FAO Fisheries Synopsis No. 84 FIRM/S84 (Distribution restricted) SAST - Haddock - 1,48(04),O1O,O1

SYNOPSIS OF BIOLOGICAL DATA ON HADDOCK Melanogrammus aeglefinus (Linnaeus) 1758

Prepared by

R.W. Blacker

FOOD AND AGRICULTURE ORGANIZATION OF THE UNITED NATIONS ROME, 1971 DOCUMENTS OF THE FISHERY DOCUMENTS DE LA DIVISION DES DOCUMENTOS DE LA DIRECCION RESOURCES DIVISIONOF FAO RESSOURCES HALIEUTIQUES DU DE RECURSOS PESQUEROS, DEL DEPARTMENT OF FISHERIES DEPARTEMENT DES PÊCHES DE DEPARTAMENTO DE PESCA DE LA LA FAO FAO

Documents whicharenotofficial Des documents qui ne figurent pas Esta Subdirección publica varias series FAO publications are issued in several parmi les publications officielles de la de documentos que no pueden conside- series.They are given a restricted distri- FAO sont publiés dans diverses séries, lIs rarse como publicaciones oficiales de la bution and this fact should be indicated font seulement l'objet d'une distribution FAO. Todosellostienendistribución ifthey arecited.Most of them are restreinte, aussi convient-il de le préciser limitada, circunstancia que debe indicarse prepared as working papers for meetings, lorsque ces documents sont cités. Il s'agit en el caso de ser citados. La mayoría de or are summaries of information for use le plus souvent de documents de travail los títulos que figuran en dichas series of member governments, organizations, préparés pour des réunions, ou de ré- son documentos de trabajo preparados and specialists concerned. sumés d'informationàl'intentiondes para reuniones o resúmenes de informa- gouvernements des pays membres, ainsi ción destinados a los estados miembros, que des organisations et spécialistes inté- organizaciones y especialistas inte- ressés. Ces séries sont les suivantes: resados.

FAO Fisheries Report FIR IS (No.) FAO Fisheries Circular FIR ¡C (No.) FAO Fisheries Synopsis FIR ¡S (No.)

Special groups of synopses are iden- Des catégories spéciales de synopses Grupos especiales de sinopsis se dis- tifìed by symbols followed by classi- sont identifiéesàl'aide de symboles tinguen con las siglas siguientes, seguidas fication numbers based on indexed code suivis des chiffres de classification basés por números de clasificación que se basan of"ASFA": sur le code d'indexation de la « ASFA»: en las claves de los índices de la «ASFA»:

SAST Data concerning certain species SAST Données sur certaines espèces et SAST Datos relativos a ciertas especies and fish stocks. populations de poissons. y poblaciones. MAST Information on methods and sub- MAST Renseignements sur des méthodes MAST Sinopsis sobre métodos y mate- jects. et des sujets. rias.

OT Oceanographic data. 01 Données océanographiques. 01 Sinopsis sobre oceanografía. IT Limnological data. IT Données limnologiques. IT Sinopsis sobre limnología. and et y CART Information concerning fisheries CART Renseignements sur les pêcheries CART Información sobre los recursos and resources of certain countries et les ressources de certains pays acuáticos vivos de algunos países and regions (FIDIS). et régions (FID/S). y regiones (FID/S).

FAO Fisheries Technical Paper FIRT/T (No.)

Special groups of TechnicalPapers Des catégories spéciales de documents Grupos especiales de documentos téc- are identified by: techniques sont identifiées à l'aide des nicos se identifican por las siglas siguien- symboles suivants: tes:

RE Indexed lists of experts and insti- RE Listes indexées d'experts et insti- RE Listas índices de expertos y de tutionsdrawn fromRegisters tutions tirées des registres tenus instituciones tomadas de los re- maintained by the FisheryRe- à jour par la Division des ressour- gistros que se llevan en la Direc- sources Division. ces halieutiques. ción de Recursos Pesqueros. CB Lists of periodicals, special sec- CB Listes de périodiques, des sections CB Listas de periódicos, secciones es- tions of " Aquatic Sciences and spéciales de la «Aquatic Sciences peciales de la «Aquatic Sciences Fisheries Abstracts (ASFA), and Fisheries Abstracts (AS FA) », and Fisheries Abstracts (ASFA) », specialbibliographies and pa- des bibliographies particulières et bibliografías especiales y trabajos pers concerning documentation des articles sur les problèmes de relativos a losproblemasde problems. documentation. documentación. MFS Provisionaleditionsof" FAO M FS Editions provisoires des e Manuels MFS Edicionesprovisionalesdeos Manuals inFisheries Science." FAO de science halieutique ». e Manuales de la FAO de Ciencias Pesqueras ».

Some documents also have another Certainsdocumentsportentparfois Algunos documentos tienen también identification, if, for example, they have d'autresnumérosd'identification,par otra identificaciónsi, por ejemplo, son been contributed to a meeting for which exemple s'ils ont été préparés pour une contribuciones a unareunión cuyos papers have been numbered according réunion dontles documents ont été documentos hansidomarcados con to another system. marqués àl'aide d'un autre système. arreglo a otros sistemas. FAO Fisheries Synopsis No. 84 FIRM/S84 (Distribution restricted) SÄST - Haddock - 1,48(04),O1OQO1

SYNOPSIS OF BIOLOGICAL DATA ON HADDOCK

Melano9rammus aeglefinus (Linnaeus) 1758

Prepared by

R.W. Blacker Fisheries Laboratory Lowestof t Engi and

FOOD AND AGRICULTURE ORGANIZATION OF THE UNITED NATIONS Rome, December 1971 PREPARATION OF THIS SYNOPSIS

The present synopsis uns prepared as a follow-Up of the recommendation of ICES at its 54th Statutory Meeting (Copenhagen 1966) that FAO should extend its Species Synopses Series to cover sorne of the species of major commercial importance from Arctic and Sub- Arctic regions.

The details set out in this paper are based on data collected by the author in the course of his personnl research work on the species and also, on information received from various sources0 most of which are listed in the bibliography.

Distribution Bibliographic Reference

FAO Department of Fisheries Blacker, R,W. (1971) FAO Regional Fisheries Officers FAO Fish.Synop., (84):pag.var. Regional Fisheries Councils and Synopsis 0.1 biological data on haddock Commissions Melanograinmus aeglefinus (Linnaeus) 1758 Selector SM Author Nomenclature, taxonomy, morphology, distribution, bionomics, life history, nutrition and growth, behaviour, population dynamics, exploitation, fishing gear, fishing areas, protection and management, AME, ANW. FIRM/S84 Haddock CONTENTS

Pag e

1 IDENTITY 1:1

1.1 Nomenclature i

1,11 Valid name 1,12 Objective synonymy

1,2 Taxonomy

1.21 Affinities

1.22 Taxonomic status 1

1,23 Subspecies 1

1,24 Standard common names, vernacular names 1

1.3 Morpholojy 3

1,31 External morphology 3 1.32 Cytomorphology 3 1,33 Protein specificity 3

2 DISTRIBUTION 2:1

2,1 Total area 1

2,2 Differential distribution 1

2.21 Spawn, larvae and juveniles 2.22 Adults

2.3 Determinants of distribution changes 3

2,4 Hybridization *

3 BIONOMICS AND LIFE HISTORY 3:1

3.1 Reproduction 1

3,11 Sexuality 1

3.12 Maturity 1

3,13 Mating 1

3,14 Fertilization 1

3,15 Gonads 1 3.16 Spawning 3 3,17 Spawn 3

3.2 Pre..adult phase 4

3.21 Embryonic phase 4 3.22 Larvae phase 4 3.23 Adolescent phase 4

3.3 Adult phase 3:7

3.31 Longevity 7 3.32 Hardiness 7 3.33 Competitors 7 3.34 Predators 7 3,35 Parasites diseases, injuries and abnormalities 7 iv FIRM S84 Haddock

3.4 Nutrition and growth 3:8

3.41 Feeding 8 3.42 Food 8 3,43 Growth rate 9 3.44 Metabolism 10

3.5 Behaviour 10

3,51 Migrations and local movements 10 3.52 Schooling 10 3.53 Responses to stimuli 10

4 POPULATION 4:1

4.1 Structure 1

4,11 Sex ratio 1

4,12 Age composition i 4.13Size composition 3

4,2 Abundance and density 3

4.21 Average abundance 3 4.22 Changes in abundance 3 4.23Average density 3 4.24Changes in density 3

4,3 Natality and recruitment 6

4,31 Reproduction rates 6 4.32 Factors affecting reproduction 6 4,33 Recruitment 7

4.4Mortality and morbidity 7

4.41 Mortality rates 7 4.42 Factors causing or affecting mortality 7 4,43 Factors affecting morbidity 9 4,44 Relation of morbidity to mortality rates 9

4.5 Dynamics of population (as a whole) 9

4.6 The population in the community and the ecosystem 9

5 EXPLOITATION 5:1

5,1 Fishing equipment i

5,11 Gears 1 5.12 Boats 1

5.2 Fishing areas 1

5,21 General geographic distribution 1 5.22 Geographic ranges 1 5.23 Depth ranges i 5.24 Conditions of the grounds i FIJM s84 Haddock

Page

5,3 Fishing Seasons 5:1

5.31 General pattern of seasons 5.32 Dates of beginning1 peak arid end of seasons* 5.33 Variation in date or' duration of season*

5.4 Fishing operations and results i

5,41 Effort arid intensity 1 5,42 Selectivity 2 5,43 Catches 2

6 PROTECTION AND MANAGEMENT 6:1

6.1 ulatoryJleislative measures i

6.11 Limitation or reduction of total catch 6.12 Protection of portions of population

6.2 Control or alteration of physical features of the environment *

6.3 Control or alteration of chemical features of the environment *

6,4 Control or alteration of the biological features of the environment*

6.5 Artificial stocking

7 POND FISH CULTURE*

8 REFERENCES 8:1

* As no information was available to the author1 these items have been omitted from the text,

FIHM S8Haddock 111 i IDB.NPIPY - Speoif io 1.1Nomenclature L1elanojrammua ne lefinu(Linnaeua) 1758 (Fig0 i) 1,11Valid name Type Gadus ae lefinus L., 1758 Melanogrammue aeglefinus (Linnaeua) 1758 I D 15-16, II D 19-21, III D 19-22, I A 23-24, II A 22-23, gill rakera 24-27, 1,12Objective synonymy vertebrae 52-57.First dorsal fin high and sharp tipped,Jaw bones greatly shortened Gadus aeglefi.nua L, 1758 length of the upper jaw 28.6-33,1, lower jaw Morhua aeglefinus Fleming, 1828 35.3-39.1 percent of head length,In percent Morhua punctatuo Plemíng 1828 of body length:anteanal distance 41.7-45.5, Ae lefinus lirinei Malmgren, 1877 antedorsal distance 26.1-28.2, length of pectoral fin 14,1-16,6, length of pelvic fin 1,2Taxonomy 8.9-10.9, base of first dorsal fin 12.2-13.8, base of second dorsal fin 19,7-23.5, third 1,21Affinities 13.6-16.4, base of first anal fin 21.1-22.8, base of second anal 13.8-16.5, height of first - Suprageneric dorsal fin 13.4-18,5, height of first anal fin 9,6-10,6, length of caudal pethmole 4,3-5.3, Phylum Vertebrata its depth 10.4-12.4, head length 24.6-25.7. Subphylum Craniata In percent of head length:length of snout Superclass Geathostomata 34.5-38.9, lateral diameter of eye 20.0-28.2, Series Pisces width of forehead 17,9-22,8.Upper part of Class Teleostomi body and head dark grey with violet shading, Subclass Actinopterygil sides lighter, belly and lower part of head Order Gacliformes milky white,Lateral lino black,A large Family Gadidae black spot under the lateral line in the Subfamily Gadinae region of the pectoral fins on each sida, Pectoral and dorsal fins same colour as the Generic doreuin,Maximum length about 1 m (ter Svetovidov, 1948) Neianog'ammus Gill, 1863. Type G-aclus ae1efinum L., 1758. Only one species. The generic description given by Svetovidev 1,22Taxonomie status (1948) is: Morpho-speoies "Dorsal and anal fins separated by inter.- spaces.First anal fin short, beginning beneath 1,23Subspecies the origin of the second dorsal fin or slightly behind it,Caudal fin slightly notched.Barbel None on chin weakly developed, shorter than the dia- meter of the eye,Mouth small, rear end of 1,24Standard common names, lower jaw beneath the centre of the eye, upper vernacular names (from ICES, jaw extending beyond the lower.Teeth in 1-2 1966) rows on the dentary, in several rows on the pre- maxillary, a group of a few teeth at the apex of English Haddock the vomer,Lateral line dark, in the shape of a Danish Kuller shallow arch, to -the beginning of the third Dutch Sohelvia dorsal fin, placed above the central axis of the Faroese Hi-sa body, extending without interruption to the end of Finnish Kolja the body,There are pores along the lateral line French Eglefin canal on the head:13 pores in the preopercular- German Schellfisch mandibular canal, 7 pores in -the infraorbital Icelandlo Ysa canal,i pore in the supraorbital canal and i NorwegianIyse, Kolje pore in the supraorbital commissure.Mucous Polish Plamiak, tupaoz cavity on the skull closed in front,Prootio Portuguese Arinca with a notch in front on the sides of the skull, Spanish Eglefino Crests over the foramen of the recurrent dorsal Swedish Koija branch of the facial nerve scarcely developed. Russian Piksha Supraorbital with a high crest.Bones of the pectoral girdle greatly thickened,The opercular process of the hyomandibnlar somewhat shorter than the lower process." 1:2 FIRN/S84 Haddock 13

