Oieg V. Karaiiiushko, Amilansai. Karaiiiiishko3

CM 1997/CC: 14 Biology arid Bchaviour (TI) (CC)

, .. ''- ... RE~RODUCTIVETRAITS OF MARINE FISH ~ RELATION TO THEIR MODEOFOVWOSnnONANDZOOGEOGRAPmCDffiTmrnUTION

Jm-gen S. ChristiansenI, Svem E. Fevolden2, Oieg V. KaraIIiushko, ami Lansa I. KaraIIiiIshko3

. 1. Norwegian InStituteofFisherles ami AqiiacUlture, N-9()()5 Trema, Norn:äy 2. The NorwegianCollege ofFishery Science, UniversitYofTroriisa, N-9037 TrOmSo, Norway • 3. MUrmaDsk Marine BiologiciU InStitirte, 183010 MUrriiäiisk, . . . RuSsm

ABSTRACT

. 1t is generatly. reported that polar fish species produce. relativety few, luge, eggs . .comparoo with that offish species ät·lower hititudes. However,egg siZe.and fecundity(i:' e. the total number ofeggs to be' spawned) in fish is ihought also 'to be Iiilked.to tlieir mode öf.oviposition. This commuilication outllnes how egg siZe,.feeundity, arid henee, . .reproduCtlve output of j~ marine fish species cOvanes with Doth zoogeography (L e. polar vs boreal) an.d mode ofoviposition (i. e.' pelagic vs demersal). Tbe results siiggest . that i) only.pelagic polar .fish,produ,ce relatively few, large eggs compared with their .. boreal counterparts ü) substrate' spawning .species produce relatively few, läfge eggs irrespective ofzoogeographie' regions, arid ili) maiemal output, i. e. the product ofegg .'voluriie aiid feCWidity, does not differ between polar arid boreal specles within the same mode ofoViposition. Consequently, behaviouriU arid ecologicaI factOfs arid not hititiide per se seem to be the major deteriniIiarits iIi the reproduCtlve triütS cfiriarine fish species.

K~y 'words: Egg size, feCiinditY, rnateriiaI output, mode cfovipoSi~ori, zoogeography

'. . , 2.

INTROnUCTION

In oviparous fish, the combination of egg size and ,fecundity (i. e. the total number of eggs to be spawned) that propagates the largeSt ninnbers ofviable offspring is deemed to be favoured by evolutionary processes. Polar manne fish species appear to produce fewer arid also lärger yolk rieh eggs compared with their tower latitüde coUntei-parts. In his paper from 1941 RaSs propolinded: "Tbe size of eggs of aquatic ammaIs is in accordance with their breeding teinperatures" and ce••• the eggs beilig greater, the nearer to the pole lies the habitat ofthe spedes". Rass' statement that the physiciU environment (i. e. temperature) .Conditions the increase in egg size. towards higher latitudes, was, however, opposed by Mai"shall (1953). He argued that the llititudinal cline observed in egg size and fecundity couId be aseribed to sundry beh8vi6uraI and ecologica1 factors, .e. g. availability of food, rather than to a single physica1 mable sueh as breedirig temperature per se. Tbc egg incubation period (i. e. the time span between fertilizatiori , and hatching) does. not seem to con-elate with egg size within fish species,at any, one temperature (Blaxter'1988).but large eggs häieh with lüge larvae (Blaxter 1988; Koek , 1992). Large larvae subsist longer on endogeneous met, are less affectoo byvariations in • exogeneous food supply and this may reSUIt in more viable offspring (MaiteinSdottir arid Ahle 1992).

Polar fish are subjected to an extreme seasonality in food availability since a short period ofa high production cfplRnktonic orgamsms is followed by long perioelS with littIe or no food in the water colunin (Ursin 1984). Tberefore, according to Märshall (1953), the adaptive value oflarge egg size,and !bus, greater lärvae survival iS partlcuIarly imPOrtaIlt in habitats. which are clulracterlzed by relatively sparse food concentrationS e. g. the , deep-:sea, open oCearllC witers, arid seasoIlaIly, the polar seai " '

