in different areas
West EuropeBaltio and drainageUpper Volga systemWhite Sea, Karelia
Siberia, Far East
Atlantic oats ofNorth America
TABlE XV
Spawning season
MarchApril - MayApril - MayMay, early June
Juno, early July
From February(the south bound-ary of the area)to the beginningof June (the Mir-amichi River tn1northwards)
according to McKenzie (1964) in the Mirarnichiriver it lasts 5-10 days. There may be severalspawning peaks, depending on weather conditionsand population heterogeneity (subpopulations).
As a rule smelt spawn at night (Domrachevand Pravdin, 1926; Kendall, 1927, Hoover, 1936;Liovenso, 1954, Rupp, 1959, McKenmie 1964, andoriginal data).
The majority of workers have noted a de-crease in the size of smelt during the spawaingrun: the older and larger individuals spawnfirst, Changos in the sex-ratio on th spawn-ing grounds aro discussed below.
According to Rupp (1959) the character ofthe spawning run depends on ice conditions:the spawning run of smelt in Maine, U.S.A. (in80 percent of casos) begins within the first 10days after the ice has broken. The spawningrun breaks off when the weather is stormy orthe moon is very bright. Colour of water doesnot influence the spawning run.
All over its great range(see Table xv) thesmelt begins to spawn when the water temperatunis about 400 (sometimes 120 lower or higher).The spawning peak oocurs at water temperaturesof 6.-9°C (Chumayevmkaya-Svetovidova, 1945;Marcotte, 1946; Marcotte and Tremblay, l943;Dryagin, 1949; Schetinina, 1954; Ivanova,
1955; Stofanovekaja, 1957; McKenzie, 1964,etc.). Spawning breaks off at a sharp de-crease in water temperature. It lasts longerin cold years than in 'warm ones, The spawn-ing smelt usually avoids temperatures lowerthan 4°C and higher than 12°C. Spawning
Lillelund, 1961Probatov, 1927 and otherFedorova, 1953 and othersNelyantzev, 1946; Balagurova,1957, and many others
Agapov, 1941; Amstislavmky,1963; Ivanova, 1955;Kuznetzova, 1962MoKenie, 1964
begins earlier in warm years with high waterlevel, The smelt enters rivers and spawns athigh tide. Sometimes the spawning run is de-layed because of ioe movements,
Esox lucius, Leuciscus idus, Rutilusrutilue,, Misgurnus fosailia, Acerina oernuaspawn elmost at the amo time as smolt infresh waters of eastern Europe (Meshkor andSorokin, 1952).
As a rulo smelt ascend rivers to spawn(see 3.51), but some freshwater smelt popula-tions spawn near river mouths and do not as-oend. the rivers (Melyantzev, 1946; Fedorova,1953; Sohetinina, 1954, etc.). Many authorsnote that water depths at smelt spawninggrounds vary from several centimetres to soy-oral metres, According to Stofanowskaya(1957) smelt eggs occur at the depths of 17 min some Karelian lakes.. The eggs are deposit-ed on stones, pbblos, water plants, submergedparts of bushes, grass and other things, Theydo not occur on muddy bottoms, The currentvelocity in the spawning grounds of the WhiteSea smelt is between 0.3 and 2 rn/sec (originaldata),
Spawning areas are larger in the yearswith high water level. Rupp and Redmond(1966) noted that smelt taken from spawninggrounds in rivers and introduced into somelakes of Maine did not enter other rivers butspawned near the banks of the lakes,
A number of authors, including Kendall(1927), Baldwin (1948), Lillelund (1961),Amstislavsky (1963), and unpublished data re-
FRm/578 Smelt
over according to Unanyan and Soin (1963) onlydead egga of the White Sea emelt become de-tached from the aubetratum
The vitellum io round or a little oval inchape. A thin layer of cytoplasm ourrouncioit. The vitelline membrane le thin, and flume-roue oil 1obules are dieperood all over thevitellus (see 1.32).
32 Pre-adult phae32I Embryonic phase
The embryonic phase is clesorihed by Unanyanand Soin (1963). After extrusion into water,the egg immediately swells. Its diameter in-creases about 0.2 mm, perivitelline apace formabetween the vitellus and the membranes. In 4 h(temperature of 805°C) the blastodimo forme atthe animai isole of the egg (Fig. 10.1). In 8 hafter fertilization 2 blaFitomeres are present(Fig. 10.2) in 13 h 30 mm, 8 blastomeree arepresent (Fig. 10.3), In 2,5 days after fert-ilization (12°c) the biastula has formed (Fig.10.6). Formation of the embryonic plates be-gins 2 days 14 h after fertilization (Fig. 10.7).In 3 days gastrul atio is complote (Fig. 10.8).In 4 days 5 h (10.5°c) the embryo has eyecups(Fig. 11.3 4); in days 5 h after fertiliza-tion (io°c5 segmentation of the mesodermn begins,the main parte of the brain form, auditory veo-toles and c:rystallins lonsea in the eyes arepresent. The embryo has a ohorda, 30-32 rnyo-tornee, and the caudal part begins to separatefrom the yolk sac (Fig. 11.5,6,7). In 7 days10 h the embryo ham a long tail separated fromthe yolk sac, the eyes have developed blackpigment; the embryo showm twitohing movementowithin the egg (Fig, 11.8). In 9 days and 16hafter fertilization (9.5°C) hatching glands arepresent all over the body surface, particularlyon the head and i.eo±oral fins; the parts ofthe brain are well formed; t}1e black pigmentin the eyes become stronger and yellow pigmentappears. The yolk sac is egg-shaped. Theembryo han olfactory vesicles; oemiolocularcanales are preoent in th auditory vesiclem.The head is not distinct from the yolk sac.The mouth i at the lower side of the head.Four rudimentary gill slits and, a rudimontaryoperculurn are present. Melonophores are foundon the ventral aide of the caudal myotomos,under the ai isentary canal, and on the yolk sac.The heart pulsates regularly; it consists oftwo ohamhe:ra. The blood has no corpuscles.There are 42 pre-anal and. 17 poet-anal myotomes(Fig. 12_i)
Thring embryonic development there are nodifferences between migrant and small freshwatersmelt except the body size and number of myo-tornee (Orib, 1946).
3:9
Egg predators includo insects and fishes(etickiehack, smelt itself and, others). Eggsmay become parasitized by Saprolegnia.
Nany authors (including Schneherger, 1937;Neehknv and Sorokin, 1952; Unanyan and Soin,1963) have remarked on the high rate of survi-val of smelt eggs during incubation. Accord-ing to LiJlelund, (1961) the rate of mortalityinoreasos only at the end of the incubationperiod, when about 86 percent of eggs of theElbe river smelt die, However Rothschild(1961) ha oLated that lose of smelt eggs inthe Dean-Brook river, Maine, U.S.A., is 99.5percent. The main causes are drying after t}water ìvel ham dropped, overwarmning and pre-dation (see above). Influence of these fac-tors is least in cold. years with high waterlevel.
Rothschild (1961) and McKenzie (1964)found that the number of larvae produced perunit area morcases up to egg densities ofabout 11-13 per and declines when the den-sity of eggs is. higher. The relativo larvalproduction is highest (hatching rate 3.6%)whom the average density of eggs is 0.5 per cm2and declines to 0,03% at an egg density of 194PST cm2.
Before hatching, the embryo begins to movevery energetically, the caudal part moving es-pecially sharply. The membrane bursts and thetail comas out first, Moving energeticallythe embryo frees itself, the egg-case romaineon the substratum or is torn off during hatch-ing.