1.3Morpholo 1.33 Protein specificity

1.31 External morphology (see 1.21 Individual differences in tranaferrin and Figure 1). types in haddock have been located, but as yet definite transferring type determinations bave I 32 Cytomorphology not been possible (Niller and Naevdal,1967), Intraspecific variations in hae3noglobin com- Chromosome numbers 2n = or 7 pairs 46, 6 ponents have been detected in haddock from the metaceritrio,16or 17 pairs telocontric (Purdom, Norwegian coast, but the full sii. ifioanoe of personal communication), these variations has not been determined (ller and Naevdal,1969),

FIRM S8Jddock 2j 2 DISTRIBUTION The larvae are uaually most concentrated between 10 and 40 n depth (Coiten, 1965). 2,1Total area (Fig. 2) Juveuiie The haddock isa continental shelf species of the north-east Atlantic (ARE) and north-west According to Svetovidov (1948) in the Atlantic (ANW), rents Sea O-group juveniles are pelagic until autumn cooling starts;then they take to the In the north-east Atlantic it ranges from bottom,In the following spring they become the northern part of the :ay of Biscay to the pelagic again and move inshore,In the next north of west Spitsbergen, including the Irish autumn they move offshore again and become Sea, the North Sea, the western part of the wholly bottom-living, Intheir third year they itic and the basics off the continental shelf start seasonal migrations, moving to the western - Porcupine Bank, 'Farm" area, Rockali Bank ents Sea when autumn cooling begins and back (Blacker, 1962 and 1963), Lousy Bank (Pawsey and o the eastern Barents Sea during the feeding Davis, 1924), Farce Bank and Farce Plateau. season,In these migrations, the largest fish In the Barents Sea it oocurs as far east as move furthest east (Maslov, 1939), Goose Bank and the shores of southern Novaya Zemlya;some dead ones have been found in Thompson (1929) gives the areas of dis- -the Kara Sea in recent years (Svetovidov, 1948). tribution of North Sea haddock at successive At Spitsbergen, juvenile haddock have been ages from hatching, taken as far north as 80°30'N (Berger, Popov and Oheremisina, 1962),In the southern part In the north-west Atlantic juveniles seem of its range the haddock a-t present makes only to occur on the sane grounds as the adult sporadic occurrences in the southern North Sea otock (Schuck, 1948;Colton, 1965), and the English Channel and it is doubtful if i-t occurs in northern Biscay nowadays. 2,22Adults The haddock is abundant at Iceland except Mainly bottom-living but known to be in the colder areas of the north and north-east pelagic at times (see 3,5), (Saomundsson, 1949), but at Greenland it is comparatively rare on the west side, and very Seasonal migrations are undertaken between rare on the east coast (Konstantinov, 1968). spawning and feeding grounds:from the shallower parts of the Barents Sea to the north- In the north-west Atlantic the haddock west arid north Norway coast for spawning,At ranges from Cape May (38°N) to the Strait of Iceland spawning is concentrated off the south- Belle Isle (52°N),It is nos-t abundant from west and west coasts and feeding grounds are off Cape Cod to the southern part of Grand Bank, these north-west, north, south and south-east Haddock are much iosabundantinthe Gulf of coasts, St. Lawrence, off the west coast of Newfoundland (Bigelow and Schroeder, 1953) and off the east In the north-west Atlantic migrations are coast of Newfoundland (Leim and Scott, 1966). not extensive,In the Gulf cf Maine there is a movement into shallower water in spring and 2,2Differential distribution summer and againinautumn,Those haddock withdraw from shallow water when the temperature 2,21 Spawn, larvae and juveniles risos above 10°C in summer and when it fails in winter (Bigelow and Schroeder, 1953;Coitos, - Spawning areas and seasons 1955;Needler1930;Schroeder, 1942; (see Table I) Vladykov, 1935), The eggs are pelagic, mainly freni O-20 in In the North Sea, adults spawn in deep (Coitos, 1965), water in the northern North Sea and migrate to shallower water to the west and south for Larvae feeding,Haddock probably also spawn off the Scottish north-west coast, in the Butt of Lewis Larvae are pelagic and late stage larvae region (R. Jones, personal communication). may perhaps be associated with jellyfish Largest fish migrate furthest (Thompson, 1929), (e,g, C-flea) (Svetovidov, 1948; Coitos and In years of high stock abundance movements are Temple, 19.1), greater;for example, the very abundant 1962 year-class spread into the southern North Sea arid the English Channel, where haddock do not usually occur, 600 40° 200 00 200 490

30° 600 80° 40° Fig. 2 The diatribtttion of 20° 0° g. a 58' ITaddbck 2t3

TABLE I

Haddock spawnin' areas and seasons

Stock Spawning area(a) Spawning season Peak season Authority

1. North Sea Northern North Sea March to mid-MayLate March to Saville (1959, 1965) and and shelf W of f irot half April N Scotland Orkney and Shetland Schmidt (1909) MAFF 2, Rockafl Iockall Bank Nay (unpub1ihed data) 30 Proe NW, N and NE of' Late February toEarly April Saville (1956) Faro e mid-May

4, renta SeaNW Norway, 6368°N* Late January to March to April ound (1970) June

5.Iceland SW and. W loeland May io June Saemundsscn (1949)

6, NW Atlantic (a) Georges Bank, February to May March toApril Bigelow and. Schroeder Browns Bank, (1953); Walford. (1938) coastal water Golton and. Templo (1961 of Gulf of Maine

Scotian Banks April Leim and Scott (.1966)

Grand Bank May to Juno June dder (1965)

* According to Svetovidov (1948) haddock also spawn off the Finmark and W, Nunnan coasts, but there appears to be no recent evidence of this.

2.3Determinants of distribution changes The usual depth rango is 10-450 in, Svetovidov (1948) gives the depth range as Like the cod. (Gaduz morhua) the haddock is 10-1 000 in, but the latter depth probably closely associated with the Gulf Stream and its refera to a record of Goode and. Bean (1895) branches where mixing with Arctic currents and which is open to doubt, other waters takes place in shallow depths. Sonina (1969) describes changes in the Generally haddock occur in water tempera- distribution and migrations of haddock in tures between i and 1000. the rents Sea from 1927 to 1965. She attributes these to variations in 'the hyd.ro- In the nc.-west ktlantic the salinity graphic conditions and a change of feeling range of haddock is from 315 to habita. (Bigolow and Schroeder., 1953), but in the Barents Sea the salinity range is higher, 34_ 35°/co (Svetovidov, 1948)

FIRM S84 Haddock :1

3 BIONOMICS AND LIFE HISTORY 3.14 Fertilization

3.1 Reproduction External.

3.11 Sexuality 3,15 Gonads

The sexes are separate. - Relation of gonad size and, egg nunbar to body length and weight Hormaphroditism may coeur as in other gadoids, and to age but from the absenoe of published reoords it must be very rare, The relation of fecundity to length is shown in Figure 3.The low values of Earll There are no external sexual charaoters, (1880) compared with those of Raitt(1933) and fodder (1965) suggest thatsome eggs may have 3.12 Maturity been released before his sampleswere obtained,, although Earll was aware that spawningof' The ages and sizes at first maturity for individuals was spread overa long period. various populations are given in Table II. The relation of fecundity toage is given 3.13 Mating in Table III,

Probably polygamous and promiscuous (seo also 3,16),

TABLE II Haddock size and age at first maturity

Stock Age (years) Size ( ) Authority

North Sea 3-5 23-29 Thompson (1924, 1929) Rockall about 30 Personal observation Farce 2- 27-42 Thompson (1924) Barents Sea 4-7 37-73 Personal observation; Sonina (1969) Norwegian 24 26-32 Personal observation coastal

Iceland 4-5 50-55 Saomundsson (1949); Thompson(1929a) NW Atlantic 3-6 36-54 Thompson (1939); Bigelow and Schroeder (1953); Georges Bank 23 30-50 Clark (1959)

TANLH III Relation of fecundity to age (thousands of eggs)

Age (years) Stock Authority 3 4 5 6 7 8 9 10 11 12 North 100 224 Sea* 31 159 278 Raitt (1933)

Grand. - 48-. 195 198 514 481 496 742 805 846 Hoclder (1965) Bank 154 298 298 550 514 1148

* Ages adjusted to 1 January birthday 3:2 FI.4»/884

C

o o X o0o

0 O o X o o o X

Oo

O X

Raitt.North Sea. o Hodder.6rand Cank. x Ecirit.Gulf ofiaine

20 30 I.0 50 60 70 0 90 Length (cm)

Fig. 3The fecundity/length relationship for sorne haddock P15WS84 Haddóck 3:

-Coefficient of fecundity - Relation of time of breeding to that of rdated. species For Nor-th Sea: F 0.2032 z (Raitt,1933) Usually three weeks or so later than cod in the same area, with an overlap between late- For Grand Bank: F 0.001391 z spawning cod and earlyspawning haddook (Hodder,1965) (Saville, 1956; Walford,1938; Marak and Colton,1962) -Number of eggs produced. by an individua]. Location and type of spawning ground - In a year For location see 2,21, See above. Type of ground: in the Northwest - In a lifetime Atlantic, spawning takes place over broken ground where sand, gravel, mud and rooks inter-. A haddock may survive to spawn in six or mingle (Bigelow and Schroeder,1953). more successive years after reaching maturity, in which case the total egg production would be -Variations and causes of in excess of 3 million. variations of spawning grounds

- Variation in fecundity with No information, health or condition Ratio and distribution of No information. sexes on spawning grounds

- Correlation between the number No information (see 411). of eggs produced and the nature of environment Nature of mating act

Hodder (1965) discusses the possible effects Courtship and mating have been observed of temperature on the fecundity of Grand Bank in the aquarium (Hawkins, Chapman and Symonds, haddock, 1967).A malo and female paired off and sperm and eggs were released with the two fish in 3,16Spawning close contact, belly to belly, swimming vertioally upwards, - Number of spawnings per year Variation in mating behaviour Only one spawning per year, but observations of spawning in an aquarium show that the eggs are No information. released at intervals over a period of about three weeks (Hawkins, Chapman and Symonds, 1967).Earll Nesting habits (1880)and Raitt (1933) had already suggested repetitive spawning. No nest produced,

- Spawning seasons Reproductive isolation

See Table I and2,21. None.

-Spawning time of day - Induction of spawning

No particular time of day from aquarium No information, observations (Hawkins, Chapman and Symonds,i967) 3.17Spawn Sequence of spawning of individuals in a population Eggs buoyant, transparent, without oil globule and. with a narrow perivitelline space. No information. Diameter 1.19-1.72 mm, In early stages indistinguishable from eggs of cod and witch - Factors influencing spawning (Glyp-tocephalus cynoglossus) (Bigelow and time Schroeder, 1953; Walford,1938).

Closely related to temperature changes on Georges Bank (Colton,1968). FIRL

3.2 Pro-adult phase - Effect of environment on rates of development and, survival 321 Embryonic phase The variation in survival of hasiiock

McIntosh and Masierman (1897) describe the brpods in relation to wind strength and direa-- develo ont of haddock eggs and larvae, but no tión and currents has been studied. by temperature details are given, According to them, Carru'thers, Lawford and Veley(1951), gastruiation, the formation of the notochord and Carruthers, Lawford, Veley and Parrish(1951), the appearance of the first pient spots take Chase (1955), Rae, K.M. (1957), Saville(1959, place on the fifth day; the blastopore closes 1965)and Walford(1938).Carruthers, Lawford, on the eleventh day; embryonic fin rays and eye Veloy and Parrish(1951)found a correlation pigment appear on the seventeenth day; and between year-class strength of North Sea hatching occurs after20days. haddock and wind direction and strength, poor year-classes during the period 1920 to1949 The rate of development varies with temperar- being associated with predominant south-.west ture, The rates given by different observers winds during the spawning season. Saville are shown in Table IV, The figures given by (1959, 1965)found no such correlation for the Dannevig(1895)and Walford. (1938) agree closely, period1952to 1957, and Gu].land (1965) pointed but those of Bigelow and Schroeder(1953)differ out the difficulties of attempting to correlate considerably from those of the other authors. the survival of young stages of fish with environmental conditions. 3,22Larvae phase 3.23Adolescent phase Post-larval and young stages are illustrated in Figure4. Development of the basic adult colouration is complete at a size of45-50mm. At this Newly-hatched larvae are3o5-4.1mm long stage the juveniles are still pelagio. (McIntosh and Nasterman,1897).The yolk-sac is absorbed in about ten days at5C when the larvae See2,21for distribution of juvenile reaohe a size of5.5mm (Bigelow and Schroeder, stages. 1953). -Rates and periods of develop- Haddock larvae are distinguished from those ment and survival and factors of cod, saithe (Pollachius virens) pollack affecting these (p. pollachius) and polar cod Boreqgadus salda) by the absence of media-lateral piauontation. The growth rates of various populations They are distinguished from those of other are shown in Figure5eThe adolescent phase gadoids without medio-latera]. piient by their may last from two years (Faroe and North Sea) pigmentation and the position of the anua. 'to seven years (Barents Sea) (see Table II). Schmidt (1905) gives keys for the identification of larvae of different sizes. 0-group, 1-group and occasionally larger haddock are preyed upon by cod, saithe, whiting The black shoulder patoh is readily seen in and other predatory fish (Bigelow and Schroeder, larvae of45-50mm. 1953; observations from MAFF routine research vessel samples), See also3,34,