AIthough it is gene~y' reported that,polar fish produce relatively few, larg~ eggs, egg size and fecundity ~ fish is StrongIy linked totheir mode ofovipositiOli (BlaXier 1969). ' Fish that spawn freely floating eggs (i. e. display pelagic ovip6sition) ofteri produce snianer eggs and, are more' fecund crimpared to species that deposit theireggs ontormto . 'Substrates (i.: e. display demerSaI oviposition) (Tyler and Sumpter 1996). :nie ritimber of studies' deaimg with egg size' arid feciindity in fish' ~ 'increased, e eonsiderably in recent years, and it also includes reviews on Aritarctic fish reproduction (Kock and Kellennan 199i; Hubold 1992). In the light" ofthe available information, this communicatiori ouilines' how egg size, fecuIiditY, arid hence, reproduCtive output of marine fish species cOvarieswith both zoogeography (i. e. polar vs boreal) and mode of oviposition (i. e. pclagic vs demersat). . . :METHons

Tbe data base

A total of 35 fish species, represented by 58 data sets, is comprised in this study. All species are marine and predominaIitly members 'of the Gadidae (northem hemisphere) .. " and 'Notothenüdae (Antarctic) faInilies. Fish were examined with' .respect to their zoogeographie distribution and mode cf oviposition (Table 1). Zoogeographica1Iy, fish were divided into two broad catt~gories vii. polar (i. e. Arctic and Antärctlc combinoo) . . ". '.:. , 4

arid boreal species. The' distiriCtion betWeen'boreal arid gellUinly ArCti6 species follows the authority of Äridriyashev and Chemova (1995). For rwons of siniplificiition, we have trea.ted .A:ritarctic fish aS belonging to thei same geographie region (i. e.' witmn the boundary of the AntarCtic Convergence)although Kock and Kellennann (1991) and Kock (1992) hRve made a fuitherdistinction betWeen species living in the High-AntilrCtic . Zone iilid the SeasoriaI Pack-ice Zone respectively.

Moreover, flsh were grouped accofding to therr mode of oViposition Vii. peiaglc spawners iri. which eggs develop freely in the water column arid substrate spawners in which eggs develop at or elose to the sea floor (Table i). FOT example, althmigh a bottom-dwelling speCies in adult stages, Liopsetta liiiuinda was. put into the. pelagic group smce eggs, aS iri most flatfish, develop iri thei water column. By coritnisi, MallaluS vi/lasus was categorlzed as demersaI; it is a truei pelagic species in adult Stages but eggs develop in large clusters onto substi-ät~s.

Fulther, our data base includes average values regardIDg body length (m min) ofsexually matUre fish· and, the corresponding egg diameter' (in mm)and feciIDdit}r (Table ) .. •• Information häs been gatbered ritainly from the works by Rass (1941), M8rsliall (1953), Kock arid Kellenilann (1991), Hubold (1992) arid Rass (1992).

Essentially, the reproduCtlve ~utPut byfeinale fish depends on tw~ key-parameters viz. egg siZei '.. arid. fecundity. ASsuming thRt eggs ... are ., spherieat,. egg .diameters were 3 trariSformed into volurries (in mm ). We here define the reprodtictive output by afemale' fish, i e. maternal output (MO), as: '

MO = EV*F wher~ " EV is the average volurne ofa single egg and F is fecundity (i e. thei total riiJmber ofeggs to be spaWned per iridividuat fish). Thus, MO represents the total biomaSs of 3 'releasaI>le.egg produ~s arid is expres'sed in,rom •

. , Statistieat analysis .

The dependence of thei reproductive ~ariables (ie. egg size, f~dity and maternat. • output) on body lengtll arid combiriations ofzoogeography (polar vs boreal) arid mode of oviposition (petagic vs demersat) was tested by. means of arialysis of vanance , using . zoogeography/oviposition as a flXed class variable and body length as a continuöus variable (Table 2). The GLM procooure ,iii SAS was used to estiritate F-vaIues arid sigiii.ficance levels (SAS IriStitute Inc. 1985). To meet tbe requirements' of nonDalitY fectiDdity arid maü~rnal output was log-transforriiCd in the variance ariälyses.