3.22 Larval phase
The period of incubation varios in differ-ent smelt populations (Table XVI). Generallysmall forme have a shorter period of incubationand the new:ly hatched larvae are smaller.Naturally the sum of day-degrees from fertili-zation until ha'tohing is not constant, but in-oreases exponentially with decreasing watertemperature.
A newly batched pro-larva has the yolk saoabout 0.7 mm in length. There is an oil glob-ule in it. The globule resolves after theyolk. Grib (1946) described 4 stages in thelarval development of the Nova river smelt:
1. Pro-larva with yolk-sac. This etagelaste 7-9 days. The dorsal fin fold begins atthe second segment. The pro-anal fin is welldeveloped. Pectoral fins aro present. Pig-mentation is on the ventral part only, Thebody is transparent. The blood has no cor-Spuedes. The mouth becomes terminal at theend of 'the stage.. The pro-larva begins feed-ing while the oil globule is still presont.'l'ho 'o. loi"v measures to 7 mm in lngth(c"ig. 12
Fie. 10 Early stages of development of eggs of theWhite Sea smelt (to the end of gastrulation)(from Unanyari and Soin, 1963).
Fig0 11 Late stages of development of eggsof the White Sea smelt (to hatching)(from Unanyan and Soin, 1963)
.FRJS78 Smelt
Locality
Pskov-lakeValday-lakeRybinmk-waterbodyElbe rivereva river
Thite SeaOb estuaryirasichi river
Larva without yolk-mao, 6.8-13.4 mrs.Blood oorpusolem are present. The fin foldis reducod, dorsal and anal rays are developing.Pigmentation is similar to that of the pro-larva; a pigment spot of two melanophores lepresent under the anas, The alimentary canalis tubular. A rudimentary gas-bladder can beseen at the end, of the stage. This stagelasts from 10-14 days after hatching (Fig. 13),
Larvae of 13-13 mm. The body is trame-.parent. Th dorsal, anal and adipose finshave formed, Bays are present in the unpairedfins, The caudal fin is rounded. Tho pro-anal fold begins from the eighth pre-anal nag-ment.
Larvae of more than 18 mrs (28 mm inaverage), Such. larvae appear in the Novariver in July-August. The caudal fin is in-dented, The final number of raye are presentin the dorsal arid anal fins. The embryonicfin fold can be distinguished on the rentra.].aide in fish of up to 30 mm. The ventral finsare situated more posteriorly than earlier as aresult of growth of the gas bladäier. Upperand lower vertebral arches are present. Theoperouluni covers the gills completely, Pig-ment is concentrated mainly on tho ventral side,Melanophores appear on the operculum, on thetop of snout, under the ayes, along the chorda,on the body sides and on the dorsal part cf thtail to the end of the stage, Larvae andadults have similar nieristic and plastic cha-racters. The number of myotomes does notchange during the larval period.
According to Raes (1949) in general thesmelt larvae belong to the "herring type" hav-ing a relative short caudal part, isometricpigmentation, low disposition of the pectoralfins, homocerca]. tail and so on.
TABLE XVI
Duration of period of incubation and body length athatohing in different smelt populations
Sum of Body length atday-degrees hatching (mm)
Authority
80 3,8 Mohkov and Sorokiri, 1952110 6,2 Chumayovskaya-Sve tovi.dova, 1945138 4.2-4,9 Sohetinina, 195460-110 5.0-6.0 Lilleluni, 1961
140-180 5.4-6,0 Grib, 1946aht.170 6,0 Unanyan and Soin, 1963
132 - Amatisiaveky, 1959174 5.0 Ì4oKenzie, 1964
Smelt oggo and larvae arc fed on by thethree-spine stickleback, gobiido, herring,perch and, other fishes (including the r"eltitself) and larvae of Inseota. One o hamain causes of egg mortality is SaJ:
Unanyan and Soin (1963) sta;;:d .t lr.aer" tho White Sea ornait begin food within6 days of hat chirig and aro on -. r l.- 4cpmndenion external feeding 9 days pf cAccording to Gri (1946) the '''L1rcrbegins to food 7-9 days after ) ch 'c Thecorresponding time for Pekov-laka "rich i" 7-8days (Meshkov and Sorokin, 1952),
Lilelunrl (1961) stated that a cro-larvawith yolk-sac may live without fc ri.::.r. 'or 10-14 days at a temperature of 10-12
Larval smelt feed ort email pinetonicforms: young stages of Copepcda hotatoria,larvae of Gastropoda and Lainoliibr:cichiata,
3.23 Adolescent phase
According to Grib (1946) the adoJeecenphase in the Neya river smelt bog r-j nSeptember when the first scaler ,'rr'e (c 4 cm),Ehrenbaum (1909) gave the eate dc for .hs E]beriver smelt. However McKemz'a (9') e'aectthat first scales in the Mircrr o r - - cr rmaitform at a length of 20-25 mrs (g 2 'c) Bythis time the pigmentation of 1r do'sal parbecomes moro intenso.
The main mortality occurs rhr rg the 'irr hmonths of life, Lillolund, (t3) otmc hrI,the mortality coefficient is ghoe frrrsMay-August, its value decrcrsmr ;r' flnoPec rcithen it is of little iwporteucc
3x12
3
o
7
1mm
o.- ,,ge,
Fig. 12 1. Embryo of White Sea smelt9 eggmembranes removed
2,3, Prolarvae of White Sea smelt (from Unanyan and. Soin, 1963)
1mm
-
Fig0 13 Larvae of White Sea smelt after resorbtion of yolk(from Unanyan and Soin, 1963)
PRm/S75 Smelt
The young smelt feeds on planktonic Orum-tacoans (copepoàa, Cladocera) and, other groupeof zooplunkton.
3,3 Adult phase (mature fish)
3.31 Longevity
Longevity varies greatly in various popula-tians of Osmerus eperlanus (Table VIII). Gen-erally small freshwater smelt do not eurvivfor more than 3 years. Individuals of 12 and,oven 15 years old may be found in the smeltpopulations of estuaries of the Yenisey, Lena,Anadyr and Khatanga rivers (Agapov, 1941;Mikhin, 1941; Lukyanohikov, 1964). The max-1mal age of malos is usually 2 years less thanthat of females.
3.32 Hardiness
The environment of smelt populations isdiscussed, in section 2,1.
According to LilloluncI (1961) experimentsmade in Hamburg harbour showed that eggs developin spite of sewage water. Only in the lastphase of development where there is very littlecurrent is there a great mortality of eggs,caused by the accumulation of detritus, proto-zoa and algae on the eggs. Eggs of smelt arefertilo up to a salinity of 16 o/oo. The sua-oeptihi.lity of eggs to salinity during theperiod of incubation depends on the stage ofembryological development. A salinity below10 0/00 15 neither injurious to eggs nor tolarv-ae, The upper limit of temperature toler-ance of early eggs lies in the range 17,7-20.7°C,for later stages it is 21-24°C. Comparabledata for adult smelt are not available.
3.33 Competitors
The main food competitors of Siberiansmelt are Coregonidae and Acerina cernua feed-ing on Chironomidas larvae and nektobenthicOruetacea (Pirojnikov, 1950, 1955; Amst1elavçy,1963, and others).