TABLE IV

Hatching time (days) of haddock eggs at various temperatures(°c)

100 120 140 Authority 6° 8°

Dannevig 92/3 23 173/4 15Y2 13 10/4 83/4 (1895) 20Y2

Walford 21 18'V2 163/4 13Y2 io/4 (1938)

Bigelow and Schroeder 15 13 (1953) II

i :i

14

Fig. 4 Postlarval aM young stages of haddock (from Schmidt, 1905) 80

70

60

- - FaroeBarentsIceland Sea EL)40 50 - IC.N.A.F Sz North Sea (fast) - / ' ,"-4 Rockati - - IC.NÁF 4 x 10 , ,y_'d ._- North Sea (sL.L.._ _..- North Sea iones ami Jermyzi, Blacker,Rookall wipiiblished.; Jones,Paroe MAFFBarents market Sea, sanpling Iceland Henneimzth,Browns Bank Grossleinsto. (IcNiP and 4x) 1968; 1 I 2I 3 I 4I 5 J 1962; 6 I uniblished;data for Age (years) 7 I 1969, 8I 9 I 10 I 11 I 12 I 13 t 14 Data sources: Fig. 5 Growth curves Lor various haddock stocks McCracken, 1964; Georges Bank (IcNAF 5Z) 1962.Jensen .. Wise, Y1m4/584 Haddock

Effects of environment, sub- (1965a)also recordsd cod and, haddock hotb populations, density, on raies of feeding on spawning capelin on the South-east development and survival Shoal of the Grand Bank in July and AngLLst, bwt -the haddock contained a, greater percentage of Growth rates of different populations vary capelin eggs than the cod. At Iceland both (Fig. 5)and there are also variations within species sometimes feed on san-leels (Brown and, populations (Thompson, 1929; Jones, 1962).These Chong, 1946) and this also occurs in the North-- may be environmental effects. See also3.43. west Atlantic (Kohler and Fitzgerald,1969). In all these instances the extent of competi- -Differences from adults in tion is probably small because of the great diet, feediug methods, etc. abundance of the food organisuw at the time when cod and. haddock are both feeding on them. Basically -there is no difference between the diet and feeding methods of adolescent and 3.34Predators adult haddock as both feed mainly on benthos (see 3.42), but the smaller adolescents (o-. and Fish up to about30cm are eaten by cod., 1-group) feed on smaller species and speoimene coalfish (Bigelow and Schroeder,1953)and than the large adults. other carnivores,Larger fish are taken by seals (Rae, B.B., 1968), but man is the prin- 3.3 Adult phase cipal predator at this stage. 3.31Longevity 3.35 Parasites, diseases, injuries, and abnormalities - Average life expectancy - Parasites and diseases Varies with different populations. The following parasites of haddock are -MTimu1nage listed by Reichenbach-Klinke(1966),

22years in the Barents Sea (personal Protozoa: observation). yxoaoma aeglefini in cranial bones Nosema branchiale on gills years in the north-west Atlantic 14 obolus aeglefini (Bigelow and Schroeder, 1953). Eimerin in swimbladder Haemogregarina ae lefini in blood years at Iceland (from routine otolith 16 Leucocytozoon sp. in blood samples, Fisheries Laboratory, Lowestoft). Platyhelminthea: 11 years in the North Sea (Thompson,1923). Diclidophora gadi on gills 3.32Hardiness Bucephalopsisracilescens Lepiclapedon rachion Temp3rature range1-12°C,salinity range Derotrema abyssorum 31 .5 .34.5 /00in the North-west Atlantic, up to Cx7pt000'tyle lingua metacercaria in 'the 35,2°/ooin the North-east Atlantic (Bigelow and skin Schroeder,1953). Orillotia erinacea larvae (adu,lts ja sharks and. rays) Haddock r r'ly occur in water colder than Parabothrium bulbiferwn i C and. colder mperatures cause large Abothrium mortalities (Templeman,1965). In acuaria Botbriocephalus scor ii. haddock survived. at 14'-'20°C for severa], months (Bigelow and Schroeder,1953),but such hi Nema-toda: temperatures are not normal in areas where Cucullanus cirratus in gut haddock occur. Contracaecum clavatum in liver, gut, mesen-. C. aduncuin taries, stomach and 3.33Competitors abdominal wall Ascarophi s morrhuae in gut There is, perhaps, a certain amoun-t of competition between cod and haddock (Brown and Cheng,1946; Kohler and Fitzgerald,1969; Pet rova-Gririkevich,1944). Petrova.-Grinkevich found that competition between cod and haddock was greatest in spring and early summer in the western and central Barents Sea, when both fish were feeding on capelin or krill,Templeman 3:8 FIRM!- S 1 Haddock

Crustacea: Variation of feeding habitB Bomoloohus soleas on gills with availability, season, etc. Caligas curtus on skin and fins Lernaeocera lusci on gills Larvae and juveniles are planktonic feeders. ZL.branchialisJ11 on gills Epibrachiella imdica Adults feed less intensively in the 2 (i1r11r1ihi ) in mouth cavity and spawning season,Usually they are benthic C. brevicollis/ ) pharynx feeders, but at times haddock concentrate on spawning shoals of capelin, feeding on the fish To this list should be added:Tripartiella and eggs (Templeman, 1965a), on herring melanogrammi (Protozoa, Ciliata), found on the spawning grounds (Bowman, 1923), and on sand- gills and fins by Stein (1961), and Ascarophia eels (Bigelow and Schroeder 1953; Brown and filiformis (Nematoda) and Pyraxnicephalue Cheng, 1946). Sonina (19695 described long-. phocarum (Cestoda), recorded on haddock by tenu changes in the feeding habits of Barents Dogiel, Petrushevski arid. Polyanski (1958). Sea haddock: a change from feeding mainly on. benthos in the east and southeast Barents Few of these parasites have been studied Sea to feeding on euphausiids and capelin in detail in the haddock2 but the effects of further west. Irligley (1956) described food Lernaeocera obtusa have been described by habits on Georges Bank. Kabata (1958). Infection is by a planktonic stage from the intermediate phase in the lemon 3.42 Food sole (Microstomus).Adults reduce the weight of the host by 10 to 30 percent. - Types

- Injuries and. abnormalities Planktonic copepods during pelagic larva]. and juvenile stages (Olgilvie, 1938; Marak, Xanthochroism occurs occasionally,Apart 1960). Adults feed, on a vast variety of from the golden colour there appears to be no benthic animals, especially Crustacea, adverse effect on the fish, Polychaetes, Lamellibranch molluscs, and Echinoderms (Ophiuroids and Echinoids in 3,4Nutrition and. rowth particular) (Brown and Cheng, 1946; Ritchie, 1937; Bigelow and Schroeder, 1953; Hertling, 3.41 Feeding 1940 Kohier and Fitzgerald, 1969; Wigley, 19565. -. Time of day Fish eaten by haddock include herring, Unknown. capelin, sandeels, small mackerel, young eels and. silver hake (Bigelow and Schroeder, 1953). - Plane Cannibalism has also been recorded (Wigley, 1956). Capelin spawn and herring spawn are Anhero in the geographic range. also eaten (see 3.41).

- Manner - Volume of food eaten

The method of feeding has been observed in Ritchie (1937) gives the volume of each aquaria by Tseeb (1962, 1962b). Nouthfuls of of the constituent groups in the diet for the bottom are tIsievedt in the mouth; sorno various size groups of haddock from the North particles are passed out through the gills and Sea. Thompson (1939) gives the following others spat out of the mouth, thus reducing the average volumes of food per 100 stomachs: intake of mineral particles, Nova Scotian Banks590 cc (average haddock - Frequency length 49 cm) Newfoundland Banks 934 cc (average haddock Unknown, length 56 cm) North Sea 562 cc (average haddock length over 40 cm).

J)'Kabata (1958) separated L. oPltsa, found only on haddock, from L. branchialis, Kabata (1963) reviewed the systematics of Clavella spp. in European waters and concluded that the many species described are all ecotypes of Clavella adunca. FIRM S8i Kaddook 3.43 Growth raie - Relation of growth to feedi spawning, eto. - Relative and absolute growth patterns and rates The slow growth rate in the deep central North Sea and at Rookall is probably due to poor The growth rate varies according to area. feeding conditions. Curves given in Figure 5 show that growth is rapii at Iceland, at Faroe,itthe Barents Sea, Templeman and Squires (1956) showed that and on Georges Bank and slow in the deep central in the Grand Bank area the growth rates differ North Sea arid at Rockall.GrowthOrdiffcvent for males and females:males grow faster at populations in the North-west Atlantic is giva first and mature earlier than females, but after by Hennemuth, Grosslein and McCracken (1964). maturity is reached their growth slows down. Jones (1962) also showed that female haddock Andersson (1938) noted variations in the were significantly larger than males of the same growth rate of different year.classes of haddock age in the North Sea and at Faroe in 1950 and entering the Skagerrak. 1951. Jones (1962) illustrated considerable annual - Relation cf growth to popula- variations in growth in the North Sea and compared tion density the growth rates found by Thompson (1929) with sore recent data (Table V),He also found con- The growth rate of the extremely abundant siderable variations for Faroe haddock. North Sea 1962 yeaxi_.class was siiificantly lower than average in i-ta earlier years (Jones The following parameters of the Bertalanffy and Jexyn, 1968).This may have been caused by growth curve have been obtained: the great density of this yean-class. L00(om) K Beyer-ton and Holt (1957) and Raitt (1939) North Sea, slow 48.3 0.28 discussed the effects of haddock population growth area Jones density on growth rate for various areas in the North Sea, fast 58.1 0.24 (1962) North Sea and showed that the growth rate growth area decreases with increasing population in some Faroe 77.9 0.22 areas. ICNAF Subarea 3 55-60 022 (Grand Bank) An inverse relationship between the growth ICNAP Subarea 4 65-700.17-0.22 Beverton rate and population density of the Barents Sea (Scotian Shelf) (1965) haddock population has been described by SoriAna ICNI4F Subarea 5 73 0.28 (1965, 1967, 1969).She also showed a relatioiz (Georges Bank) øhip between growth rate and changes in feeding habits which have happened at the same -Lime as - Condition factors the present deorease in population density in the Barents Sea. 0.85-1.2 (R. Jones, personal communication; TTtler, 1969).

TALE V The variation in growth rate of North Sea haddock, shown as mean lengths in centimetres.(After Jones, 1962)

Slow growth area Age ast growth area (years) 1926-30 1936-39 1947-57 1926-30 1936-39 1947-57

0.75 14.5 15.3 15.7 17.0 16.9 17,8 1.75 21,4 23.5 23.9 24.2 25,8 27,1 2.75 25.9 29.4 29.4 29.4 31.9 34.6 3.75 29.3 34.0 33.9 33 T 36,4 4017 4.75 32.9 37.8 37.3 37.6 39,9 45.8 3:10 p1pJ41584Haddock -Food-growthrelations - Mixing between species No information. In the catches haddock often occur mixed with codandcoalfish, but there is no evidence 3.44 Metabolism to prove that they are not in separate shoals on the sea-bed. - Metabolic rates -Patterns of schools The following values have been obtained by Ty-tier (1969, and personal communication). No information.

-Standa:rd - Vertical movements

+A 10°C, 5,9±12 mg 02/kg/hour; at15°C, At times haddock are known to be pelagic 150 - 40 mg 02ftgiour. and well off the sea bed (Trout, 1962; Woodhead, 1964, 1965).In the North Sea they - Active have been caught and observed on television at 27-36 m where the depth was 128 m (Margette, Mean rate at aswimming speed.of twobody 1966). lengthsper secondiat 10°C, 268 -48 mg 02/kg/hour;at 15 C, 289 ± 66 mg 02/kg/hour. - Size, density and, behaviour of schools in relation to time -Endocrine systemsandhormones of day, geographic location, seasons oceanographical factors, No information. physiological conditions - Osmotic relations Diurnal vertical migrations take place in some areas (Woodhead, 1964, 1965;Colton, No information. 1965).Thereare seasonal variations in catchabilitywhichare presumably due to 3,5Behaviour changes in the behaviour of the fish, for example the change from "dark" to "daylight" 3.51 Migrationsandlocal movements fishing at Faroe in the spring. In mostareasthere are migrations to and - Aggregation (local concem- fromthe spawning grounds. These may be tration of several separate extensive as in the Barents Sea (Saetersdal, schools of some species or 1956, Sonina, 1969) and at Iceland (Saemundsson, various species) 1949; Thompson,1929a),orshort movements as in the North-west Atlantic (Bigelow andSchroeder, Noinformation. 1953; McCracken,1965). See also 2.22. 3.53Responses to stimuli Thompson(1929)described thedifferences in migration distances and direction with - Environmentalstimuli increasing age of haddock in the North Sea. - Mechanical (pressures, 3.52 Schooling currents, sound) Entent of schooling habits Haddock produce sounds (Hawhins and Cha*nan, 1966) so presumably theycanrespond At spawning timeandwhen feeding on to sound stimuli. concentrations of capelin or sandeels, haddock form dense schools, but Jones(1963)suggests - Chemical (cflfactory, gastator thatit is not normally a shoaling fish. salinity gradients) Composition of stocks by size, No information. age and sex -Thermal See 4.1 No information. - Mixing ofstocks - Optical No evidence of mixing ofstocksanywhere. No information. Fi

- Artificial stimuli Photographe taken with cameras mounted on the trawl show haddock swimming in the direo- Fishing gear components, model -tion of the tow in front of the trawl as well as gear when they are under the netting (Blacker, Observations on the behaviour of haddock unpublished), in the trawl have been made by Clark (1958) and Livingstone(1962).Haddook in the funnel of - Electrical the cod-end swam mainly in the direction of the tow, making some side-to-side movements with-. Haddock respond in the same way as other out touching the net at the sides.A small fish, swimming toward the anode In high percentage swam straight into the cod-end,The currents paralysis or death occurs (MoRae and majority of fish seen escaping from the ood-end French,1965) went through the meshes easily, but some escaped only after a struggle. -Variations in reactions ocourrin with , size, etc. Blax-ter and. Parrìsh(1966)desoribed tank experiments with nets etc. and found -that No information, haddock responded to visual stimuli in daylight and some undetermined stimuli at low light levels.