~ '- , RESULTS

Detaileid inforinätion regaiding the reproduCtlve parameters, i. e. egg siie, fecimclltY, and rnate1lüU outpu~ is preserited in Table 1. .More than one data set was obtameid for same species: Dtie to reaSons ofclarity we mve not Ideniifieci the mdiVidual species in Figtires .1 tö4. .',. • t 4 . c. Eggsize

Correlations between egg size (polar vs boreal; pelagie vs demersal) 80d body lerigth are shown in Fig. 1 A arid B. Egg size in marine fish speeies seenis to be a eharaCteristic cf both zoogeography and mode of oviposition. Within the group of pelagie spawning' species, polar species produce significantly larger eggs tluin do boreal species (Fig. 1 A; Table 2). By eontrast, the size of demersally deposited eggs do nöt differ. between zoogeographie regions (Fig. 1 B; Table 2). Further, egg size inereases with body length of speeies irrespective ofzoogeography 80d mode of oviposition With the exception of boreal pelagie spawners. Por this group cf species, egg size is consistantlY,low at 80Y body length (Fig. 1,A). Consequently, i) the ovefall egg size appem to mcrease With ·body length, and ü) substrate spawners pröduce similar sized eggs irrespective of zoogeography whereas egg sizes of pelagie spaWning speeies are largest towards the poles.

Fectiodity

Correlations between fecundity '(polar vs boreal; pelagie vs demersal) 80d body length are shown in Fig. 2 A and B. Feciundity increases with body length irrespective of • zoogeography 80d mode ofovipositioo (Fig. 2 A arid B; Table 2). Further, fecunditY data of pelagie spawners cluSter inta .tWo. distinet groups with '. polar species being significaritly less fecurid at agiven body length ccimpäred With that ofboreal species (Fig. 2 A; Table 2). This is in cOntraSt to substrate spawners where fecuridity does riot differ betWeen polar and boreal species (Fig. 2 B; TabeI2). '

Matemal output

Tbe correlations be~een m~teriial. output (polar vs borea.t; pelagic vs 'demersal) 80d . ·body lerigth are shown in Fig. 3 A änd B. Mateciial outPut, i. e.the product of egg volume and fecundity, increases with body lerigth itTespective of zoogeography 80d mode of oviposition. Moreover, there are no zoogeographie differences in rnatenial output within modes of oviposition(Fig. 3'A 80d B; Table 2).. However, when the "., matemal output betWeen modes of oviposition, i. e. pelagie vs demersal spaWners, is eompared (Fig. 4), the maternal output ,for pelagie spawners tuins out to be significantly . higher than that seen amongst substrate'späWnmg species irrespectlve ofbody.1ength (F • =33.49, P =0.0001; analysis ofvari8Oce). . '

DISCUSSION

Data regarding the modes ofoviposition in Antarctie fish are sparse. Most members of the family Nototheniidae are considered to be substrate spawners. As they approäeh sexUal maturity the development offully mature gonads seeins to require twO succesSive seasons, and eggs are probably deposited in one bateh amiually then~after (Everson 1977, Kock and Kellerman 1991).

Morphological eharacteristies of. egg-sh~lls are. specles-specific 80d developing pelagic eggs have been identified from a few species i. e. Notothenia rOsS;i arid N. neglecta ·(coniceps) (White et w. 1996). Other species' sueh as Chainpsocephalus guiuiari 80d pisiostichus eleginofdes niay also produ.ce pelagic eggs .(White et 8J.. 1996) and they 5 •• have, therefore, heen consi"dered as heing pelagic spawners in this st11dy' (Table 1). On . the ether harid, we, have cluir-acterized. P/eurairamma antarcticum (G. HUDold, pers. conUn.) and DissostichUs iizäwsoni. as heing"demersal spawners although the latter have many featUres in conUnon With the pelagic spawning species, i. e. a high fecuridity arid materriaI output (Table 1).

•.-

•-,'\,':' .' • t 6 ., cciIToborai'e the suggestion made by M~shal1 (1953), tlult behavioural' and ecological fuetors seem to be the major detenninarits in the reprodtietive traits cf marine fish speeies. .

ACKNOWLEDGEMENTS

We wish to thank Professor Aßdrew Clarke, Dr. Gerd Hubold, and Dr. Karl-Hermann Kock for valuable coniments.

LITERATURE CITED

Andriyashev, A P. arid Chernova, N. V. 1995. Arinotated list offishlike vertebrates arid fish oftha Arctic Seas and adjacent waters. JoWna1 ofIchthyology 35 (1): 81-123.

,Blaxter, J. H. S. 1969. Develop~~nt: eggs and llirvae: In: Flsh Physiology (W. S. Hoar andD. J. Randall, eds), Vol. 3: 177-252. AcademicPress, New'York.