Young of the White Sea smelt compete forzooplankton with herring, adults compete forAmphipocla, Mysidacea and some Cumacea withherring in April-May and. September-October(Epstein, 1957). There is great similarityin diet (especially on the basis of Polychaeta)between the White Sea smelt and Eleginus navaga,yoxocephalus M. uadricornis (Russa-nova1 1963)Thasterosteus aculeatus Timakova,1957), Pleuronectos flesus and Liopsettaglacialim (Kudereky and Russanova, 1963).According to Abdel-Malek (1963) there is nofood competition between adults of smelt andthree-opined stickleback. In the early summerthe Finnish Gulf smelt competes for planktoniccruataceans with young and partly adult
Cyprinidae (Rutilus rutilus, Alburnus alburnus,Abranis brama, Leuciscus idus and others) andperch (Perca fluviatilis) (Kojevriikov, 1955).The young of the Elbe river smelt compete forfood with the young of Clupea finta from May toAugust (Lillelund, 1961). The fresh watersmelt of the north-western part of USSR occupiesthe food niche of a plankton-feeder in openwaters and competes for zooplankton with Core-
albula, Abramia ballerus, Alburnus albur-nus, young of Perca fluviatilis, sometimes withRutilus rutilue and Acerina cernua (Petrov,1940; Meehicov and Sorokin, 1952, and others).
3,34 Predators
Sea mammals, birds and fishes (cod inparticular) food on migrant spawning smelt insubarctic waters. Ezox lucius, Luciopercaluo1operca and large individuals of Perca f lu-viatilis feed on smelt in European fresh andbrackish waters, Salvelinus fontinalie,Cristiwomer nanaycush, gnue clupeaformis,Stizorstodion vitreum, Perca flavescens, Anguillarostrata, Ì4icropterus dolomleu, Norone americanaand. Lota ,yiaoulosa feed on smelt in North American
wato'"rKendall, 1927).
There is sorne evidence that the specificsmell of ornelt stock scares away other fishes.
3.35 Parasites, diseases, injuriesarid abnormalities
According to Petrow (1940) 28.5-36.6% ofthe Pokey-lake and 20% of the White laico smeltare parasitized. Parasitism io less intonsivin other lakes of West Europe. Shulman andShulman-Aihova (1953) list 16 species of para-sites for the White Sea smelt,
The list of smelt parasites recorded fromwaters of the USSR is given in Table XVII
3,4 Nutrition and growth
3.41 Peoding
Anstiolavoky and ussynina (1963) statedthat the Ob river smelt feeds during day andmight but with different intensity: maximumfeeding takes place at 13 and 21 h correspondingto vertical migration of zooplankton; minimumfeeding is at 5 h.
Migrant smelt feeds in coastal and offshorewaters of seas and in river estuaries; land-locked forms feed in open waters of lakes.
The smelt searches in the water taking foodorganisms selectively (Table xviii),
According to Timakova (i 957) the White Seasmelt feeds on fish (herring) and Crustaoeaduring the post-spawning period. Intensity of
3r13
Class orOrder
Miorosporidia
Cestoidoa
Species
Glugea herwigi*Weissenberg,1921
TriaenophoruBcrassus Forel,1880
Triaenophorus *nodulosus(Pallas, 1781)
Eubothriumcrassum Bio oh,1779
.h liobothrjumatum L.,175;
Diphyllobothriumnorveicum Vik,1957
Proteocephaluslongicoll is
1925)
Stage of para-site infeotirigsmelt
Adult
Larva (enoys-ted pleuro-cerco ida)
Adult
Larva (pleu-rocercoids)
Encys tedPleuro cerco ids
Adult
TABLE XVII
Parasites of smelt (for territory of USSR)!!
usoles, gills,
Liver andother organs
Intestine
Musolea, liverand otherorgans
Tissues ofintestine andstomach
Intet me
adult is aparasite ofp ike
Any Copepods,the finalhosts aro pike,perch andother predators
Pleurocercoidsin intestine operch, pro car-coids in anyCyclops
The final host-mammals
Final hostsmammals andbirds
JJ Table based on "A key for identification of parasites of freshwater fishes of USSR",edited by B. Byhhcveky (1962)
Baltic, White
All over USSR
Within the ararea of Sal-moflida
North-We sternUSSR, VolgaYenisey, ObRivers, basins
Not recorded
North-Wes ternUSSR, SiberiaKajnchatka,Ainur River
Not recorded
Far East,Kainchatka
Basins of theWhite, Barentz,Japan Seas
ÇLeder, 1800)
Trematoda Leicithaster sp. Adult Intestine
Orientophorus Adult Pylorio oaeoatrovi
(Layman, 1930)
Podocotyle Adult Intestinereflexa(Creplin,
me menteriesetc.
and BarenizSeas and theirbasins, lakesof the UpperVolga, Amur.
Larva (plouro- Muscles and Proceroolds Area of Esoxcercoids) internal in body oavity lucius and
organs of Copepoda, Esci reiohorti
3214 FRín/S78 Suieli
Class orOrder
Trematoda
Nematoda
Spe oie s
Buoejhalue01 mor hum
Baer, 1 27
Cotylurum rile-atus (Run,, 1802)
TotracotyleintermediaHughes, 1928
Diplostomum **31
spathaoeum(Rud., 1819)
Contracaeownaciuncum(Rud., 1802)
Porro cae ownep eri ani(Linstow, 1019)
Cystidicolfar io ni sFisher, 1798)
Camalianuslacustris(Zoega, 1776)
Camailanuçjtruncatus1iidT7il 4)PhilomotramanguineaRud,, 1917)
St of parai-site infectingsmelt
Adut
Cyets
Larva
Larva
Adult
Adult,larva
Adult,larva
Acilt,iorra
i rva
Adult,larva
Infected Other hostsorgans and stages
in them
Gill tissues,under skin
Tissues of Final hosts-gas bladder birdsand otherinternaiorgans, bodycavity
He art
Crystalline Final hosts-lens of oyes birds
St mach, ceso-phagus, pylo-rio oaeoaLiver, n.ry, muscles
Dorsal musclesunder skin
Gas bladder
Intestine
Intestine
Body cavity,the skin bot-ween fin yo
Adult unkno
Are a
Baltic and WhiteSeas basins,Siberian and FarEast Rivers
Baltic and WhiteSea, VolgaSiberian RiversRybinek waterbor,Kare lia
Not recorded
ot recorded
North-Wee ternUSSR, Ob and AmurRivers, Japanand Okhotsk Seas.basi mci
North-WesternUSSR
Baltic, White,Barentz, ringSeas basins
Evsryuhere, e-copia the AmurRiver basin
'ytihere, ex-the Amur
recorded
FRm/578 Sm1t 3x15
TABlE XVII (continued)
3:16
Copepoda
Banchiura
clavula (Bu-jardin, 1845)
Mete ohinorhyn-chue salmenis
TiiTier, 1780)
Ponrnhorhyuchuulaovie(MUller)
Co ryno s orna.