FIRM SBHaddock

4 PoPuLATI at first capture

4.1 Struotu.re This vrioe with the growth rate and the mesh size in use. In the North Sea, at Faro 4.11 Sex ratio and Georges :, haddock usually enter the fishery late in their second year; at Iceland Research vessel samples off the north-west the Barents Sea and off Iova Scotia the of Norway coast in 1963 gave a ratio of58percent entry into -the fishery is usuall' 3-4 years, males to 42 percent females; 95.4 percent of but a few fish are caught when 2Y2-3 years old, the fish were mature (MAFF, unpublished data), - Age at first maturity Acoording to Thompson (1924) in the over- all population the numbers of each sex are about See 3.12, Pable II equal, He found variations lxi the sex ratios on different grounds with females predominating - Maximum age in the shallower inshore areas and males in the offshoregrounds, bu-t this is probably not Twenty -two years recorded from the ;&rente cnerally true. Sea (personal observation),

4,12 Age composition - Density of actgroups

The age of haddock may be determined from Highly variable, In the North Sea the scales (Thompson, 1923; Jensen and Clarke 1962 year-class was the most abundant ever 1958; Jensen and. Wise, 1962) or otoliths recorded, whereas the 1956 year-class was -the (Saeteredal, 1953). Comparison of age deter- poorest recorded. since 1916 (when records minat ions from scales and otoliths shows that began). The results of annual surveys of the up to the age of eight years or so agreement northern North Sea made by Scottish research is good, but after -this the scalo ages become vessels are published. in Annales Biologiques, increasingly unreliable (Kohier and Clark, 1958; Saeteredal, 1958), The results of Russian surveys of the ìrents Sea for 0-, I-, II- and. III- age-group - Age composition of the haddock have been published annually since population as a whole 1959 by Bararionkova (1961, 1962, 1963), ranenkova and i-ranova (1964), Nizovtsev and. There is little information for the Baranova (1965) and Bara-uova (1966, 1967, populations as a whole because O- arid. I-group 1968, 1968a), These surveys and the inter-- haddock aro not sampled adequately by commercial national 0-group fish surveys of the rents gear, but data are available for the northern Sea (Drasund, 1970; ICES, 1969b) show wide North Sea from Scottish research vessel surveys variations in the density and distribution of made annually since 1919 (except for the war young haddock, years 1939-45) (Raitt, 1939; Jones 1962), arid for the north-ost Atlaritlo from U.S., Surveys of the distribution and abundanoe U.S.S.R., and Canadian research vessel surveys of young fish have also been made at Farce in made since 194.8 (Grosslein, 1969; Richter and. some years (Joensen, 1956, 1957, 1959), Vinogradov, 1969; Lux, Richter and. Grosslein, 1969), Results from surveys of the North Sea For the north-west Atlantic, da-ta on the since 1947 and of the Barents Sea sinoe 1958, abundance and age and size distribution for the have been published annually in Annales distribution for the following IGNAF subareas Biologiques (ICES, Copenha,gen). have been published:

- Age distribution of the Subareas 30 ad 3N(Gran ) Templeman and calch Hod.der (1965 Hodder (1966 Ita for various areas are given in Pable VI, Subarea 3Ps (st, Pierre Templeman and. Bank) Hodder(1965a) Ad.d.i-tional information for some stocks is given each year in Annales Biologiques (ICES, Subarea 4W (fleerald arid McCracken Copenhagen) and the ICNAP Sampling Yearbooks, Sable Island Banks) (1968) Age distributions of ca-tohes at Iceland from 1960 to 1966 are given in ICES, 1969, Subarea 4X (Browns and Hennemuth, Le Havre Banks) Grosslein and. - Variations with depth, etc. McCracken (1964) No data, Age distribution (nunbers per thousand fish) of ha&iock landings, tota], catch and effort for 1968 TABLE VI Total Total Subarea 2 3 390.64 5 Age (years) 6 120,07 8 9 10 11 older12 and tone)(metricoatci fishing hou-va' lia* i.ys Norway fished0VaI ooast Iceland. rents Sea 73,50.2 352,6 14.2 0.2 401,1 82.4 331,5376.4 82.8 42,391.138.6 357.7 19,3 117.8 20.950,0 14.6 2,46.3 2,11.11.2 2.61.40,8 1.91,11.7 20 842 9 628554 212 427 6984 762806 jJ IV Vb Northern1. North Searoe Trawl 13.3 1,6 446.9522.3 232,9 45,7 80.6 5.7 60,2 2,4 483.5 57,9 31.2 1.7 0,69,4 3.2 0.9 11 9075 578 9653 913 617 L" North1.2, Central North Sea TrawlSeine 18.117.5 426.5 226.9 42.0 4.5 0.7 222.7506.2 2,0 0,6 10 069 904 8810 175 615 5Z Georges2, Bank Seine 10.9 121.4531.7702,2 28.656.6 - 71,7 5.6 566,2 0.3 223.5137.8 27,2 0.7 17.9 0.1 29.2 - - - 2544512 864 133 598 9096'4J 4/ jJU.S. English landings/ Scottishlandings (IcNAP, (NAPF,landings 1970) unpublished, (IcEs data)Scottish landings (Jones and Jeriuyn 1970) 1969) FIRM ¡ Ha4dook z

Subarea 5Z (Georges ;nk) Colton (1955); 4.2Abundance and densi (of population) Honnesiuth (1969); Zukowaki (1968) 4.21 Average abundance

The bounderies and subareas of ICES In the north-east Arctic the estimated ICNAF and NEFC aro shown in Figure 6. available population of Aroto-Norwegian haddock has averaged 823 million during the period O-group surveys of Georges Bank and the 1949-1968, The highest figure was 1 859 million Gulf of Maine have been made in most years in1952 when the outstanding 1950 year-class since 1948 (Colton, 1955; Grosslein, 1969, entered the fishery. The lowest was 388 million 1969a; Graham, 1969), in 1968, after a succession of poor or moderate strength year-classes. These estimates do not 4,13Size composition include O- and I-group fish (Garrod, personal communication). - See Table VII for length composition of the catch for According to Hennemuth (1969) the available various areas. population of Georges Bank haddock from 1935- 1960 was 132 million, but in 1968 it was Wells (1968) gives factors for the con- reduced to 47 million. The estimated average version of standard, fork and total lengths number of recruits (age 2) for the period for haddock from St. Pierre Bank, ICNAF sub- 1935-1960 was 49 million. The numbers of 0- and area 3Ps. I-group haddock would each be greater than this figure, so the total population for the period Additional information for north-east 1935-1960 probably averaged about 300 million, Atlantic stocks is pubLiahod annually in ICES "Annales Biologiques", and for the north- 4,22Changes in abundance west Atlantic in ICNAF Sampling Tearbooke. See 3.22 for the effects of environmental - Size at first capture conditions on brood strength, the variation of which is one of the main causes of fluctuations Varies with the mesh regulations for in abundance of haddock in all areas, each area. The effects of food competition and pre- - Size at first maturity dation on the abundance of haddock are negligible compared with the effect of fishing, See 3.12. Hennemuth (1969) describes the decline in abundance of haddock on Georges Bank, Raitt - Maximum size (1939) and Parrish and Jones (1953) show the effect of fishing on North Sea haddock, and 112 cm, 14.5 kg gutted weight, from Gulland (1961) gives data for the Iceland stock, Iceland (Thompson, 1929a). Recent changes in the Barents Sea and Iceland stocks have also been studied (ICES, 1969, - Density of size groups 1969a).

Varies with the succosa of year-classes 4.23Average density but the actual density of any particular size group is difficult to determine for any - Annual mean density stock. See 4.2. See 4.24. - Length and weight relation- ship 4,24' Changes in density

See Table VIII. - Landings per unit of fishing effort

For the period 1917-1968, landings from Georges Bank (IGNAF subarea 5Z) varied from the peak of 22.6 metric tons per day's fishing in 1927 to 2.9 metric tons per day's fishingin 1968 (Hennemuth, 1969). aength distrilmtjon of gnglish haddoek lm1ings for 1968 (MAFF, unpthlished). TABLE VII Number of fish per 100 hours' fishing 30-4 6 35-9 40-4 45-9 50-4 Lh-group55-9 (an) 60-4 65-i 70-4 75-9 80-4 85-9 90-4 Total lia VaI IcelandLroEroxite Sea coast Length and weight relationship of haddock from various arean TABLE VIII shown as the weight (g) of fish of lengths 20-80 cm 58 2 209126 67 446734 665 1 o