Blaxter, J. H. S. 1988. Pattern and variety in developinent. In: Fish Physiology (W. S. Hoar and D. J. Randall, OOs), Vol. HA: 1-58. Acadeniic Press, New York. •

Chemova, N. V. 1987. About breeding of hump-backed sriailfish Liparis" gibbus Bean (Scorpaenuormes, Liparidae). In: Ecology-morphological investigationS of commercial fishes in the northern basin. Leningrad, NaukR: 87-94. (in Rtissian).

Chemova, N. V. 1991. Snailfishes (Liparidae) from the Eurasian Arctic. APatity. 112 pp. (in Russian). ..',

.. , .. '.... Clarke, A, ,1987. Temperature, latitude and, reproductive eifort. Marine ,Ecology Progress Series,~8: 89-99. ", ... '. Daoulas, Ch. and Ecoriömou, ,A N• .1986. SeasorW variation ofegg siZe in the sardine, Sardina pilchaTdus Walh., cf the Saronikos Gulf: causes and a probable exphination. . ' .Journal ofFish Biology 28: 449457.

, . Everson, I. 1977. The living resources ofthe Southem Ocean. FAD, GLO/S0I7711: 1': 156. ,..... " .... .' • Gerasimchuck, V. V. 1987. On the fecUntllty of Antarctic sidestrlpe, Pleurogramma antarctieum. VoproSy Ikhtiologii 5: 858-860. (in Russian).

Gusev, E. V. 1995. Individual fecundity of wolffish family Anarhichas iri the Barerits Sea. In: The problems,of fishery scierice in creation of yoting scientlSts. PINRO, Mumiansk: 252-270. (in Russian).

Hislop, J. R G. 1984. A comparison ofthe reproductive tactics and strategies of cod, häddock, whitIDg and Norway pout in the North Sea. Iri: Fish Reproduction: Strategies arid Tactics (G. W. Potts and R J. Wooton, ecIs), Aeadenuc Press, London: 311-329.

Hubold, G. 1992., ~ur Ökologie der Fische ~ Woodellmeer. Berichte zur Polarforschung 103: 1~157.

.. . .' . " ' , . 7

.' "

Kock, K-H. 1992. AIitarciic fish arid fisheries. (Studies iri polar research). Cambrldge University Press, Cambridge. 359 pp.

Kock, K-H. arid Kellennann, A 1991. Reproduetion in Antarctic notothemod fish. Antarctic Science 3 (2): 125-150.

Marshall, N. :B. 1953. Egg size in Aretic, Antaretic and deep-sea fishes. Evolution 7: 328-341.

Marteinsdottir, G. and Ahle, K. W. 1992. influence,ofegg siZe on embrYos arid laivae of "' .. FuiuJulus heteroclitus. Joiiriial cifFish Biology 41: 883-896.

Mazhirina, G. P. 1994. Reproductive charaterlstics of blue whiÜng, Micromesistius pouicissou, in the northeastem Atlantic. Journal ofIchthyology 34 (1): 132-139. .

Pozdnyakov, Yu. F. 1957. About fecunclity of eapeliii cf the Barents seä. Doklady • ., .. Alaidenili NaUk SSSR 112 (4): 777-778. (m RUssian). Rass, is. 1941. AiWogous or para.Ilel variations in strtictUre and development offishes mnorthem and ai-ctl6 seäs. Jubilee PubliCätions af the Moscow Society of NatUralists 1805-1940: 1-60.

Riss, T~ 8.1968. Spawrrlng arid development ofpolar ead. leES Rapport P-v Reunion 158: 135-137.

Rass, i S. 1992. Metamerism ofba.ckbone arid. siZe of eggs in specinieris cffish from . nortlH:m seäS. Doklady Akademii N8uk 324 (5): 1131-1135. (in RUssian).

., " SAS Insti~te Ine. 1985. SAS User;s Guide: StatiStlcs, Version 5 Editi~n. Cary, NC: SAS InstitUte. ., .'

Tokfa.nov, A M. 1988. Some aspects of the biology of the K3.fficha.tkan sCulpin, Artediellus camchaticus, on the eastem coast ofKanlcha.tka.. Voprosy Ikhtiolcigii 3: ~15 • 420. (in RUssian).'

Tseyeb and Pozdnyako~, Yu. F. 1975. The ichthyofauna of Lake MagnIloye. m: M3.gilnoye Lake. Aka.demii Nauk SSSR; Murmansk Marin BiologiCal Institufe. (ul . Russian).