semermo(Forsoell, 1904)
Coryn000mU t3.'IUUO CUIR
(Ruth, ThO2)
iraoiJio.ant Markouitsh,932
Ergasilis sio*boldi Nordmann,1832
Gussov 1951
Ax' lus :í:o *
cous L.,1(56)
Adult
Adult
Larva
Larva
Adult
Adult
Adult
.du1t
* May causo mass mortality of fishCauses loss of weight
** Couses blincthess and mortality
TABLE XVII (continued)
Intestine
Intestino,livor
Body cavity
Body o'iy,loulicles, lu-
terial orUano
GiLls
Gills
Gills, skin
Gills andskin
Other hostsand stagesin then
Not shown
Intermediateho s ts-Gamînarus
pulox and.Fontoporeiaaf f luis
Intermediatehost-Ponto-porela affinis
Gammars pule
Final hostssea marinaisrarely birds
Final honte -esa luamnalerarely brclo
Are a
Not recorded
Everywhere
North-WesternUSSR, Volga
North and EastSeas of USSR
North and EastSe of USSR
ar East
FRrn/78 Smelt
2a1tio, Siberia,Amur River
Baltic, Japan SeaSiberia
Europe, Siberia
Class or Stage of para- InfectedOrder Spe oies sito infecting
smeltorgans
Acantocephala Echinorhynohue Adult Intestineadi Mil1or,
i 77
Pseudoo chi- Adult lutes tine
chua
FRm/S78 Smelt
Components
CopepodaCiado ceraMys idao
AmphipodaIsopodaCumacea
51.232,9
feeding increases until July. Polychaeta (45%)and herring eggs (27%) aro the main food cornpo-nents. In summer older individuals feed onPolychaeta, Decapoda and fishes, in autumn theyfeed on Polychaeta and Amphipoda. Intensity offeeding decreases until December. lu winter itis low, and food consists of nektobenthic Crus-tacea, Amstislavsky and Brussynina (1963)noted that the Ob river smelt feeds on Amphipodain winter. Thiring the vegetative season Mysi-dacea, Copepoda, Cladocera and young smelt areimportant in smelt diet. The Lena river smelthas the same diet (Pirojnikov, 1950, 1955), TheElbe river smelt foods on Eurytemora from Aprilto October and Gammaridae from November to March(Ladiges, 1935). Planktonic Crustacea dominatoin the diet of small freshwater smelt. Theirimportance in the diet depends on seasonalabundance.
According to Tiniakova (1957) intensity offeeding of ritales and females of the same agevaries with the seamon after spawning femalesfeed much more intensively than males; in Julyand August maies feed more intensively,Females oonsumo more fish than males.
The assortment of food components, theirsizes and quantity change with growth. Youngof all populations feed on zoopiankton. Fishesaged. 2 years eat Nysidas, Amphipoda, Inseotalarvae and fishes, The importance of fishesincreases with :e.
In winter the smelt feeds very little.As a rule smelt abstain from feeding during thespawning and pro-spawning period (Nikolsky,
1956, and others).
3.42 Food
Young and adults of small freshwater smeltfeed on planktonio CruBtaoea: Cladocera(Bosmina sp., Daphnia sp. etc.) and Copepoda
TABLE XVIII
Selection of food organisms by the Lena riversmelt (after Firojnikov, 1955).
Fishes were 6-11 years old and measured 161-317 mm
share (in %) Share in thein plankton in nekton diet of smelt
24.211,9
1 .7700
Oo
67.311,4
oo
3:17
(Cyclops 5go, Diaptomus sp. etc.). The maimfood groups of adults of different populationsare Nysidacea, Amphipoda and young fish. Thesmelt of the Atlantic coasts of North Americafeeds on pelagic Isopoda (Kendall, 1927). Thediet of the White Sea smelt consists of 60 com-ponents (Kudersky and Russanova, 1963),Polychaeta playing the main rolo (mainly Nereidawhich do not occur in smelt diet in other areas).
3043 Growth rato
According to :lyanina (1968) all smeltpopulations may be divided into 2 groups basedon character of the growth rate during ontogeiys(i) populations whose individuals grow mostintensively during their first year of life;then the growth rate decreases comparativelysharply (smelt populations of West and EastEuropean basins, oxoludin the extreme North,and North-American waters);(2) populations whose individuals grow com-paratively slowly during their first year whiletheir rate of growth increases more or lessstarply during their second year. Then itdecreases gradually (smelt populations of theWhite Sea, the Cheshskaya Bay and Siberiancoasts). Sizes of individualrj of differentsmelt populations in successive years of lifeare given in Table VIII. Valuen of the lineargrowth (in % to maximal length) of some popula-tions of both groups are shown in Fig. 14.
Condition factor (Ponderal index)
The well-known length-weight relation infish may be expressed by the following equatn
Q Al', where Q weight, i . length,
A const., n - near 3
Values of "n'e for different smelt populationsare given in Table XIX.
3:18
(50
IIQ
20
Values of ??I for different smelt populatione
Looality
1 2
1 2 4 5 (5
yesro ei. life
TABLE XIX
(50
L'o
20
A. Populations of the 1group
r;Q
40
20
B. Populations of the 2group
Fig, 14 Linear growth of fish in successivo years oflife (as % of maximal length) of differentsmelt populations (from Belyanlna, 1968).
n Authority
1 2 4 5 6
years of life
Yenisey River
FRmJ78 Smelt
Landlocked
Sea,.-migrant
White lakeRybinck waterbodyKuriahea HaffGreat lakee
Elbe riverF!tramioh! riverhito SecKi1uIciha BayDrina Buy
lonisoy rIver
2.70
2.912.872,82
3.283.20
3.483.363.15
Sohetinina, 1954Schetinina, 1954Marro, 1931Lillelund, 1961
Lilleiund, 1961McKenzie, 1958f Lillelund,
Original dataKirpiohnikov, 1935Tyurin, 1924 and otberm
1961
7 8
Ryhinek watorbody Finnish Gulf
Locality
The populations of the first group liveunder temperate climatic conditions0 Theirseason of growth lasts 6-7 months. They livein waters with a good supply of small zoopiank-ton whioh the fry and young feed. on teplankton biomasa is about 300-1300 mg/rn , asmay be concluded from various literaturesources0 The rate of growth of freshwaterlandJocked populations decreases sharply duringontogeny (Belyanina, 1968 Tablos VIII, IX,XIX, XX Fig0 14). PoDulations of the secondgroup live under severo sub-arctic conditions.Their season of groui,h lasts 5-6 months (WhiteSea smelt) or only iaonths (East Siberianpopulations)0 The average zooplankton biornassis about 50-300 mg/n3 dulng the season0 Thelatei' spawning period (May-June and even July)compared to that of the European and Americanpopulations (March-April), the lower tempera-ture during the season, the earlier cold Inautumn, the poorer food resources * all themefactors c1.00rmine the slow growth-rate of theyoung smelt of these regions0 However, rate
ThBLTI XX
Rate of growth in different smelt populationa.(from data in Table VIII)
3.22.51.32.50,7
1,5 1.52.5 loi 2,91,2 209 2.2 2.50,6 1,2 004 0.3 0,5 0.51.44.2 2.3 1.2 2.3 1.2
3,9 3.9 2.62.5 204 1,6
4.4 407 4,0 4,2309 3,7 2,2 2,4 3.1 2.8 0.75,9 3.8 2.4 2,0
4.4 2,7 2.5 208 1.74,3 3.5 2.7 200 106 0,5 1.1
3Q8 2.9 1.2 1.3305
Length increase in successive years cf life3 4 5 6 7 8 9 10 11 12
of growth increases during the second year oflife and greatly exceeds that of the first year(Fig. 14). This is evidently a result ofchanges in ooding habits; the smelt begins tofeed. on lrc C:custaoeans (Amphipoda, Mysiciacea).The Kandalakaha Day smelt (white Sea) is remark-able for iba most rapid growth (linear andweight) th'cin the socond (and. third.) year oflife (Tablao VUI, tX, xIx), This populationlives under conditions of higher salinity (25-.28 O/oo) than do ali other smelt pop'lations,The food resources of the White Sea are richer
}1r' those of estuarios of Siberian rivera; theaverage biomasa, of benthos of the White Sea isabout 10-30 g/rn2 (excluding Mollusoa and Eohino-derynete, which smelt does not feed on), whilethis valuo is only 4-5 g/m2 for the estuariosof the Siberian rivers (Oreas, 1957, and others),Also the Kandalaksha Bay smelt feeds intensivelyon heteronsreid. stages of Polychaeta, mainlyNerelu virons (Abdel-.Malek), 1963; Kudorsky andilusoonova, 1963) which is an eaiiy accessibleand very abundant food in this region.