1 938 1 640

073 2

5 857046539 71 928 048

419 1

4 040256093 942132300 219 7763 19 2771 4111 8 43 760 2 01 19 38 567 783 h (an) AreaGulf of )ine Airthorityh,rll (1880) 20 - 25 -- 30 35 - 40 - 45 (491 077 an) 55(561 815 an) 260 145(61 an) 65(662 950 an) 470 335(72 an) 75 1WNorth Atlantic Sea ClarkSchucklittRussell and(1933) (1947),(1914) 7267 198138134 242339238 508418388 586763629 1 073 843902 1 169440220 1 8921 570650 2 400200 - 2 965 - 3 700 - 4 490 - - North Sea M&1TDietach and. (1959) D (1969) - - 253 - 619 - 1 266 - 2 194 - 3 430 - 5 120 II 41 AZM4TAL £1141 *114 IIQJ(CTIDI NORTH ATLANTIC (IT(ICI IT 4I I. I$ I 'o. IV 1V 7V 1V (t 43. s Vila :\12 - 3r XII 5Zw : r' 7v5 / 3i 432! X 'g. 4 6 H .. Fig. 6 Areas and Subareas of ICES, ICNAF and NEAFC (from ICNAF Statistical Bulletin) FIPJ'V584 Haddock 'In the Barcnts Se. (ICES cua I) the 4.3Natalitand recruitmen catch per unit effort for Lng1ieb trauler3 during the period 946-1967 varicd from 170 4.31Reproduction rates to 970 tons per m±llion ton-hours0 For the Norway coast fishery (subarea lia) the - Annual e production figures for the same period were 180 to 7 900 tons per million ton-hours0The peak Prom egco and larvae surveys a-t roo, figures tiere for 1946 when fishing was reauined Saville1956) estimated egg production ae after the 1939-45 war0For the period 58 x 101in 1950, 70 x 10'' in 1951, 67 i 1011 1950-1967 the highest catch-ra es tiere .420 in 1952 and 49 x 1011in1953, tons per million ton-hours in theorenta Sea and 1 100 tons per million ton-hours off the Walford (1938) estimated the number o± Norway coast (Icii, 1969a), ec produced by the spatjning stock on Georges Bank in 1932 as 80 x 1011, At Iceland (subarea Va) Ehglish trawler catches deulined from 757 tons per million Beverton and Holt (1957) estimated the ton-hours in 1946 to 74 tons per million mean annual egg production for North Sea haddock ton-hoursin1966 (iCES, 1969), as 37,5 x iO1for the period 1922-1937, the range being from 7.9 x 1012 to 65,5 x 1012, Haddock catch-rates in the northern North Sea during the period 1947-1967 varied - Survival rates from about 3,81 tons per 100 hours' fishing to a peak of 14,07 tons por 100 hoursin1966, Saville (1956) found no clear relationship These are the catching rates of Aberdeen between the number of eggs produced and, the trawlers fishing the most important dis- number of surviving larvae atroe,Beyer-ton tributional area of the haddock in the North and Holt (1957) also showed that there is no Sea,Catch-rates in the middle North Sea direct relation between the number of eggs are very much lowers produced and the subsequent annual recruit- ment to the stock,Hervingtcn (1944) stated At Faroe the British catch per unit effort that large spawning stocks usually result in since 1947 has varied from 780 tone per poor year-classes, because the survival of million ton-hours (1947) to 250 tons per larvae from large broods is reduced by com- million ton-hours (1961) (B,W. Jones, personal petition for food. communication), - Forecasting of potential - Variations with depth yields Haddock under-bake seasonal migrations in Indices of year-class strength have been most areasusually from deep water in winter obtained from surveys of 0-grpup fish in the to shallow feeding grounds in summer (McCracken, north-east Arctic and in the north-west 1965;Temploman and. Hodder, 1965b;Schuck, Atlantic (see 4.12 - Density of age-groups), 1952), but there may be local variations In Forecasts of yield have been based on these in the pattern of movement, as in the Gulf of recent years for New England waters (Graham, Haine (see 2,22), 1967), - Seasonal variations in 4.32 ]ctors affecting reproduction araïlable stock - Density dependent factors Trout (196a) described the variationsin Sonina (1965, 1967) and others (see 3043) catches associated with the seasonal migration have shown that growth of haddock may be patterns, density-dependent.As fecundity is related to In the northern North Sea and in other the size of the fish, the potential reproduc- areas the available stock increases toward the tion rate is obviously affected by any factor end of the year and the end of the growth which reduces the growth of the fish, season, when the recruit year-class usually Predation is no doubt a cause of larval reaches saleable size (see 3,43). mortality;the extent is unknown, Schuck (1949) describes seasonal changes in relation to fishing in New England waters. - Physical factors iluro of year-classes in some areas has been attributed to water movements caused by wind carrying eggs and larvae into unfavourable areas (see 3.22), FIRM/884 Haddock Rollefeen (1930) showed that rough - Relation of recruitment to be directly responsible for the stock size and reouctive ra. and death of cggs of cod.Presumably haddock would be sw300ptible in the same way. See 4.31. 4.33Recruitment 4.4 1r11t.r and morbidity - Mean recruitment to the 4.41Mortality rates fishable stock - Annual mean mortality rates For the North Sea the mean annual recruit- ment for te period. 1922-1937 was calculated an For all areas the annual mortality rate 849,5x 100 (Beverton and :.1t, 1957). lies between 40 and 75 percent (see Table IX for authorities), Hennemuth (1969) calculated the mean annual recruitment of 2year-old fish on - Instantaneous mortality rgeo Bank as49x 10° for the period coefficients 1935-1963. See Table IX. - Footors determining recruitment - Variations in mortality The age at which hadcìook attain market- st of the authorities quoted in Table IX able size is probably the main factor in show the year-to-year variations in mortality determining recruitment to the fishery (see coefficients, and the variation of mortality 4.12), rite with age. - Seasonal pattern of McCrcken (1968) also shows seasonal recruitment es in the mortality rate of haddock of the central Scot inn Shelf, In the North Sea recruits usually enter the fishery in the autumn and winter of their Jon (1966) gives a ranof total second year (age 1Y2 years), but before that mortality coefficients in the North Sea1 from large quantities of the pro-recruitment s 0.8 to 1.0inthe northernmost part to 1.5 to of abundant year-classes may be caught and 1.6inthe heavily fished. arca off the Scottish rejected..ThisiBalso true of Georges Bank east coast, haddock (Bigelow and Schroeder, 1953; Graham, 1952) and probably applies to other - Density dependence of stocks as well. natural mortality - Variation in annual No information. recruitment 4,42Dhctors causing or affecting For the North Sea during the period mortality 1922-1937 the calculated numera of annual re9ruits varied from 39 x 10to 2 807 x - Predators 100 (Beverton rd Holt, 1957). There are no data on the extent of Estimated aual recruitment of mortality cauaed by predation, but larvae and 2-year-old ad1ook on Geges Bank varied fish up to 30 cm long are eaten lr a variety from I x 10to 502 x 10for the 1956-68 of predn.tors (eee 3.34). year-classes.The decline in recruitment has been most marked since 1965.The - Food of larvae and post-la estimated. recruitment as 2-year-olde from the and its availability 1965-68year-classes is: 1965, 2 i 106; 1966, 16 z 10'; 1967, 1 i io6;1968, 16XiO6.This Food availability may be one of the is to be compar9d with the mean annual recruit- canses of tho variation in year-clacu st h ment of49i 10° for the period 1935-63 (Herrington, 19ì;Suville, 1956) (uee 431) (Hennemuth, 1969). - Physical factors Estimates of total numbers of recruits are not available for other areas or periods, See 4.32. but indications of the variation in year.- class strength can be obtained from a number of sources (see 4.12), Estinates of rtality ooeffioients tiy ooiiot74.hthg (x, z) for £uthori7 rioua (1966)and cnd Jones (1953 ii (1957 Iorh Saa 19221939 .ortali*- Total Di*lit7 - Jc31ft.rith B.W, zt (perooiml j.j (1953)ir-iah and Jonc'i (1953) 3zc IC (1969s)(1969) 1947-1950 H:-rJi,3Hme.u-th Groaslein (1969)Ci_-_cen (1964); NrCracken (1966) and(1968) * br age't3 2-9 age-groupa 69 1958-1951947'1965 1.0 1020 1.02 o (ezaiu2ing1926-1950 1%l-: 0.15 0.59-1.41 0.'4-11201000 .56 1951-19681941-19G 0.18 'r 0.21-0.45 Ix 0.32-1.280. .63 4 Io.1*nd .ck oIou 1964-19651956-19631960-1966 0,200.200,300.15 0.65**0.03-0.98'0,. 0.33-1. 1.s" 0.85**0.19-1.17' 4 0.80 1.00 6.5o (4W,Bootian(5z) 4X) .1f 1948-19651965-19661935-1963 0,20020 0.800.50 0,.s1.000,70 a 73 (3)Grand nk 1955-1964 0,75-1.00 FIRMJS84Haddock

- Direct effects of fishing 4,6The o ule,tion in the coinmunit and the ecos sem Pishing is the main causò of mortality (see Table IX). - Physical features of the biotope of the community - Indirect effects of fishing The haddock is normally a bottom-living No information. fish at moderate depths in areas where the predominant hydrographic feature is -the 4.43Pactora affecting morbidity mixing of Atlantic water wi-th another water ss e.g. Arctic water in -the rents Sea, - Parasites and diseases - Species composition of the See 3.35. community

4.44 Relation of morbidity to A wide variety of other fishes are caught mortality rates with haddocks but the relative sizes of their populations are unicnown, No Information. - Interrelations of the 45 Dynamics of population (as a whole) population of the species in the community and ecosystem Because the haddock is one of the moEt important demersal species in the North The haddock is mainly a benthos feeders Atlantic the dynamics of all thé major but sometimes feeds on pelagic fish or stocks bave been studied. in recent years. plankton in midwater (see 341) Adult fish Beverton and Holt (1957) used pre-war data on are the final link in the marine chain: the North Sea haddock to demonstrate methods of population assessment. Parrish and Jones bent hic plank-tonic Pia (1953) describe -the methods they applied to flio_it the North Sea stock (1946-50) and to the Parse stock (1914-50). fish

Recent assessments of the Arcto-Norwegian Juvenile stages are often the prey of carni- stock ICES, 1968, 1969a), of the Iceland vorous fishes such ascod0 stock ICES, 1969), and of the Georges Bank stock Hennemuth, 1969) have used the method Type of fluctuations (cyclic of virtual population analysis described by - and non-cyclic) Gu].land in ICES (1965). An earlier assess- ment of the Iceland stock, showing the effect There aro no data which suggest cyclic of changes in mesh size and fishing effort, fluctuations of haddock populations, but was made by GullanzI (1961). there are great variations in population density, the reasons for which are not fully The method of virtual population analysis understood (see 4,22 and 4.24), reduces the variance of mortality estimates compared with those estimates obtained from - Changes in environmental the ratio of the catches per unit effort of factors and their effect on the saine year--class in successive years, the population because it is based on -the numbers of fish of each year-class and furthermore it does See 2.B, 3.22 and 4,32, not assume that -the fishing mortality is constant with age.

FIRM S Haddock

5 EXPLOITATION Differential abundance associated with hydrographic 5.1 Fishing equipment features

5.11 Gears Atlantic subarctic regions: 5.1,2.Barents Sea and North Atlantic intermediate boreal - Present gear regions; 5.2.1 Newfoundland waters; 5.2,2 Irminger Gyral; 5.2.3 Norwegian Sea and Farces; Haddock are caught by a wide variety of 5.2.4 North Sea and Irish Sea, Limited to gears; the most important is probably the otter areas influenced by the Gulf Stream, trawl, but seine nets, lines and traps are also used, and midwater -trawis take haddock on 5,23 Depth ranges occasion. - Bathymetric contour - Changes in types of gear during the development of the fishery Rare waters shallower than 10.-iB m (Bigelow and Schroeder, 1953) and most Changes refLect the growth of otter trawling fisheries take place in lesa than 100-360 n, in the twentieth century, Variations of d.enaity with - Use of echo sounding or fish depth detectors No information, Haddock give good echoes, but these are indistinguishable from those of cod. and many 5.24 Conditions of the grounds other species with the gear and techniques at present available. Probably feeding grounds are mainly s or sandy mud with a rich fauna of molluscs and 5.12 Boats echinoderms (Blacker, unpublished photographs),

- Type 5,3 Fishing seasons

Otter trawlers, seine-.netters, small and 5.31 General pattern of seasons large lineboats are used, depending on the geographic area of the fishing grounds. Fished at all seasons of the year,

- Size, power 5.4Fishing operations and results

All sizes from rowing boats to the largest 5.41Effort and intensity factory trawlers. In most areas haddock and cod. occur 5.2 nars together and it is impossible -to give separate estimates of effort for each species. 5.21 General geographic distribution - Units of effort Fished throughout its geographic distri- bution (see 2,'! and 2.2) apart from the extreme Catch per 100 hours, catoh per 10 hours, limits of larval distribution, catch per day' s fishing, catch per day's absence. 5.22 Geographic ranges -'Landings per unit of fishing - Distance from coast effort

Rarely caught close to coast (see 5,23) Georges Bank except in deep fjords as at Farce, and in some 2.9-22.6 metric tons per day's Henneinuth locha on the Scottish west coast. fishing (1969) Barents Sea/Norway Coast MATT and - Areas of greatest abundance Barents Sea: 11.1 tons DAFS Norway Coast: 29.4 tons ÂNE, Atlantic NE: Barents Sea, Norwegian (1969) - per 100 hours' fishing, Sea, North Sea and off Iceland. English landings, 1968 ANW, Atlantic NW: Georges Bank and Browns Bank. 5:2 FIRM s:Haddock Iceland selectivity (Blacker, 1967;Bohl, 1967; 4.5 tons per loo hours' MAFF and DAFS Hylen, 1967;Olsen, 1967) fishing, English landings (1969) - Changes in the mesh size and Northern North Sea their effects 9,6 tons per loo hours' MAFF and DAFS fishing, Scottish trawlers (1969) Gulland (1961) and ICES (1969) discuss the 6.0 tons per 100 hours' MAFF and DM'S effects of increasing mesh sizes for the Iceland fishing, Scottish seine- (1969) stock.The effects of mesh changes on the yield nattera of the Arcto-Norweg'iaxi haddock have also been estimated. (ICES, 1969a). Fishing effort per unit area, and total fishing intensity Graham (1952) and Beverton and Hodder (1962) estimate the effects of mesh changes in Jhere cod, and haddock ranges overlap, ICNA? subareas 3, 4 and 5. separate estimates of effort for each species are unobtainable.Data on fishing effort for - Other selection fac-tors some areas and some countries are given in Table VI,Further data on total fishing effort In some areas and some seasons there may be are available from Bulletins Statistiques considerable rejection o± fish just above the (ICEs, Copenhagen) and ICNAF Statistical Bul- lega]. minimum sizer because of difficulty in letins,No figures are available for fishing selling them.This is particularly likely in effort per unit area. the North Sea when a very abundant year-class first enters the fishery, - Causes of variation in fishing effort and intensity 5.43 Catches Fishing effort in particular areas varies - Total annual yields from with season, weather and economic factors. different fishing grounds For example, heavy fishing on Norway coast grounds may result in diversion of trawlers Annual yields from the major haddock from Iceland and the North-west Atlantic.Poor fishing areas are given in Table X.Figure yields in the Barents Sea in some years have 7 illustrates the yield from the more important resulted in increased effort by British and areas. Soviet vessels in the North-west Atlantic, (Data from ICES Bulletin Statistique, Prolonged spells of bad weather and ice ICNAF Statistical Bulletin,) conditions may close some grounds such as those off North-west and, North Ice1and The total catch of haddock from all areas, in metric tons, live weight for some recent 5.42 Selectivity years Was:

- Selective properties of gear 1962 1963 1964 Jb 5 00 5Tò0 5öO Selection factors for haddock: FAO (1967, 1965 i2.. 1967 1968) Double-braided 3,0-4.1Blacker (1967); 747 000 729 000 4ò0 manila ood end Pope and Hall (1966); Maximum equilibrium yield Bohl (1966); McCracken (1963); Parrish and Jones (1953) gave estimates Clark (1963) of the maximum yield for North Sea and Faroe Double-braided 3,234 Hylen (1967); haddock for 'two levels of natural mortality, olyproprlene Olsen (1966, 1967) Assuming an instantaneous natural mortality (ulstron) coefficient (M) of 0.11, the maximum yields per recniit are 0.31 kg for the North Sea and Double-braided 3.6 corn- Treshev and 0.73 kg for Faroe.Assuming M = 0.22 the cor-- polyamide pared Stepanov (1968) responding yields are 0,21 kg and 0.37 kg. (Kapron) with These yields could be attained with an effort 3.1 for of about 75 percent of the 1950 level, when the manila minimum -trawl mesh size was 70 nm.Increases in the minimum mesh size would give an increased Experiments with topside chafers show that yield without the same reduction in effort. the approved ICNAF chafer (i-fr times the width of Since then the minimum mesh sizes for these and the cod-end), and ohafers with meshes about other areas have been increased (see 6.12), twice the cod-end mesh, have little effect on NWNorth Scotland Sea (IVa, (VIa) b, o) Rockall (VIb) Barents Sea (i) Norwegian Sea (lia)