Tyler, C. R and Sumpter, J. P. 1996.00cYte groWth and development in teleoSts. ReViews iri Fish Biology arid Fishenes 6: 287-318.

Ursin, E. 1984. The tropicai, the temperate and tbe arctic seas as media for fish. production. Dana. 3: 43-60.

Vacchi, M., Williariis, R., and La Mesa, M. 1996. ReproduCtion in tlu-ee species offish .. fioiri ihe Ross Sea. arid Mawson Sea. AntarCtlci Scienee 8 (2): 185-192. .' - - . ( 8 ~ •.. White, M. G., Veit, R R, Noith,'A W:, and Robinson, K.'1996. Egg-shell morphology ofthe Antarctic fish, Notothenia rossii Richardson, and the distribution and abunciance ofpelagic eggs at South Georgia. Antarctic Science 8 (3): 267-271.

Wooton, R J. 1985. Energetics ofreproduction. In: Fish energetics. New perspectives. (p. Tytler and P. Calow, eds), Croom Helm Ltd., Great Britain: 231-254.

LEGENDS TO TABLES

Table 1. Reproductive parameters of 35 marine fish species. Species are grouped according to their mode ofoviposition, i. e. pelagic and demersal, and zoogeography, i. e. boreal and polar (Arctic and Antarcuc combined). BL = body Iength; ED = egg '. diameter; EV =egg volume; FEe =fecundity; MO =materriaJ. output. .

Table 2. ANOVA tests ofdependence ofreproductive variables on body length, zoogeography, and mode ofoviposition. ns =not significant.

LEGENDS TO FIGURES

, . • Figure 1. CorrelatioDs between egg 'diameter (polar vs boreal; pelagic vs demersal) and bodylength.

Figure 2. Correlations between fecundity (polar vs boreal; pelagic vs demersal) and body Iength. Note the logaritmic sca1e for fecundity.

Figure 3. Correlations between matemal output (polar vs boreal; pelagic vs demersal) and body length. Maternal output is defined as the product ofegg volume and nuinber of . eggs. Note the Iogaritmic sca1efor maternal output. .

. , Figure 4. Correlations between maternal output of pelagic and deme~ species (polar and boreal combined) and body length. Maternal output is defined as the product ofegg 'voltime 'and numberofeggs. Note the Iogaritmic'scale for maternaI output. .. • , TABLE1• .-• Speeies EV(mm31 FEe (nollOOI MO (mm3l\ndl Soure. . . Pelaglc Boml

"'>,,. ;,... Ttisopteru$lIiScUS 150 0.97 0.478 207479 .99187 Rass 1941 Ttisopterus Iuscus 200 1.23 0.975 835997 814865 Rass 1941 Ttisopterus esmarId 150 1.00 0.524 61686 32311 Rass 1941 TtisoptenJs esmarId 250 1.13 0.756 384190 29D301 Rass 1941 Odontogadus merlangus 300 0.97 0.478 1D9358 52279 Rass 1941 Odontogadus merlangus 550 1.32 1.205 131712 158671 Rass 1941 A:JI1achius vir8ns 600 1.03 0.572 500000O 2861852 Rass 1941 PoIJachius vir8ns 9DD 1.22 0.951 8260000 7856426 Rass 1941 Melanogrammus aeglelinus 500 1.19 . 0.883 170000 15Oll57 Rass 1941 MeIanogrammus aegJefinus 750 1.67 2.440 1800000 4391242 Rass 1941 Gadus morhua tIIOIhua 500 1.13 0.756 270000D 2040632 Rass 1941 Gadus tIIOIhua tIIOIhua 800 1.65 2.353 910000D 21412073 Rass 1941

MoIvamolva '''.'.,':' 1000 1.05 0.6D6 BIaxler 1969 Gadus mothua ldJdinensIs 525 1.00 0.524 Tseyeb & PozdnyakDv 1975 Gadus mothua Idldinensis 628 1.75 2.807 Tseyeb & PozdnyakoV 1975 Merlangius merlangus • 300 1.16 0.818 Hisiop 1984 Mictomesistius poutassou 303 o.n 0.196 122DOO 23852 Mazhirina 1994 .. PeJagie Polar

