FAO Fisheries Synopsis No. 82 FIRM/Sri (Distribution restricted) SAST - toblanus - 1.72(04),002,02 Ammodytos dublus - 1.72(04),002,05 Ammodyte. amepleaT178 - 1.72(04),002,06 Ammodytes ma,'Inus - 1.72(04),002,07

SYNOPSIS OF BIOLOGICAL DATA ON NORTH ATLANTIC SAND OF THE AMMODYTES A. tobianus, A. dubius, A. americanus and A. marinus

Prepared by P. J. Reay

FOOD AND AGRICULTURE ORGANIZATION OF THE UNITED NATIONS ROME, 1970 DOCUMENTS OF THE FISHERY DOCUMENTS DE LA DIVISION DES DOCUMENTOS DE LA DIRECCION RESOURCES DIVISION OF FAO DE- RESSOURCES HALIEUTIQUES DU DE- DE RECURSOS PESQUEROS DEL DE- PARTMENT OF FISHERIES PARTEMENT DES PÊCHES DE LA FAO PARTAMENTO DE PESCA DE LA FAO

Documents whicharenotofficial Des documents qui ne figurent pas Esta Subdirección publica varias se- FAO publications are issued in several parmi les publications officielles de la ries de documentos que no pueden series. They aregivenarestricted FAO sont publiés dans diverses séries. considerarse como publicacionesofi- distribution and thisfact should be Is font seulement l'objet d'une distri- ciales de la FAO. Todos ellos tienen indicatedifthey arecited. Most of bution restreinte, aussi convient-il de distribución limitada, circunstancia que them are prepared as working papers le préciser lorsque ces documents sont debe indicarse en el caso de ser ci- for meetings, or are summaries of infor- cités.lis'agit le plus souvent de do- tados. La mayoría de los títulos que mationforuseof member govern- cuments de travail préparés pour des figuranen dichas series son docu- ments,organizations,and specialists réunions, ou de résumés d'information mentos de trabajopreparadospara concerned. àl'intentiondes gouvernements des reuniones o resúmenes de información pays membres, ainsi que des organi- destinados a los estados miembros, sations et spécialistes intéressés. Ces organizaciones y especiálistasintere- séries sont les suivantes: sados.

FAO Fisheries Report FRJR (No.) FAO Fisheries Circular FRIC (No.) FAO Fisheries Synopsis FR/S (No.)

Special groups of synopses are iden- Des catégories spéciales de synopses Grupos especiales de sinopsisse tified by symbols followed by classifi- sont identifiées à l'aide de symboles distinguen conlassiglassiguientes, cation numbers based on indexed code suivis des chiffres de classification ba- seguidas por números de clasificación of "Current Bibliography": sés surlecode d'indexation de la que se basan en las claves de los ín- <'Current Bibliography »: dices dela<'Current Bibliography. SAST Dataconcerningcertainspe- SAST Données sur certaines espèces SAST Datos relativos a ciertas espe- cies andfishstocks. et populations de poissons. cies y poblaciones. MAST Informationonmethodsand MAST Renseignements surdes mé- MAST Sinopsis sobre métodos y ma- subjects. thodes et des sujeth. terias. OT Oceanographic data. 01 Données océanopraphiques. 01 Sinopsis sobre oceanografía. IT Limnological data. IT Données limnologiques. IT Sinopsis sobre limnologia. and et y CART Information concerning - CART Renseignements sur les pêche- CART Información sobre los recursos eries and resources of certain ries etles ressources de cer- acuáticosvivos de algunos countries and regions (FID/S). tains pays et régions (FD/S). países y regiones (FID/S).

FAO Fisheries Technical Paper FRIT (No.)

Special groups of Technical Papers Des catégories spéciales de docu- Grupos especiales de documentos are identified by: ments techniquessontidentifiéesà técnicos se identifican por las siglas l'aide des symboles suivants: siguientes: RE Indexedlistsofexpertsand RE Listesindexéesd'expertset RE Listas índices de expertos y de institutions drawn fromRegis- institutions tirées des registres institucionestomadasdelos ters maintained by the Fishery tenus à jour parlaDivision registros que se llevan enla Resources Division. des ressources halieutiques. Dirección de Recursos Pesque- ros.