IcelandFaroe Plateau (Va) ana BankCm1, m)

ICNAFICNIF SubareaSubarea 543 NorthNW ScotlandSea (IVa, (VIa) b, o) Haddock landings from the major fishing areas (1938-1968) in metric tons, round fresh BarentsRockall Sea (VIb) (I) TABLE X IcelandFarceNorwegian Plateau (Va) Sea and (lia) Bank(Yb1, Vb2) 105 304 8630 986053 8 96 7827191 112394 5 581879 165670734 12 66 424 762024 995 41368 702329 871391152 632026 415699239 ICN2ICNAFICNAF SubareaSubarea Subarea 543 544768 086727657 4448 689908337 40 5335711 040400 67 238 52 419 Dataand for ICESICNAF subareas subareas from from Cons0 mt, Commnperm0 NWjut, 243Atlant0 for 1967 and 1968 data, except for Regions plo 121 1602614 719302 2 159 7282510 642 8451 547 8 431 2 301 8625 427727 20 709 159 1722725 149189 4667 000 000 46 000 i08 3444779 000 000 52 000 119 60335 145 44 046 59 398 59 114 123 3562113 791471 3 073 102 56214 431 2070 076 135 51 337 198 706 125 68122 096 1 375 7926i8 835056 9932 416 59 635 599912 996247 353 221 70032 467 268 958 29 881 27 569 19 491 981819 505 407i08 548 123 438 35 417 1 935 69 539 154 7258599 240127 8 612 87 656 r0 Fish0I arid Res,Iii whichDoc, are obtainedTer Bulletins from ICES, Statitiaues 69/21 and. Statistica]. Bulletins. 1938-1966; 1969. and FAO 18 762 23 164 15 403 157 098 1634 573693 134 40 93517 515 93 917 126 978 6660 .1410929 854 167 40820 302 104 00530 668 874 28 469 49601213 482000 000372 163 720 57 000

139 46920 469 903 Subarea 13711938 076482 1561945-6 759 2 802 931947 777 7 548 7i1948 068 7 329 55 1949231 4 839 561950 4292 867 561951 478 3 975 521952 372 8 039 18601953 313380 1954 1955 171956 857 147393261 161972 3.787 265916356 799166 2 350 941253136 329 258 3 903 100793779 785423 1 792 115 57424 95398051 840 113090 983010517 1 959 144092786 735758 3 257 103 6622014271 334 1 641 105 4161501215 605 1 402 24 948 823 2642 719122 - 2332 710984 - 5751 905035 7866 511537 - 6160 741373 - 42532354 325 351360248 48532744 659883952433 445337 37929564i 583 50535562 295 056539335 104 471504364 341525100 58518061 451436444898 Subarea 1957 1958 1959 1960 1961 1962 1963 1964 1965 1966 1967 1968 4551 948 227 6 393 Fisheries Circulare Nos, 44 066 211 300 North Sea ,K' \ 1' - _. NorwegianBarents Sea Sea y -- ;L::,Á S.-- Ø- /-o.-o,. 'b- . - Farces \ Iceland -o 1938 195-4.6 fro-- 195Q V_ò- 1960 1965 1968 Fige 7 Haddcck isc1.ings fm -the main fiMng areas for the 1938 to 1968 (see elso Table X) î95 eriod s. FIRM/S84 Ha.dook 5 5

For the Arcto-Norwegian stock a yield of Hennemuth (1969) calculated a maximum sus- 0.578 kg per recruit should be attainable at tainable yield of about 50 000 metric tons per 33 percent of the present fishing level, which year for the Georges Bank stock ai a level of yields 0.456 kg per recruit (ICES, 1969a). about 7 000 standard days9 fishing (approxi- mately 50 percent of the 1967 and 1968 fishing effort).

FIRM S Haddock 6:1

6 PROTECTION AID MANAGEMENT (b) 67°O0'W 42°20'N 67°O0'W 41°15'N For the purposes of regulation of the 65°40'W 41°2'N fisheries the North Atlantic is divided into two 65°40'W 42°0O'N (ICNAF, 1969, areas: the northeast Atlantic comes under the 66°00'w 42°20'N p 2728) NorthEast Atlantic Fisheries Commission (NEAFC) and. the north-west Atlantic is wider the jurie-. Closed. seasons diction of the International Commission for the Northwest Atlantic Fisheries (ICNAF). Fishing is prohibited in the above areas during March and April 1970, 1971 and 1972e 6,1 Regulatory (legislatç) measures (IcN', 1969, p. 27-28)

The problem of management of fish stocks, - Limitations on size or including haddock in the North Atlantic, is efficiency of gear or craft reviewed in ICNAF, 1968. Subsequently the regulations outlined below were brought into The mesh size of trawls, and other nets force in 1970 (ICNAF, 1969). used for catching demersal fish, is regulated in both theIGNAF andNEAFCareas, 6.11 Limitation or reduction of total catch The minimum mesh sizes are given in Table XI. Mesh sizes are measured from knot to knot - Limitation on the efficiency stretched., anti apply to both wet and dry nets, of fishing units In theICNAFarea and Region 1of the None to date, NEAFC area topside chafers are permitted on trawl codends, provided that their specifi- - Limitation on the tota]. catches cations and. attachments are in accord with one of the following: ICNAF regulations impose the following total catch limits each year during 1970, the chafer is rectangular, of a mesh size 1971 and 1972: not less than that of the codend, and is at least one and. a half times the width of the Division4Xof subarea4: 18 000 metric tons codend, It must be attached. to the codend only along the forward and lateral edges, so Subarea (which includes 5 12 000 metric tons that if there is a splitting strop the chafer Georges Bank): is attached. not more than four meshes forward and ends at least four meshes forward of the Details of the method. of reporting catches to codline mesh,If there is no splitting strop be used. by the Contracting Governments are also the chafer must not be more than one third the given(ICNAF,1969, p26-28), length of the codend; or 6.12 Protection of portions of flaptype chafers of the same mesh and. population width as the codend, attached only along the forward edge and not more than ten meshes Closed. areas long, The aggregate length of such pieces must not exceed twothirds the length of the ICNAFregulations prohibit fishing with codend.; or gear capable of catching demersal species during March and April 1970, 1971 and 1972 in the a rectangular chafer, the same material following areas: and width as the codend, but with mesh dimensions 'twice that of the codend, fastened That part of division 4X of subarea 4 that 1 only along the forward and lateral edges so lies between 42°00'N latitude and. 43000'N lati- that each mesh coincides with four meshes of tude and between 67°0O'W longitude and 64°30'W the codend, longitude. - Restrictions based on sex or 2, In areas of subarea bounded by straight 5 condition lines connecting the following coordinates in the order listed: None.

(a) 70°O0'W 42°10'N 69°10'W 41°10'N 68°30'W 41°35'N 69°20'W 42°30'N 6:2 Ii/S84ddock

TABLE XI

Minimum mesh sizes (St. Inst.1968, 1969)

Minimum mesh size, width of flat gauge (mm)

IGNAF Subareas EAFC area Net

1 2,3,4 Region 1* Faroe North Sea and.5 (subarea Vb) (subarea Iv)

1. Seine net 110 100 110 f05 70

2, Any trawl made of cotton, hemp, 120 105 120 105 75 polyamide fibres or polyester fibres

3e Any trawl made of manila, sisal or 130 114 130 110 80 any other materia]. not mentioned in 2

* NEAFC region i includes subareas I, lia, lib, Va, XII and XIV.

- Restrictions on use of fish In all countries within the NEAFC area there are lega]. minimum sizes for haddock; In industrial fisheries in the NEAFC area and the landing of' haddock less than 27 orn tota]. oertain fisheries in the ICNAF area, in which the length from near-water fisheries, or 31 cm use of small-meshed nets is legal, the catch of from distant-water fisheries, is illegal. haddock must not exceed 10 percent by weight of the total catch. FIRN/584 Haddock 8:1

8 REFERENCES

Anderson1 K.A., An investigation into the alterations in the growth-rate of the haddock. Rapp. 1938 P.-v,Réun.Cons.perm.int.Explor.Mer, 108(1):85-7

Baranenkova, A.S., Soviet investigations on small haddock of age-groups 0-III, Annls biol., 1961 Copenh., 16(1959):148-9

Soviet investigations on small haddock in the Barents Sea. Annls biol., Copenh,, 1962 17(1960):145-6

Soviet investigations on young haddock in the Barents Sea. Annls biol., Copenh., 1963 18(1961):107-1O

Baranenkova, A.S. and S.F. Baranova,Soviet investigations on young haddock of age groups O, I, 1964 II and III in the Barents Sea. Annls biol., Copenh., 19(1962):88-9

Baranova, Z.P., Soviet investigations on young haddock (O-3 age-groups) in the Barents Sea. 1966 Annis biol., Copenh., 21(1964):91-2

Soviet investigations on young haddock of the O, I,II and III age-groups in the 1967 Barents Sea in 1964/65. Annls biol., Copenh., 22(1965):92-4

Soviet investigationson young haddock of the O, I, II and III age-groups in the 1968 Barents Sea. Annls biol. Copenh., 23(1966):108-9

Soviet investigationson young haddock of the O, I, II and III age-groups in the 1968a Barents Sea. Jnnls biol,, Copenh., 24(1967):102-3

Berger1 T.S., B.A. Popov and R.A. Cheremisina, New data on the northern limit of occurrence of 1962 young cod, haddock, Norway haddock and herring in the Barents Sea. Dokl.Akad.Nauk, SSSR (Eng. Transl,biol.Sci,Sect.) 143(4):372-4 (Russian pages 965-967)

Beverton, R.J.H., Catch/effort assessment in some ICNAF fisheries. Res.Bull.int.Commn NW. 1965 Atlant.Fish., 2:59-72

Beverton, R.J.H. and V.M. Hodder (Ecis.), Report of working group of scientists on fishery 1962 assessment in relation to regulation problems. A.Proc.int,Commn NW.Atlant.Fish., 11 Suppl. :1-81

Beverton, R.J.H. and S.J. Holt, On the dynamics of exploited fish populations. Fishery Invest,, 1957 Lond.(2,19:1-533

Bigelow, H.B. and W.C. Schroeder, Fishes of the Gulf of Maine. Fishery Bull.Fish.Wildl.Serv. 1953 U.S., 53(74):1577

Blacker, R.W. Rare fishes from the Atlantic Slope fishing grounds, Ann.Mag,nat.Hist.,(13),5: 1962 261-71

Haddock: Rockall stock. Annls biol., Copenh., 18(1961):134-6 1963

Tests of a large-meshed topside chafer. Coop.Res.Rep.int.Coun.Explor.Sea (B), 1967 1967:17-22

Blaxter, J.H.S. and B.B. Parrish, The reaction of marine fish to moving netting and other 1966 devices in tanks, Mar.Res., 1966(1):1-15

Bohi, H., The effect of the approved ICNAF topside chafer on codend selectivity, Redbook mt, 1967 Coman NW.Atlant.Fish., 1966(3):122-31

Bowman, A., Spawny haddocks. Scient.Invest,Fishery Bd Scotl., 1922(4):1-15 1923 8:2 FIRM/S84 Haddock

Brown, W.W. and C. Cheng, Investigations into the food of the cod (Gadus callarlas L.) off Bear 1946 Island, and of the cod and haddock (G. aeglefinus L.) off Iceland and the Hurman coast. Hull Bull.mar.Ecol., 3(18):35-71

Carruthers, J.N., AL.. Lawford and V,F.C. Veley, Fishery hydrography: brood strength fluctua- 1951 tions in various North Sea fish, with suggested methods of prediction.Kieler Meeresforsch., 8:5-15

Carruthers, J.N. et al., Variations in brood-strength in the North Sea haddock, in the light of 1951 relevant wind conditions, Nature, Loud., 168:317-9

Chase, J., Winds and temperature in relation to brood-strength of Georges Bank haddock.J.Cons. 1955 perm.int.Explor.Mer, 21:17-24

Clark, J.R., Underwater television observations on the effect of chafing gear on escapement of 1958 haddock. A,Proc.,int.Commn NW.Atlant.Fish., 8:101-2

Sexual maturity of haddock. Trans.,Am.Fish.Soc., 88:212-3 1959

Size selection of fish by otter trawls. Results of recent experiments in the 1963 Northwest Atlantic. Spec.Publs int.Cominn NW.Atlant.Fish., 5:24-96

Clark, J.R. and E.L. Dietsch, Length-weight tables for Northwest Atlantic haddock. Sampl.Yb.int. 1959 Com,nn NW,,Atlant.Fish., 2(1957):25-37

Colton, J.B., Spring and summer distribution of haddock on Georges Bank. Spec.scient.Rep.U.S. 1955 Fish Wildl.Serv.(Fish.), 156:1-65

The distribution and behaviour of pelagic and early demersal stages of haddock in 1965 relation to sampling techniques, Spec.Publs int.Commn NW.Atlant.Fish,, 6:317-33

A comparison of current and long-term temperatures of Continental Shelf waters, 1968 Nova Scotia to Long Island, Res.Bull.int.Commn NW.Atlant.Fish,., 5:110-29

Colton, J.B. and R,F. Temple, The enigma of Georges Bank spawning. Limnol.Oceanogr., 6(3):280-91 1961

Dannevig, H,, The influence of temperature on the development of the eggs of fishes.Rep.Fishery 1895 Bd Scotl,, 13(3):147-52