Antarctic " "'i'" , Notothenia toSsii ·510 4.50 47.731 19271 919829 Kock & Kellermann 1991 Notothenia fOSSil 780 . 5.00 65.475 9933D 6503632 Kock & Kellermann 1991 Notothenia neglecta (coriiceps) 340 4.30 41.646 6897 287231 Kock & KeIIelmaM 1991 Notothenia neglecta (ooriiceps) 550 4.70 54.382 41024 2230987 Kock& KelJermann 1991 Dissosticus eleglnoides 930 4.30 41.646 238314 9924769 Kock& Kellennann 1991 e Dissosticus e/et;}inoides 1310 4.70 54.382 545665 29674620 Kock & Kellermann 1991 ChampsocephaJus gunnarl 340 3.50 22.458 3094' 69485 Kock & Kellermann 1991 ChampsooephaIus gunnarl 500 4.10 36.101 11664 421080 Kock & Kellennann 1991

Arctic " Bot8ogadus saida 237 1.57 2.027 16625 33700 Owndata Bot8ogadus saida 120 1.55 1.951 30000 58517 Rass 1941 Bcx80gadus saida 160 1.84 3.263 67000 218622 Rass 1941 Liopsetta Iimanda 350 1.40 1.437 125CXXl 179663 Rass 1992

D8mersal Bornl uPans inontagut 125 1.19 0.883 ,793 700 ~1941 Cyc;Jopterus lumpus , 600 2.60 .9.206 194112 1787055 Rass 1941 Anartlichas tupus lupus 1250 6.00 113.141 .12740 1441414 Rass 1941

Agonus cataphractus " ," ,." 200 2.23 5.809 2400 13941 Rass 1941 MyoxocephaJus scaplus scorpius 250 2.51 8.283 2742 22712 Rass 1941 '<" .•.• ' .. MaJlotus vlJIosus , 153 1.20' 0.905 11874 10747 PazdnyakoV 1957 Msdiellus camchaticus 160 3.40 20.587 370 7617 Toknmov 1989 Anarhichas minor 63D 2.70 10.310 4200 43302 Gusev1995' Anarhichas minor 1200. 4.00 33.523 . 35200 1180017 Gusev1995 Dem.rs.1 Po" " "'., Antarctic· . ,~ .,.": Pleuragramma antarctlcum 154 1.80 3.055 4315 13181 Gerasimchuck 1987 PIeuragramma antarctlcum 256 2.00 4.190 1m4 74480 , Gerasimchuck 1987 Dissosticus mawsoni . 1200 4.00 33.523 471900 . 15819598 Kock& Kellennalin 1991 e . Dissosticus mawsoni 1800 4.30 41.646 1343380 55946090 . Kock& Kellermann 1991 • Harpagifer antarcticus 110 2.50 8.184 400 3274 • Kock & Kellermann 1991 Harpagifer antarcticus 110 2.50 8.184 1000 8184 Kock & Kellermann 1991 Nototheniops nudiftons 115 2.00 4.190 544 2280 Kock & Kellermann 1991 Nototheniops nudiftons , 191 2.50 8.184 3110 25453 Kock& Kellermann 1991 Pagolhenia borchgtrNenId 165 2.40 7.241 1500 10862 Kock & Kellermann 1991 Pagothenia borchgtrNenId 235 3DOO Kock & KelIermann 1991 Tremalemus bemachII 200 3.60 24.438 1200 29326 Kock & Kellermann 1991 Trematemus bemachII 288 3.80 28.742 Vacchi et BI. 1996 Trematomus hansoni 320 3.10 15.605 9600 1498D3 Hubold 1992Nacchi et al. 1996 Trematemus haIISOIIi . 440 3.60 24.438 9600 234609 Hubold 1992Nacchl et BI. 1996 '

Arc:tlc EJeglnus navaga 180 1.50 1.768 2DDOO 35357 R§S1941 EJeglnus navaga 200 1.90 3.593 62DOO 222750 Rass 1941 Uparis gIbbus 135 1.13 0.756 4000 3023 Chen'1lMl1987 Uparis gibbus 180 1.70 2.573 12000 30881 CheIllCMl 1987 Catepn:x;tus lafIu/a 54 3.50 22.458 Chen'1lMl 1991 Rhodichthys reg;na 310 7.00 .":, 179.663 Chen'1lMl1991 • t

• •

Table 2. ANOVA tests ofdependence ofreproduetive variables onbody length, zoogeography. and mode ofoviposition. ns =not significant