2
Frochwaters
'(,2
5.25036.0
2.93.04.6205
Pskov-1 alce
limen lakeValday lakeWhite lakeRyb1nk wtb 5.9 2,8
Kurishee Haf f 6.3 4.2Darley lake 7.1 3.6Lazniiaclon lake 8.2 1.1Lad.oga lake 8.0 1,5Onega lake 6.3 2.5
Michigan 9.2 6.5Upper lake 6,5 8.3
Sea-migrant
Elbe river 7.1 6.3Finnish Gulf 7.8 3,3
White SeasOnega Bay 407 5,4Dvina Bay 4.1 5.2Kanciaiaksha Bay 4,7 8,2
Chesha Bay 3.5 4.6Yenisey river 4,5 5,3Lena riverAmur river 505 6.0
FRs/S73 Sm I 3 19
32O FRni373 niei&
The length of the growth season is differ-ent for different age groups young f leh beginto grow earlier and finish the season laterthan older individuals (Lillelund, 1961 and.others). This may be related to spawning inspring or in the beginning of summer.
Smelt grow faster in warm years and slowerin cold ones (Kirpiohnikov, 1935; Meshkov andSorokin, 1952, original data on the White Seasmelt). The effect of temperature on growthis seen more clearly in younger fish.
Generally strong year-classes grow moreslowly than weak ones (Lillelund, 1961;Morosova, 1960, and original data). Accordingto Abrosov and Agapov (1957) the rate of growthof the small smelt of the Jijitzkoye lake de-creases with increasing population density.
3.44 Metabolism
The metaboljm of smelt has not beenstudied.
Leviyova (1952), Kleimenov (1962) and morneothers give data on the chemical composition ofsmelt (Table XXI), Data on fat are shown inTable XXII, Siberian smelt have the most fatin the body and reproductive products. Thesmall smelt of the Upper Volga system have t'nelowest fat content. During the spawning periodthe fat content of the body and internal organsof males (except gonads) is higher than that offemales. Chechenkin (1952) reported that thefat of the Pakov lake smelt ccnsiets mainly ofnon-saturated fatty acids, with oleic aciddominant.
Fatness chances during life and by seasons(Belyanina, l966b). Localization of fat insmelt (in por cent to total fat) is shown inTable XXIII.
Ripening of the female gonads results inconsiderable changes in the localization of fatand its breakdown, The fat of the intestineand mosenteries is used up almost entirely.This process is much less intensive in males.Acoordíng to Belyanina and Ì4akaroira (1965) afemale smelt uses up about 65% of initial fatduring epawning, 53% being removed as fat ofextruded eggs and 12% being used in activity.A male uses up about 47% of its fat, fat inextruded reproductive products being 6-10%;a great part of the fat is used in activityduring spaming.
Sudocrine systems and hormones and osmoticrelat,ionc of the smelt have not been studied.
3.5 Behaviour
3.51 Migrations and local movements
During most of the year smelt stay nearshores. After spawning the fish migrate todeeper water, evidently to avoid high tempera-tures, In autumn fish of all age-groupe keeptogether. Smelt tend to form local popula-tions. Feeding migrations of smelt are short.They spend the winter at river mouths. Insprin, they migrate to spawning areas Upstream.Extent of spawning migrations is given inTable XXIV, season of migration in Table XV.After spawning adult smelt actively move down-river, prolarvae drift passively.
Differences in the migratory behaviour ofdifferent populations are mentioned in Section3.12,
3,52 Schooling
Evidently smelt form schools during allstages of the life cycle (Kendall, 1927 andothers). By our observations only the largestand oldest examples of White Sea smelt (6-8 yrold) do not form schools but search for fishalone. Spawning and wintering concentrationsare rather dense, and feeding schools are ofi :ms densìty. In autumn and winter, smeltconcentrations consist of fish of both sexesand different age groups. In spring, matureindividuals form pawning schools, grouping bysize or age.
Gohiids, flatfishes and other speciessometimes occur in fishing note together withthe White Sea smelt; herring, stickleback,sprat nd others with the Finnish Gulf smelt,
Larvae and young smelt feeding on planktonperform vertical movements (Amstislavsky, 1963;McKenzie, 1964).
The behaviour of spawning schools is dis-oumsed in Section 3.16.
3.53 Responses to stimuli
Kendall (1927), Baldwin (1948), Dryagin(1948) and others have noted that adult smeltavoid strong light. Strong light destroysdeveloping eggs. Hatching larvae reactpositively to light (Unanyan and 5oin 1963).
Unanyan and Soin (1963) noted that sea-water of 26 O/<, salinity has a harmful effecton the reproductive products of smelt, prevent-ing fertilization. Seawater of salinity above
FRrn/s Sri t
TABLE XXI
Chemical composition of smelt(from Levlyeva, 1952 and Kleimenov, 1962)
Contente in mg/bog
IC Ca Fe 1MG
388 83 07
Aminoaoid oontent as % of protein
There ae 2340 international units oi viosin D in i ; of omoltlivor fat and 170-200 i.u. in i g o:' :'r of uitomc'l ognnr
3:21
Isolouoino
4.8
Leuoine
.8
Lysine
11,1
Methionino
2.8
Phenilalanine
4.0
Treonino
4.3
Va].ino
5.4
Organ Month Chemical composition (%) CalorificvalueWater Fat Total
NProtein
NExtractive
NTotal protein
..N6.2Ash
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 2,30 85.2-89.7
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 45June 80.6 2.9 2,05 1,48 0,57 12,81 4.41
livers May 74.9 7.6June 79.8 3,9
internal May 59.7 27.3organs with-out reproduo-tive produots
June 78.3 8.3
ar East
e nt ire May 77-78 4 4- 16,1-17.0 1 O- 110.65.0 1.1
eggs 62 12,0 23,0 2.0
te st is 66 7,0 24,0 3.0
3:22 FRm/578 Smelt
Fat content of opaming smelt from different populations
Sex Organ
White Sea
Uchinskoye Females bodywaterbody gonads
and other internal organsRybinolz
waterbody Malos bodygonads
other internal organs
Females bodygonads
other internal organs
ales bodygonads
other internal organs
PABLE XXII
TABLE XXIII
Localization of fat in smelt (as % of total fat)
SeasonAjilMa31st0ctobe
44-53 59-6545-57 2-3to 4 33-38
78-88 63-796-12 4-6
to 16 20-33
727
21
Locality Fa conten in % of we subs tance Authoritybody eggs testis
White Sea 1.7-3.9 8-9 5 Bolyanina, 1966bNova river 2.5-3.5 - Leviyeva, 1952Siberia 4.7 - 5.3 12 7 Kyzevetter, 1949Uoh inskoye
waterbody0.5 - 2.4 48 - 6 Original data
North Germany 10.2 5.1 orawa, 1956
36 7764 8
very little 15
61 787 7
32 15
FRm/s78Srnel t
Locality
Yenisey riverLena riverAmur riverSuyfun river and other
rivers of the Far EastElbe riverRivers of the White Sea
basïnSaint lawrence riverMiramiohi river
13 0/00 results in pathological changes indeveloping eggs: flowing out of yolk, deforma-tien of body of embryo, absence of orystallinolens in eyes, destruction of brain9 delay inhatching, sometimes death of embryos. Earlierstages of development (till complote gastrula-tion) aro mora sensitivo to salinity than thoselater (after the embryo body has formed).Hatching prolarvae react positively to salinity
TABLE XXIV
Extent of upriver migrations in different smelt populations
Extent ofmigration
km
1000l80-200
270
16-.18120
2-3
30050
Authority
Tyurth, 1924, and othersPirojnikov, 1950Kuznetzova, 1962
Ivanova, 1955Lillelund, 1961Balagurova, 1957 and others
Magnin and alieu, 1965McKenzie, 1964
3:23
by migrating downstream0 Froehwater has abad effect on the survival of smelt larvae inlater stages of development (Unanyan and Soin,
1963). Lillelund (1961) stated that salmi-ties below 10 0/00 have no harmful effect ondeveloping eggs and larvae, The upper limitsof thermal tolerance during early stes ofdevelopment of eggs lie within the range 17.7-20.7°C, at later stages within the rango 21-
24°C0
4 POPULAfPION
4.,I Structure
4,11 Sex ratio
According to Kirpichnikov (1935) malespredominate in the majority of smelt popula-tions. The number of maies decreases in olderago-groups because of their shorter life cycle(Table xxv). A decrease in the relativenumber of males in each age-group may he seenon comparing the sex composition of spawningand feeding concentrations, This may beossod by the higher total mortality rate ofmales during spawning (Tremblay, 1946).