Dogiel, V.A., G,K. Petrusevski and lu. I. Polyanski, Parasitology of fishes. Leningrad 1958 University Press English translation by Z. Kabata, Edimburgh, Oliver and Boyd, 384 p. 1961

Dragesund, O. (Ed.), International O-group fish surveys in the Barents Sea 1965-1968, Coop.Res. 1970 Rep.int.Coun.Explor.Sea (A), 18:1-81

Earli, R.E., A report on the history and present condition of the shore cod-fisheries of Cape A' 1880 Ain, Mass., together with notes on the natural history and artificial propagation of the species. Rep.U.S.Commnr Fish., 1878:685-740

FAO, Yearbook of fishery statistics, 1966. Yb.Fish.Statist., 22(1966):pag.var. 1967

Yearbook of fishery statistics, 1967. Yb.Fish,,Statist., 24(1967):pag.var. 1968

Fleming, J., A history of British animals. Edinburgh and London. 1828

Gill, T., Notice of a collection of the fishes of California, presented to the Smithsonian 1863 Institution by Mr. Samuel Hubbard. Proc.Acad.Sci.Philad., 1862(5):274-82

Goode, G,B. and T,H. Bean, Oceanic ichthyology. Spec.Bull.U.S.natn.Mus., 553 p. 1895 FIRM/S84 Haddock 8:3

Graham, H.W., Mesh regulation to increase the yield of the GeorgesBankhaddock fishery, A,Pz'oc, 1952 int,Commn NW,Atlant,Fish., 2:23-33

United States research report, 1966. Redbook int.Commn NW,Atiarit.Fish., 1967(2): 1967 136-51

United States research report, 1968. Redbook int.Commn NW.Atlant,Fish.,, 1969(2): 1969 121-36

Grosslein, M.D., Groundfish survey program of BCF Woods Hole. Comml Fish,Rev,., 31(8-9):22-.30 1969

Haddock recruitment predictions from bottom-.trawl catches of O-group fish in 1969a Subarea 5 and Division 4X. Res.Doc,Int,Comrnn NW.Atlant,Fish,, (69/89):1-14

Gulland, JA., Fishing and the stocks of fish at Iceland, Fishery Invest.Lond,(2), 23(4):152 1961

Survival of the youngest stages of fish, and its relation to year.class strength, 1965 Spec.Publs int,Commn NW,Atlant.Fish., 6:363-71

Hawkins, A.D. and C.J. Chapman, Underwater sounds of the haddock, Melanogrammus aeglefinus, 1966 J.mar,biol,Ass.U.K,, 46(2):241-7

Hawkins, A.D., C,J. Chapman and D,J. Symonds, Spawning of haddock in captivity. Nature, Lond,, 1967 215:923-5 (Issued also as Scott,Fish,Bull., (28):7-1O)

Hennemuth, R.C., Status of the Georges Bank haddock fishery, Res,Doc,Int,Coinmn NW.Atlant.Fish., 1969 (69/90):1-21

Hennemuth, R.C., M,D. Grosslein and F.D, McCracken, Abundance age composition of landings, and 1964 total mortality of haddock caught off southern Nova Scotia, 1956-1961, Res.Bull.jnt, Commn NW.Atlant,Fish., 1:43-73

Herrington, W,C., Factors controlling population size. Trans.N,Am,Wildl,Conf,, 9:250-263 1944

Hertling, H.0 Untersuchungen über die Ernährung von Meeresfischen, 2,Quantitative Nahrungsunter- 1940 suchungen an Schellfischen (Gadus aeglefinus), Wittlingen (G. merlengus) und Dorschen (c. morrhua) aus der Nordsee unter besonderer Berücksichtigung des Wärmewertes der Nahrung. Ber.dt.Wiss.Kominn Meeresforsch,, 9(2):274-317

Hodder, V.M., The possible effects of temperature on the fecundity of Grand Bank haddock, 1965 Spec.Publs int.Commn NW.Atlant.Fish., 6:515-22

Trends in the haddock fishery of ICNAF Subarea 3, Res.Bull.int.Commn NW,Atlant, 1966 Fish., 3:55-63

Hylen, A., Selectivity experiments with a large-meshed topside chafer.Coop.Res.Rep,int.Coun. 1967 Explor.Sea (B), 1967:23-5

ICES, Estimation of mortality rates, Annex to Arctic fisheries working group report of meeting 1965 in Hamburg, 18-23 Januâry 1965. ICES, CM 1965/3:1-9

ICES list of names of fish and shellfish. Bull.statist.Pch,marit,, 49(1964):71-80 1966

North-east Arctic fisheries working group. Report of the meeting at Copenhagen, 1968 4-14 December 1967. ICES CM 1968/F:19:1-1O

Report of the North-western working group, 1968, Coop,Res,Rep.int,Coun.Explor, 1969 Sea (A), 1968(10):1-32 8:4 FIRM S84 Haddock

ICES, North-east Arctic fisheries working group. Report of the meeting at Copenhagen, 13-17 1969a January 1969. ICES CM1969/F:2:1-21

Preliminary report of the O-group fish survey in the Barents Sea and adjacent 1969b waters in August-September 1969. ICES CM 1969/F:34:1_14

Nominal catch and fishing effort. Stock record data 1968. Demex'sal species. 1970 Statist.Newsl,, (46):154 p.

ICNAF, Report of the working group on joint biological and. economic assessment of conservation 1968 actions. A.Proc,int.Commn NW,Atlant.Fish,, 17(4):48-84

Report of 19th Annual Meeting of the International Commission for the Northwest 1969 Atlantic Fisheries, Warsaw, Poland, 2-7 June 1969, A.Proc.int.Commn NW,Atlant.Fish., 19(2): 14-28

Sampling yearbook: groundfish and flounders - 1968. Sampl.Yb.int,Commn NW.Atlant. 1970 Fish,, 13(1):1-269

Jensen, A,C, and J.R. Clark, Time of formation of scale annuli. Spec.Publs int.Commn NW.Atlarit. 1958 Fish., 1:193-7

Jensen, A.C. and J.P. Wise, Determining age of young haddock from their scales. Fishery Bull. 1962 Fish Wildl.Serv.U.S,, 61(195):439-50

Jensen, J.S., Gadoid Fish: young stages: Faroes. Annls biol., Copenh., 11(1954):90 1956

Faroes: pelagicO-group cod, haddock, whiting and saithe. Annls biol,,. Copenh., 1957 12(1955) :129

Faroes: pelagicO-group cod, haddock, whiting and saithe. Annls biol,., Copenh., 1959 14(1957):106

Jones, R., Haddock bionomics. 2. The growth of haddock in the North Sea and at Faroe. Mar.Res., 1962 1962(2):1-19

Some investigations of haddock movements. Spec.Publs int.Commn NW.Atlant.Fish., 1963 4:84-8

Post-war changes in the North Sea stock of haddock. ICES CM 1966/G:20:1-.2 1966

Jones, R. and A.S. Jermyn, Scottish investigations of the North Sea stock of haddock in 1967. 1968 Annis biol., Copenh., 24(1967):107-8

Scottish investigations on the Faroe stock of haddock, 1968. Annls biol., Copenh., 1969 251968):128-129 rabata,Z., Lernaeocera obtusa n.sp,, its biology and its effects on the haddock,Mar.Res., 1958 1958(3):1-26

Clavella (Copepoda) parasitic on British Gadidae:one species or several? 193 Crustaceana, 5(1):64-74

Kohier, A.C, and J.R. Clark, Haddock scale-otolith comparisons. J.Fish.Res,Bd Can., 15:1239-46 1958

Kohier, A.C. and D.,N. Fitzgerald, Cómparisons of food of cod and haddock in the Gulf of St, 1969 Lawrence and on the Nova Scotia Banks. J.Fish.Res.Bd Can., 26(5):1273-87

Konstantinov, K.G., Some information on adult fishes taken during NORWESTLANT 1-3, 1963. 1968 Spec,Publs int.Coinmn NW.Atlant.Fish., 7(1):219-24 8:6 FIRM/S84 Haddock

Olsen, S., Norwegian mesh selection experiments in 1963 and 1964. Coop.Res.Rep.int.Coun.Explor. 1966 Sea(B), 1965:164-8

Experiments with a topside chafer of double mesh-size. Coop,Res.Rep.int.Coun. 1967 Explor.Sea(fl), 1967:26-8

Parrish, B.B., Brood fluctuations in the North Sea haddock population.Challenger Soc.Abstr., 1950 3(2):18

Parrish, B.B. and R. Jores, Haddock bionomics. 1. The state of the haddock stocks in the North 1953 Sea 1946-50 and at Faroe 1914-55. Mar.Res., 1952(4):1-27

Pawsey, E.L., et al., Report on exploratory voyages to Lousy Bank and adjacent areas. Fishery 1924 Invest.Lond., (2), 7(2):1-.22

Petrova-Grinkevich, N.S., The feeding competition between haddock and cod in the Barents Sea. 1944 Trudy poliar.nauchno-issled.Inst.morsk.ryb.Khoz.Okeanogr., 8:416-28 (in Russian, English summary)

Pope, J.A. and W.B. Hall, Selectivity of polypropylene cod-ends. Coop.Res.Rep.int,Coun.Explor. 1966 Sea(B), 1965:173-87

Rae, B.B., The food of seals in Scottish waters, Mar.Res. 1968(2):1-23 1968

Rae, K.M., A relationship between wind, plankton distribution and haddock brood strength. 1957 Bull.mar.Ecol., 4:247-69

Raitt, D.S., The fecundity of the haddock. Scient.Invest,Fisher'y Bd Scotl., 1932(1):1-42 1933

The rate of mortality of the haddock of the North Sea stock, 1919-1938. 1939 Rapp.P.-v.Rêun.Cons.perm.int.Explor.Mer, 110(6):65-79

Reichenbach-Klinke, 11.-H., Krankheiten und Schadigungen der Fische. Stuttgart, Gustav Fischer 1966 Verlag, 389 p.

Richter, V.A. and V.1. Vinogradov, Results of a trawl survey carried out on the Scotian Shelf 1969 and Georges Bank in July-August 1968. Redbook int.Coznmn NW.Atlant.Fish., 1969(3): 129-43

Ritchie, A., The food and feeding habits of the haddock (Gadus aeglefinus) in Scottish waters. 1937 Scient.Invest.Fishery Bd Scotl., 1937(2):94 p.

Rollefsen, G., Observations on cod eggs. Rapp.P.-v.Héun.Cons.perm.int.Explor.Mer, 65:31-4 1930

Russell, E.S, Report on market measurements in relation to the English haddock fishery during 1914 tT years 1909 to 1911. Fishery Invest.Lond.,(2), 1(1):135 p.

Saemundsson, B., Marine pisces. Zoology Iceland, 4(72):1-150 1949

Saetersdal, G.S., The haddock in Norwegian waters. 2. Methods in age and growth investigations. 1953 FiskDir.Skr., (Ser.Havunders.), 10(9):46 p.

Gadoid fish: haddock. Age distribution and migrations. Annls biol., Copenh., 1956 11(1954):97-100

Use of otoliths and scales of the Arctic haddock, Spec.Publs int,Cominn NW.Atlant, 1958 Fish., 1:201-6

Saville, A., Eggs and larvae of haddock (Gadus aeglefinus L.) at Faroe. Mar.Res., 1956(4):1-27 1956 FIRM S84 Haddock 8:5

Leim, A,H, and W,B. Scott, Fishes of the Atlantic coast of Canada, Bull,Fish.Res,Bd Can,, 1966 (155):1-485

Linnaeus, C,, Systema Naturae. Tom, 1. Regnum animale Ed. X Holmiae, 824 p. 1758

Livingstone, R., Underwater television observations of haddock (Melanogrammus aeglefinus 1962 Linnaeus) in the cod-end, J.Cons.perm.int.Explor.Mer, 27:43-8

Lux, F.E., V.A. Richter and M.D. Grosslein, U.S..A.-U.S.S.R. otter trawl survey, Fall 1968. 1969 Redbook int.Commn NW.Atlant.Fish., 1969(3):144-56

McCracken, F.D., Selection by codend meshes and hooks on cod, haddock, flatfish, and redfish, 1963 Spec.Publs int.Conunn NW,Atlant.Fish,, 5(12):131-55

Distribution of haddock off the eastern Canadian mainland in relation to season, 1965 depth and bottom temperature. Spec.Publs int.Commn NW.Atlant,Fish,, 6(A3):113-29

Size, age and recruitment comparisons for haddock of the central Scotian shelf. 1968 Res.Bull.int.Comjnri NW.Atlant.Fish., 5:99-109

McIntosh, W.C. and A.T. Nasterman, The life-histories of the British marine food-fishes. 1897 Cambridge University Press, 516 p.