Bodylengtb PolarIBoreal F P F P Pdagie spawnen Egg diameter 12.82 0.0014 57.79 0.0001 Fecundity 18.78 0.0003 37.23 0.0001 Materna1 output 67.16 0.0001 0.41 ns

Demersai spawnen Egg diameter 10.89 0.0029 0.42 ns Fecundity 30.36 0.0001 1.85 ns Matemal output 87.90 0.0001 2.81 ns •

•

: " . .. . , .' Fig.'1A. PELAGIC SPAWNERS 6-r------,

5 0 0 0 0 0 0 4 0 -E 0 E 3 -'-' S CI) E 2 .!!! 0 "C • 0 0 • • C) 0 C) • • • • W 1 • • • •• • • • • Boreal o Polar 0 • 600 800 1000 1200 1400 0 200 400 Body length (mm)

Fig. 1B.DEMERSAL SPAWNERS

•

o o o o o • o ••

'. Boreal I '.I C;;Polar O-!-'-'--~--+---_+---_+_---I__--_+--'--_+_---I__--_+-----l o ,200 400 600 800 1000 1200 1400 1600 1800 Body length (mm) . ,

Fig. 2 A. PELAGIC SPAWNERS

10000000 . • • • • 1000000 • • 0 • 0 0 • ~ 100000 • • 0 ~ co 0 "0= 0 "> 0 0 :s 10000 0 .-C 0 _0 0 C -~1000 "0 C • Boreal =CJ o Polar CI) LL 100 • o 200 400 600 800 1000 1200 1400 Body length (mm)

Fig. 2 B.-DEMERSAL SPAWNERS

10000000

1000000

0 • ~ 100000 CO 0 "0= • 0 0 "> • 0 :s 10000 0 0 • C - 00 ci ~ce • c 0 0 -~- 1000 0• -"0 0 • Boreal ·c 0 • =CJ o Polar CI) -LL 100 0- 200 400 600 800 1000 1200 1400 1600 1800 Body length (mm) • •

• • .Fig. 3A. PELAGIC SPAWNERS

100000000 ...------• 0 10000000 .0 • o ":' 0 • -'ii • ::J 1000000 0 "C • 'S; 0 0 0 • :s 0 c 100000 • • • 0 f') 0 E 0 • E • • -'$ 10000 Q. ::J .() 'ii 1000 • Boreal E ~ o Polar :!: 100-1-----1-----+-----+----+-----+ o 200 400 600 800 1000 1200 1400 Body length (mm)

._- -_.

• Boreal o Polar

1400 1600 1800 • •

• t Fig. 4. PELAGIC vs DEMERSAL SPAWNERS ""

100000000 • • 0 ":" 10000000 Ci- • •• :::l • 'C •0 • • 0 ~ 1000000 0 'C • • ..,.5 • .0 • • 0 o· E 100000 • • E • 0 • • • 0 - ·~·o :::l -Co • " 10000 Ibo :::l o 0 -0 Ci 00 "...c:: " 0 CI) ." 1000 0 CU ::E.- • Pelagic o Demersal 100 • o 200 400 600 BOO 1000 1200 1400 1600 1Boo Body length (mm)

ERRATUM

1) Instead ofLiopsetta limanda the species should read Liopsetta glacialis throughout the text .

2) The eorreet referenee to Tseyeb and Pozdnyakov (1975) is as folIows:

Tseyeb, R. Y. and Pozdnyakov, U. F. 1975. Eeology of the eod from Kildin !sland: reproduetion. In: The Reliet Lake Mogilnoye. Leningrad. Nauka: 227-247. (in Russian).

"." CM 1997/CC: 14

REPRODUCTIVE TRAITS OF l\1ARINE FISH IN RELATION 1'0 THEIR l\10DE OF OVIPOSITION AND ZOOGEOGRAPHIC DISTRIBUTION

J0rgen S. Christiansen, Svein E. Fevolden, Oleg V. Karamushko, and Larisa 1. Karamushko

ERRATUl\1

1) Instead ofLiopsetta limanda the speeies should read Liopsetta glacialis throughout the text

2) The eorreet referenee to Tseyeb and Pozdnyakov (1975) is as folIows:

Tseyeb, R. Y. and Pozdnyakov, U. F. 1975. Eeology of the eod from Kildin Island: reproduetion. In: The Relict Lake Mogilnoye. Leningrad. Nauka: 227-247. (in Russian).