The sex ratio on spawning grounds is dis-cussed in Section 3.16..
4.12 Age composition
The age composition of different spawningpopulations is given in Table XXVI, Catchesof feeding and, wintering concentrations in-cludo some younger immature fishes,
For age at first capture, age at maturityand maximum age see Table VIII, Most of thespawning (and fishing) stocks ars formed byfish of minimal spawning age - See Table XXVI.
TABLE XXV
Sex ratio in different age-groups ofWhite Sea smelt (original data)
Year and season Sex Ago groups Numberof sampling 2 3 4 5 6 7-8 of fishes
4,13 Size composition
])ata relating length to age and maturityin different stoc.ks are given in Table VIII.
The length-weight relationship of differ-ent stocks is given in Table XIX.
4.2 Abundance and donsit (of population)
4,21 Average abundance
According to McKenzie (1964) the smeltstock of the Mirainiohi river is estimated about375 million fish; Lillelund (1961) estimatedthe stock of the Elbe river of 60-85 millionfish,
422 Changes in abundance
Dornrachev and Pravdin (1926), Petrov(1940, 1947), Dryagin (1948) and others havenoted aharp changes in abundance in short-livedsmelt populations. Lapin (1960) gave an analy-sis of causes of that phenomenon, Sharp de-creases in abundance are related to poor densirof one or two year classes. Usually the popu-lation density recovers in several years,Sometimos a fall in density is a result of anepidemic (Van Oosten, 1947; Patterson, 1948,o' others). In most areas fishing has no
1961Spawning Females - 415 37,0. 100 100 100 184concentrations(spring)
Males - 58,5 63,0 270
Feeding Females 52,1 5405 75.0 75.0 100 100 234concentrations(summer)
Males 47.9 45.5 25,0 25,0 192
1962Spawning Females 66,0 50,0 72,7 66.7 100 335concentrations(spring)
Males - 34.0 50.0 27,3 3303 - 282
Feeding Female s 52,1 56,7 70,5 100 - 152concentra oms(summer)
Malos 470.9 43.3 29,5 127
FRrn/578 Smelt 421
Locality
Pskovoko-Chudskoyo lake 90 IO
Rybinek watorbody 17 77 6Dactey lake 74 17 8 1
Lazmiaden lake 6 37 38 15 4Ladoga lake 6 24 32 25 ii 2
Onega lakeirumiohi riveruron lakeUpper lakeNeve, riverWhite Sea*Onega BayKaridalaksha Bay
Yenisey riverLena riverChat ariga riverAnadyr river
serious effect on population density, but Moro-soya (1960) and Amatisiavsky (1963) have statedthat intensification of the fishery has led toa sharp decrease of catohee in the White lake(Vologcta district) and the Ob estuary.
4.24 Changes in density
Annual varìations in landingi per unit offishing effort for the Miramichi river, Canada,are given in Fig. 11.
Smelt occur in reilatively shallow waters.
Seasonal variations in the available stockare mentioned briefly in Section 3.52
4.3 Natality and recruitment
4.31 Reproduotion ratos
Rupp (1959) found that the survival ratoof eggs of the smelt of Maine is about 6%.According to Kojevnikov (1949) survival co-efficient (from eggs to adult) in the Novariver smelt is about 0.02%.
4.32 Factors affecting reproduction
See Sections 3,16, 3,21, 3.22, 3.23
TABLE XXVi
Ago composition (%) of epawning stocks in different smelt populatione
Age groups1 2 3 4 5 6 7 8 9 10 11 12 13 14 15
4284020 7 I66 30 454 38 8324917 I 1
1284224 3 1
7 32 41 17 3304623 1
8 20 43 23 66 14 8 39 28 5
3 10 10 19 28 25 417 21 33 13 7 4 3 1 1
4033 Recruitment
Slowly growing year classes mature andjoin fishable stocks later than those whichgrow faster. Unpublished data chow that the1958 year-class of White Sea smelt grew slowlyduring the first year of life and reachedmaturity only at the fourth year (only 20%matured at the third year of life). The 1959and 1960 year classes grew faster and maturedduring the third year of life. Similar dataare given by Morosova (1960) and Lillolund(1961).
Recruits enter the epawning stock inspring. The relative number varies from yearto year because of differences in the strengthof year-classes and in the rate of maturing(see above and 4,22).
4.4 Mortality and morbidity
4,41 Mortality ratos
According to Lillelund (1961) averagefishing morta].il;y coefficient in the Elbe riversmelt is 0,17; natural mortality coeffioientfor different age groups is as follows s
Age 1 2 3 4Mortality co- 0.90 1,08 1,30 1,59efficient
Authority
Moshkov & Sorokin,1952
Sohetinina, 1954Wilier, 1926Wilier, 1926Arkhiptzova, 1956
and othersAlevandrova, 1963McKenzie, 1964Baldwin, 1948Be.yley, 1964Kojevnikov, 1949
Balagurova, 1957Original de.te.Kravohuk, 1958Pirojnikov, 1950Luicyanchikov, 1964Agapov, 1941
FR/S78 Sme t
FRm/S78 Smelt
McKenzie (1964) stated that average annualfishing mortality, coefficient in the Miramichiriver smelt is about 3-5% of the stock.
4.42 Factors causing or affectingmortality
Predators: See Sections 3.15, 3,22, 3.34.
Food of larvae and post-larvae and itsavailability: See Section 3,22.
Physical factors: See Sections 3,15, 3,17.Sometimes stocks of smelt become frozen in iceduring the spawning migration in Siberianrivers.
Direct effects of fishing: See Section4.41.
Indirect effect of fishing:
Sa:nsonov (1910) showed that great mortal-ity of eggs of freshwater smelt is caused byfishing gear.
4.43 Factors affecting morbidity
Parasites aoci diseasez: See Table XVII
It is well known (Van Oosten, 1947, andothers) that mass morbidity and mortality ofsmelt occurred in the Great Lakes in 1942/43.
The epidemic cas apparently caused by a virusor a bacterium.
Physical factors causing morbidity of adultsmelt are not known.
4.5 Dynamics of populations (as a whole)
Lillelund (1961) gave a model of dynamicsof year-classes in the smelt population of theElbe river in 1953-60. Lapin (1960) has workedout a scheme of dynamics of age-groups in thepopulation of th small smelt of the Rybinakwaterhody (Fig. is). This model is made on theassumption that the Rybinak smelt spawns onceonly before death, The model demonstrates thatdifferent rates of growth and maturing may causechangea in the age composition of the stock.