McRae, E.D. and L.E. French, An experiment in electrical fishing with an electric field used as 1965 an adjunct to an otter-trawl net. Commi Fish.Rev., 27(6):1-11

MAFF and DAFS, Fishing prospects 1969-1970: 1-38 1969

Malmgren, A.W., G3teborgs och Bohuslns fauna: Ryggradsdjuren. G5teborg, 674 p, 1877

Marak, R,R., Food habits of larval cod, haddock, and coalfish in the Gulf of Maine and Georges 1960 Bank area, J.Cons.perm.int.Explor,Mer, 25(2):147-57

Marak, R.R. and J.B. Colton, Distribution of fish eggs and larvae, temperature, and salinity in 1962 the Georges Bank-Gulf of Maine area, 1953, Spec.scient.Rep.U.S.Fish Wildl,Serv., (398) :1-61

Margetts, A.R., North Sea experiments with pelagic trawl. Fishing News, Aberd,, 2786:7 1966

Maslov, N.A., The trawl-fishery in the Barents Sea (1936-1938). Trud.poliar,nauchno-issled.Inst, 1939 morsk.ryb.Khoz.Okeanogr,, 6(1939):3-92

Mpíller, D. and G. Naevdal, Transferring polymorphism in fishes. In Polymorphismes biochimiques 1967 dea animaux. 10th European conference on animal blood groups and biochemical polymorphisms, Paris, 5-8 July 1966. Paris, Institut National de la Recherche Agronomique, pp.367-72

Studies on hemoglobins of some gadoid fishes. FiskDir,Skr,, (Ser,Havunders.) 1969 15(2);91_8

Needler, A.W.11., The migrations of haddock and the interrelationships of haddock populations in 1930 North American waters. Contr.Can.Biol.Fish,, 6(10):241-313

Nizovtzev, G.P. arid Z.P. Baranova, Gadoid Fish: young stages. Soviet investigations on young 1965 haddock of age-groups 0,I, II and III in the Barents Sea. Annls biol., Copenh., 20(1963):118-9

Ogilvie, H.S., The food of post-larval haddock with reference to the annual fluctuations in th 1938 haddock broods. Rapp.P.-v.Réun.Cons.perm,int.Explor.Mer, 107(3):57-66 FIRM 584 Haadock 8:7

Saville, A., The plariktonic stages of the haddock in Scottish waters. Mar.Res.., 1959(3):1-23 1959

Factors controlling dispersal of the pelagic stages of fish and their influence 1965 on survival, Spec.Publs int.Cominn NW.Atlant.Fish,, 6:335-48

Schmidt, J., The pelagic post-larval stages of the Atlantic species of Gadus. Part 1. Meddr, 1905 Kommn Danìn.Fisk.-og Havunders.(Fiskeri), 1(4):1-77

The distribution of the pelagic fry and the spawning regions of the gadoids in 1909 the North Atlantic from Iceland to Spain. Rapp,P.-v.Réun.Cons,perm.int.Exlor.Mer 1O(B4): 1-229

Schroeder, W.C., Results of haddock tagging in the Gulf of Maine from 1923 to 1943, J,mar.Res, 1942 5(1):1_19

Schuck, H.W., Haddock (Melanogranimus aeglefinus). Fish,Leafl.Fish Wildl.Serv.U,S., (198):1-lo 1947

Current haddock situation on Georges Bank, Commi Fish.Rev,, 10(10):1-6 1948

Relationship of catch to changes in population size of New England haddock, 1949 Biometrics, 5(3):213-31

Offshore grounds important to the United States haddock fishery. Res,Rep.U.S,Fish 1952 Wildl.Serv,, 32:1-20

Sonina, M.A., Relationship between the growth rate and population density of haddock in the 1965 Barents Sea. Spec.Publs int.Commn NW.Atlant,Fish., 6:565-70

Some reasons for a change in the age-length composition of the haddock population 1967 in the southern Barents Sea in 1950-1965, Mater rybokhoz,Issled.sev,Bass., 8:28-44

Migrations of haddock of the Barents Sea and factors determining them, Trudy 1969 poliar,nauchno-issled.Inst.morsk.ryb.Khoz.Okeanogr, 26:1-126

Stein, G.A., On the systematics of Urceolariidae (Infusoria peritricha), Zool.Zh,, 40(8):1137-42 1961

St. Inst., Sea fisheries. Boats and methods of fishing.The fishing nets (North-east Atlantic) 1968 order 1968, Statut.Instruìn,U.K., 1968(2075):1-8

St. Inst., Sea fisheries. Boats and methods of fishing, The fishing nets (North-west Atlantic) 1969 order 1969. Statut,Instrum.U.K., 1969(628):1-7

Svetovidov, A.N., Fishes Gadiformes. Fauna SSSR, 9(4):1-218. Issued also as Transi,CoTs63-. 1948 11071) Israel program for Scientific Translations

Templeman, W., Mass mortalities of marine fishes in the Newfoundland area presumably due to low 1965 temperature. Spec,Publs int,Commn NW.Atlant.Fish., 6:137-47

Some instances of cod and haddock behaviour and concentrations in the Newfoundland 1965a and Labrador areas in relation to food, Spec.Publs int,Commn NW,Atlant,Fish., 6:449-61

Templeman, W. and V.M. Hodder, Distribution of haddock on the Grand Bank in relation to season, 1965 depth and temperature. Spec.Publs int.Cormnn NW.Atlant.Fish., 6:171-87

Distribution of haddock on St. Pierre Bank (ICNAF division 3Ps) by season, depth 1965a and temperature. Spec.Publs int,Commri NW.Atlant.Fish., 6:189-97

Templeman, W. and H.J. Squires, Relationship of otolith lengths and weights in the haddock 1956 Melanogrammus aeglefinus (L.) to the rate of growth of the fish, J.Fish,Res.Bd Can, 13(4) :467-87 8:8 FIRM/S84 Haddock

Thompson, H., Problems in haddock biology with special reference to the validity and utilization 1923 of the scale theory. 1. Preliminary report, Scient.Invest.Fishery Bd Scotl., 1922(5) :1-78

Haddock biology. 2. Frequency and distribution of the age classes in 1923. Scient. 1924 Invest.Fishery Bd Scotl,, 1924(1):1-20

Haddock biology (North Sea). A brief survey of recent data, methods and results. 1929 Rapp.P.-v.Réun.Consperm.int.Explor.Mer, 54:135-163

Haddock biology. 5. General features in the biology of the haddock (Gadus 1929a aeglefinus L.) in Icelandic waters in the period 1903.-1926. Scient.Invest..Fishery Bd Scotl., 1928(5):1-63

The occurrence and biological features of haddock in the Newfoundland area. 1939 Res.Bull.Div.Fish.Resour.Newfoundld, (6):1-31

Treschev, A. I. and G.N. Stepanov, Comparative selectivity of trawl nets made of kapron and 1968 manila. Redbook int.Commn NW.Atlant.Fish,, 1968(3):95-100

Trout, G.C., Specific seasonal density changes in cod and haddock populations. ICES CM 1962 1962 Comp,Fish Comm. 24:1-6

Tseeb, R,Ia. Observations on the feeding of haddock in an aquarium. Trudy murmansk.biol.Inst., 1962 4(8): 153-61

On the mode of haddock feeding on benthos. Zool.Zh., 41(1):11O-5 1962a

Tytler, P,, Relationship between oxygen consumption and swimming speed in the haddock, 1969 Melano9rammus aeglefinus. Nature' Lond,, 221:274-5

Viadykov, V.D., Haddock races along the North American coast. Prog.Rep.Atlant.biol,Stn, 14:3-7 1935

Walford, L.A., Effect of currents on distribution and survival of the eggs and larvae of the 1938 haddock (Melanogrammus aeglefinus) on Georges Bank, Bull.Bur,Fish., Wash., 49(29): 1-73

Wells, R., Haddock length conversion factors for St. Pierre Bank. Res.J3ull.int.Commn NW.Atlant. 1968 Fish,, 5:137-40

Wigley, R.L., Food habits of Georges Bank haddock. Spec.scient.Rep.U.S.Fish.Wildl.Serv., 1956 165:1-26

Woodhead, P.M.J., Diurnal changes in trawl catches of fishes, Rapp.P.-v.Réun.Cons.perm.int. 1964 Ex.lor.Mer, 155:35-44

Effects of light upon behaviour and distribution of demersal fishes off the North 1965 Atlantic. Spec.Publs int.Commn NW.Atlant.Fish., 6:27-87

Zukowski, C., Some data on catches and biology of haddock (Melanogramrnus aeglefinus L.) from 1968 Georges Bank and Sable Island Bank, Res.Doc.Int.Cominn NW.Atlant.Fish., (68/54):1_4

SYNOPSIS OF FISHERIES BIOLOGICAL DATA

This is one of a series of documents issued by FAO, CSIRO and USFWS concerning species and stocks of aquatic organisms of present or potential economic interest. The primary purpose of this series is to make existing infor- mation readily available to fishery scientists according to a standard pattern, and by so doing also to draw attention to gaps n knowledge. lt is hoped that synopses in the series will be useful to other scientistsinitiatinginvesti- gations of the species concerned or of related ones, as a means of exchange of knowledge among those already working on the species, and as the basis for comparative study of fisheries resources. They will be brought up to date from time to time as further information becomes available either as revisions of the entire document or their specific chapters.

The relevant series of documents are:

FAO Fisheries Synopsis No. FR/S replacing, as from 1.1.63 FAO Fisheries Biology Synopsis No. F B/S CSIRO Fisheries Synopsis No. D FO/S and USFWS FAO Fisheries Synopsis No. N M FS/S

Synopses in these series are compiled according to a standard outline described in FIb/Si Rev. 1(1965). FAO, CSIRO and USFWS are working to secure the cooperation of other organizations and of individual scientists in drafting synopses on species about which they have knowledge, and welcome offers of help in this task. Ad- ditions and corrections to synopses already issued will also be most welcome. Comments including suggestions for the expansion of the outline and requests for information should be addressed to the coordinators and editors of the issuing organizations. FAO: Fishery Resources Division Marine Biology and Environment Branch Food and Agriculture Organization of the United Nations Via delle Terme di Caracalla 00100 Rome, Italy USFWS: CSIRO: Chief, Scientific Publications Unit Scientific Editor National Marine Fisheries Service CSIRO Division of Fisheries and Oceanography Bldg. 67, Naval Support Activity Box 21 Seattle, Washington 98115, Cronulla, N.S.W. U.S.A. 2230 Australia

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

The followingsynopses in this series have been issued since January 1970: DFO/S4 Synopsis of biological data on the rainbow prawn, Parapenaeopsis scuiptiis (Heller, 1862) 1970 D FO/SS Synopsis of biological data on the school prawn, Metapenaeus macleayi (Haswell, 1879) 1970 BCF/541 Synopsis of biological data on chum salmon Oncorhynchus keta July 1970 FIRI/580 Synopsis of biological data on the eel Anguilla anguilla (Linnaeus) 1758 October 1970 Rev. 1 * FIRM/591 Synopsis of biological data on Crangon cran gon October 1970 * FIRM/S92 Synopsis of biological data on Pandalus montagui October 1970 * FIRM/593 Synopsis of biological data on the Jumbo tiger prawn Penacus monodon Fa- bricius 1798 October 1970 * FIRM/S94 Synopsis ofbiological data on the Indian prawn Ponaeus indicus H. Milne Edwards, 1 837 October 1970 * FIRM/S95 Synopsis of biological data on the prawn Pandalus platyceros Brandt, 1851 October 1970 * FIRM/596 Synopsis of biological data on the penaeid prawn Solenocera indica Nataraj, 1945 October 1970 * FIRM/S97 Synopsis of biological data on the penaeid prawn Metapenaeus dobsoni (Miers, 1878) October 1970 * FIRM/598 Synopsis of biological data on the penaeid prawn Metapenaeus affinis (H. Milne Edwards, 1837) October 1970 * FIRM/S99 Synopsis ofbiological data on the ocean shrimp Panda/us jordani Rathbun, 1902 October 1970 * FIRM/S100Sinopsis sobrelabiología del camarón blanco Penaous schmitti Burkenroad, 1936 October 1970 * FIRM/S101 Synopsis of biological data on the white shrimp Penaeus setiferus (Linnaeus) 1767 October 1 970 * FIRM/5102Synopsis of biological data on the brown shrimp Penaeus aztecus aztecus Ives, 1891 October 1970 * FIRM/S103Synopsis of biological data on the pink shrimp Penaeus duorarum duorarum Burkenroad, 1939 October 1970 * FIRM/S104Synopsis of biological data on the penaeid prawn Metapenaeus monoceros (Fabricius, 1798) October 1970 'FIRM/S105Synopsis of biological data on the penaeid prawn Metapenaeus brevicornis (H. Milne Edwards, 1837) October 1970 * FIRM/S106Synopsis of biological data on the penaeid prawn Parapenaeopsis stylifera (H. Milne Edwards, 1837) October 1970 * FIRM/Si 07Sinopsis sobre la biología del camarón nailon Heterocarpus reedi October 1970

DFO/S6 Synopsis of biological data on the greentail prawn Metapenaeus bennettae Racek and Dall, 1965 1970 DFO/57 Synopsis of biological data on the eastern king prawn Penaeus p/ebejus Hess, 1865 1970 DFO/S8 Synopsis of biological data on the banana prawn Penaeus merguiensis de Man, 1888 1970 FIR M/S82 Synopsis of biological data on North Atlantic sandeels of the genus Ammodytes (A. tobianus, A. dub/us, A. americanus and A. marinus) November 1970 FIR M/S83 Synopsis of biological data on Saccorhiza po/yschides November 1970 NMFS/S79 Synopsis of biological data on Pacific Ocean Perch, Sebastodes alutus December 1970 FIRM f S38 Synopsis of biological data on knobbed wrack Ascophyllum nodusum (Linnaeus) Rev. 1 Le Jolis December 1970 FIR M/S84 Synopsis of biological data on haddock Melanogrammus aeg/efinus (Linnaeus 1758) December 1971

* Published in the Proceedings of the World Scientific Conference on the Biology and Culture of Shrimps and Prawns, Mexico, 12 to 21 June 1967.

M /C3584/1 .72/E/2/750