4,6 The population in the community amdthe ecosystem
Pirojnikov (1955) included the Siberiansmelt in the Arctic freshwater complex of fish(Huol-jo hucho, Salvelinus fontinalis, Stenociusleuciohthys, Coregonide). Ponornareva (1949)noted that smelt larvae in the Kara estuaryoccur together with larvae of Coregonusautumnalis, Salvelinus fontinalis, Myoxocephalusscoxyius, Liopsetta laoialis and Eleginusn avaga.
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 (from Lapin,1960), Seasons aro shown on the horizontal axis, body length on the vertical.Arrows show that an age-group (or part of it) han reached maturity and spawnedin ,pring (circles signify laid eggs), The dotted line on the level of 60 mmshows that smaller fishes remain immature, Two cases (A and B) with differentinitial age structure of population are considered,
FRm/S78 Smelt
5 EXPLOITATION
5.1 Fishing equipment
5.11 Gears
In the Eibe river they catch smelt inriver-stow-nets, which lie on either side ofa cutter, or the nets may be anchored to thebottom of the river. Each cutter has one ortwo river-stow-nets (Lillelund, 1961). Inthe Groat Lakes they use nylon gill nets of25-38 mm mesh or smaller (extension measure);these are 75 m long. Otter trawis are usedalso; these are semiballoon type, with head-rope lengths of 945-1250 cm and mesh cizes of62 mm in the wings and body and 12 mm in thecod end. Shore seines are used during thespawning run; these are of 12 mm-mesh cottontwine, 180 cm deep and 76 m long. Smeltfrom rivers are captured at sea-lamprey bar-riers In weir traps of 120 x 180 x 180 einconstructed of 6 mm-mesh (bar measurement5hardware oloth over wooden frames, Spawningrun smelt are captured in commercial poundnets 210 cm deep made of cotton twine of 30 mmmesh, shrunk to about 24 mm mesh through treat-ment with prebervatives. The boom shockerconsists of two electrical wire paddies sus-pended from booms extending ahead of a 16-ftoutboard-powered boat. Power for the shockerin supplied from a portable 2.5-kw, 220-volt,3-phase, AC generator (Bayley, 1964).
According to McKenzie (1964), In theMiramiohi River "open-water fishing gear isused at the beginning of the season unless theice is strong enough to set winter fishinggear. Open-water gear consists mostly of bagnets set on stubs or pickets near shores pro-tected from the full force of storms, Thegear is fished from catamarans and man-propelledscows 6-9 m long".
In Russia, smelt is fished by specialtraps ("mereja") made of email-mesh cottontwine stretched on 4-6 hoops of wood with twowings. Traps are fixed by anchors at mouthsof spawning rivers. Shore seines of 80-170 mlong with mesh sizes of 32-30-28-26-20-18-16 mmin wings and 16-12-8 inn (inwards) in the codend (Tyurin, 1939). Large river-fixed-netsand trawls are used also.
Angling for smelt ocours also in someregions (Canada, north-western part of USSR).Baits are earth worms and young fishes,
During the development of the fisherythere has been a tendency to change from simpletraps and cotton gill nets to nylon gill nets,trawling and electrical fishing.
Echosounders and fish detectors aro notused by smelt fishermen.
Smelt is frightened by light, and effec-tive methods of attracting the fish are notknown.
5.12 Boats
In Europe smelt are usually fished frommotor boats of 12-14 m long and 4.5 n wide of40-75 hp (Lillelund, 1961). In North Americacatamarans and man-propelled soows of 6-9 mlong aro frequently used (McKenzie, 1964).
5.2 Fishing areas
5.21 General geographic distribution
Within the area of distribution (see Fig.9) smelt are fished in coastal waters of seas,estuaries of rivers and lakes. During thespawning period smelt are fished in rivers.
5.22 Geographio ranges
See Section 5.21
Smelt are moet abundant in the north-western part of U.S.S.R. and the Arnur riverbasin, Atlantic coasts of Canada and U.S.A.
New rangos of smelt fishery aree Greatlakes, where the smelt was introduced in thefirst quarter of the twentieth century(Speirs, 1951); estuaries of Siberian rivers,whore the smelt fishery did not develop untilafter 1935; artificial waterbodies of theUpper and Middle Volga system.
5.23 Depth ranges
Smelt occur In midwater or near the bot-tom in relatively shallow places. The youngstages carry out vertical migrations in pur-suit of plankton.
5.3 FIshing seasons
5.31 General pattern of season(s)
Smelt are caught mainly in spring, duringspawning. The fishing season Is rather short(see Sections 3.16, 5.33). In autumn thefishery is based on less denso concontrationsthan in spring. In winter angling is usod(McKenzie, 1964, and others).
5.32 Dates of beginning, peak andend of season(s)
The beginning of the fishing season coin-oldes with the formation of offshore feedinconcentrations In autumn and continues Inwinter. The peak of the fishing season isbased on spawning concentrations (seo Tablexv). The season onde after spawning whensmelt have migrated to deeper waters.
thousands of owt
2
22
211
20
11 0
3 2
5z2 PR/73 S
5,33 Variation in late or durationof season
The fishery for spawning smelt is influ-encod. by water temperature and other factorswhich control spawning (see Section 3.16).
According to McKenzie (1964) the fishingseason in the Miramichi river basin from 1931-1963 varied irregularly fron 77 to 136 days.This variation in length of season appears tohave little effect on the final total catch.
The highest proportion of large smelt iscaught early in the spawning season. Sizeof fish declines as the season progresses, andthe price of fish decreases.
5,4 Fishing operations and results
541 Effort and intensity
Lillelund (1961) and McKenzj,e (1964) havegiven statistics of the smelt fishery in theElbe river and the Miramichi river for manyyears, including data on catches per unit offishing effort, Data on Miramiohi annualcommercial smelt landings, 1931/32-1962/63,and catch per licensed net per season are givenin Figs. 16, 17.
9r ye
9-60
Fig. 16 1iranichi annual commercial smeltlandings, 1931-32 to 1962-63.(from McKenzie, 1964).
a
5.42 Selectivity
Bayloy (1964) reported that "the gillnets from 25-125 cm mesh captured almost ex-clusively fish of age group II or older. Theshore seine, weir trap and pound-nets, fishedon].y in the spring, sampled solely the spawn-ing population. Since segregation by maturityprecedes the spawning migrations, immaturo ape-cimens (all fish of ago-group I and part ofaçe-group II) were absent. The otter trawlsand boom shocker fished at times other than thespawning period, appeared to be the least sel-ective gears. Trawl collections usually in-oluded smelt of all sizes and age groups. Theboom shocker fished only in shallow water seem-ed, to sample adequately in the waters fished".
According to McKenzie (1964) "Increasingthe mesh size from 29-32 am increased the gill-ing 5 times arid, in the 38 mm not 7 times. Mostsmelt under 10 cm escape the commercial nets.Sise selection by the nets is one of the fac-tors causing diminution in the size of smeltcaught as the commercial season progresses",
5,43 Catches
Total annual yields for various countriesare given in Table XXVII and average yields fordifferent f ishing grounds in Table XXVIII.
years
Fig. 17 Miramiohi smelt oatoh per licensednet per season (from McKenzie, 196L1).
FRm S 8 Smelt
TABlE XXVII
Total annual yields of smelt in various countries(thousands of metric tons)
(Yearbook of Fishery Statistics FAO, 1966)
TABLE XXVIII
Average annual yields from different fishing grounds
Country1958 1961
Years1962 1963 1964 1965 1966
Total 25,0 27,0 25.0 25.0 29,0 28.0 28,0U.S.S.R. 14.5 15.1 15,3 15.5 16.1 16,7 16.6Canada 4.3 7,3 1.5 1.6 2.1 2.1 2.1U.S.A. 6.0 2.5 2.2 2.3 2.4 1.8 1.4Finland 0.1 1.1 0.6 0.4 0.7 0,8 0.9France - 0.3 0.2 0.1 0.1Federal Repubilo
of (hrmany 0.4 0.2 0,2 0,2 0,4 0.3 0.3Netherlands 0.2 0.1 0,2 0.1 0,2 0,3 0,3Norway - 0.9 - 0,1 0,6 0,2 0.2
Fishing grounds Average yield(tons)
Years Authority
Pskovsko-Chudskoyelake
limen lake
3600
363
1931-1963
1946-1958
Shlrkova & Pihu, 1966
After Kudereky, 1962White lake 261 1945-1958 After Kudersky, 1962Onega lake 623 1945-1959 After Kudereky, 1962Ladoga lake 1374 1946-1954 After Kuderaky, 1962Neya river 400 1932-1948 After Kojevnikov, 1949Finnish Gulf 640 1932-1948 After Kojevnikov, 1949Kurishes Haff 983 1948-1957 After Noskov, 1959Elbe, Eider, Weser 311.5 1955-1959 Lillelund, 1961White Sea 400Yenisey 150.7 1946-1955 Podiosnyy, 1958Ob estuary may. 1540 1960 Amstislavsky, 1963Amur 1000 1949-1958 Kuznetzova, 1962Miramiohi river 726 1931-1963 McKenzie, 1964Great lakes 7264 1960 Bayley, 1964
FRm/S7U Smelt 6l
6 PROTECTION AND MANAGEMENT
6,1 Regulatory (legislative) measures
6,11 Limitation or reduction oftotal catoh
In many oases catches of smelt may be in-creased, as only a email part of the popula-tion is removed by fishing. However, in somecases it is necessary to limit catches(Morosova, 1960; Ametielaveky, 1963).
6.12 Protection of portions ofpopulation
The size of caught smelt is limited bythe market 8 for example in the Elbe riverthey do not catch smelt snaller than 14 cm(Lillolund, 1961).
6.2 Control or alteration of physicalfeatures of the environment
Physical features of the environment maybe altered to Improve spawning grounds byclearing such impassible obstacles as brushjams, old dams, steep rapids or falls, sluice-ways, pulpwood logs as well as road oulverts.McKenzie (1964 noted that "at moderate costmuch can be done to overcome these situations,Streams full of brush and debris from lumberingoperations oan be cleared and laws enforcedto prevent recurrences. Old dams can be re-moved or if in use can In some cases be openedduring the smelt spawning season. "Smeltways"can be built also to permit passage over thedams. Grades through culvorts should be madeto conform to the slope of the original streambed with addition of baffles, if necessary, to
break the swift smooth flow, The bottom ofthe culvert at the outfall (at least whenbuilt) should be level with, or preferably low-er than the stream bed. Action to facilitatethe passage of smelt through highway oulvertsto their spawning grounds would be of consider-able importance to the smelt fishery in NewBrunswick, Smelt often escape the strong cur-rents in streams during spring froshets bygoing out over banks and spawning far from theregular stream bed, In some locations itseems practical to prevent loss of both eggsand parents from stranding by constructingdykes. In the larger Miramiohi branches andtributaries flood control by means of damswould no doubt increase the larval production".
6.5 Artificial stocking
6,52 Transplantation, introduction
Transplantation of' smelt from the Greenlake (Maine) into the Crystall lake (Michigan)was made in 1912 and in following years andhas given excellent results (Van Oosten, 1931;Creamer, 1926, 1929, and others). Smelt hasspread all over the Great lakes and formed agood fishery population there (Table XXVIII).In Russia smelt was repeatedly transplantedinto some lakes of the Leningrad and NovgorodDistricts (Píkhiy, 1941). These operationshave given good results in 42% of oases forsmall smelt but for long-lived smelt only 17%have been successful. As a rule unrepeatodtransplantation has rarely given good results.Snielt wan introduced into some Ural lakes also(Karabak, 1930), Some introduction of smeltinto the waters of Karelia were proposed byStefanovskaya (i 957).
FRs 'S Smelt 7:1
7 POND FISH CULTURE
7.9 Transport
Smelt has not been cultured in ponds./ccording to Karabak (1930) and Suvorov (1939)smelt eggs may be incubated and transported ina cool wet atmosphere. 4ortality of eggs
during 5-6 days i iena than 5% (lcarabak,1930).Good survival of transported smelt eggs andlarvae was noted by Richardson and lknap(1934), fishes of 6-10 cm length being the moathardy. Early larvae are too de].icate and theyshould not be transported. Smelt eggs arogathered on speoal i:creens during the spawn-ing period.
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*
SYNOPSIS OF FISHERIES BIOLOGICAL DATA
This is one of a series of documents issued by FAO, CSIRO and USFWS concerning speciesand stocks of aquatic organisms of present or potential economic interest. The primary pur-pose of this series is to make existing information readily available to fishery scientists ac-cording to a standard pattern, and by so doing also to draw attention to gaps in knowledge.lt is hoped that synopses in this series will be useful to other scientists initiating investiga-tions of the species concerned or of related ones, as a means of exchange of knowledgeamong those already working on the species, and as the basis for comparative study offisheries resources. They will be brought up to date from time to time as further informationbecomes available either as revisions of the entire document or their specific chapters.
The relevant series of documents are:
RAO Fisheries Synopsis No. FR/Sreplacing, as from 1.1.63 FAO FisheriesBiology 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/SiRev. 1 (1965).
FAO, CSIRO and USFWS are working to secure the cooperation of other organizations and ofindividual scientists in drafting synopses on species about which they have knowledge, andwelcome offers of help in this task. Additions and corrections to synopses already issuedwill also be most welcome. Comments including suggestions for the expansion of the outlineand requests for information should be addressed to the coordinators and editors of theissuing organizations.
FAO:W. FischerFishery Resources and Exploitation DivisionMarine Biology and Environment BranchFood and Agriculture Organization of theUnited NationsViale delle Terme di Caracalla00100 Rome, Italy
USFWS: CSIRO:L.W. Scattergood, Chief, Branch of Reports Maureen A. Wright, Scientific EditorU.S. Department of the Interior CSIRO Division of Fisheries and OceanographyFish and Wildlife Service Box 21Bureau 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 preparationwill be issued from time to time. Requests for copies of synopses should be addressed tothe issuing organization.
The following synopses in this series have been issued since January 1969:
BCF/S40 Synopsis of biological data on the Pacific mackerel Scomberja pon/cus Houttuyn (Northeast Pacific)
FRi/S30 Synopsis of biological data on the Pike Esox lucius (Linnaeus)Rev. i 1758
FRm/S43 Synopsis of biological data on the anchoveta Caten grau/ls mysti-cetus Günther, 1866
DFO/S3 Synopsis of biological data on the Tiger Prawn Penaeus escu-lentus Haswell, 1879
FRh/S35 Synopsis of biological data on West African Croakers Pseudo-Rev, i to//thus typus, P. senegalensis and P. elongatus
FRm/S78 Synopsis of biological data on smelt Osmerus eperlanus (Lin-naeus) 1758
February 1969
May 1969
October 1969
July 1969
November 1969
December 1969
MI/96024/3.70/E/1 /750