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MFS Provisionaleditionsof"FAO MFS Editionsprovisoires des «Ma- MFS Ediciones provisionales de los Manuals inFisheries Science." nuels FAO de science halieu- «Manualesde la FAO de tique «. Ciencias Pesqueras '>.

Some documents also have another Certains documents portent parfois Algunos documentos tienen también identification, if, for example, they have d'autres numéros d'identification,par otra identificación si, por ejemplo, son beencontributedto ameetingfor exemple, s'ilsont été préparés pour contribuciones a una reunión cuyos whichpapers have beennumbered une réunion dont les documents ont été documentos han sido marcados con according to another system. marqués à l'aide d'un autre système. arreglo a otros sistemas. FAO Fisheries Synopsis No. 82 FIRM/S82 (Distribution restricted) SÄST - Arninodytes tobianus 1.72 04) 002,02 Ammod.y-tes d.ubius 1,72 04 002,05 Ainmodytes aznericanus 1.72 04 002,06 Amrnod,y-tes marinus 1.72 04 002,07

SYNOPSIS OF BIOLOGICAL DATA ON NORTH ATLA1TIC

SMmEELS OF TIDE HNUS A!OODYTES

(A.tobianus, A.dubius, A.aznericanus and. A,marinus)

Prepared by

P.J. REAY Marine Reso,.u'oes Research Unit Portsmouth Polytechnic Portsmouth, U.K.

BDOD AND AGRICULTURE ORGANIZATION OF TKD UNITED NATIONS Rome, October1970 PREPARATION OF THIS SThOPSIS

This Synopsis is concerned with the biolor and exploitation of Amrnodytes in the North Atlantic, The inclusion of four species within the sane Synopsis is justified by their close similarity, and by the relatively small amount of available information on them; some data from the related Pacific species have also been included? but only in sections where no information was available on the Atlantic species, i Sandeels are currently of considerable importance as a fishmeal resource in the , and such exploitation is likely to develop in other North Atlantic areas in the near future; inshore species are less important and their present use is mainly as bait.

The author wishes to acicwwledge the advice and information received from A, Wheeler (British Museum (Natural History)) and C.T, Macer (Fisheries Laboratory, Lowestoft) during the preparation of this Synopsis,

Distribution "Current Bibliography" entry

FAO Department of Fisheries Reay, P,J. (1970) 17.-6M32O FAO Regional Fisheries Officers FAO Fish.Synops., (82):pag,var, Regional Fisheries Councils and Synopsis of biological data on North Commissions Atlantic sandeels of the genus Ammodytes Selector SÌ' (A, tobianus, A. dubius, A, americenus Author and A. marinusT

JLNW, AilE, Ammodytidae, , nomenclature. Morphology, Di etribut ion, Bionomics and life history, Behaviour, Population, Exploitation. Protection, Management, Pond fish culture. Selected bibliography, FIR1T,/S82 North Atlantic Sarideels

CONTENTS Page

i IDE2TITY 1:1

1.1Nomenclature i

1.11 Valid names i 1.12Objective synon1my*

1.2Taxonomy 2

1,21Affinities 2 1.22Taxonomic status 3 1.23Subspecies* 1,24Standard common nan, vernacular name s 3 Morphology 1.3 5 1.31 External morphology 5 1.32Cytomorphology5f 1.33Protein specificity* 2 DISTRIBUTION 2:1

2.1 Total area 1

2,2Differential distribution 1

2.21Spawn, larvae and juveniles 1 2.22Adults 4

2,3Determinants of distribution 4

2.4 I{ybridization*

3 BIONOMICS AND LIFE HISTORY 3:1

3.1Reiiroduction 1

3.11Sexuality i 3,12Maturity i 3.13Mating* 3.14Fertiliza-tion* 3.15Gonads i 3.16Spawning 2 3.17Spawn 3

3.2Preadult phase 3

3,21Embryonic phase 3 3,22Larval phase 3 3.23Adolescent phase 5

3.3Adult phase 5

3.31Longevity 5 3.32Hardiness 5 3.33Competitors 5 3.34Predators 6 3,35Parasites, diseases, injuries and abnormalities 6 iv FIRM S82North Atlantic Sandeels

304Nutrition and growth 3:7

3.4].Feeding 7 3.42Food. 3,43Growth rate 8 3.44Metabolism*

3.5Behaviow 12

3.51Migration and, local movements 12 3.52Schooling 12 3.53Responses to stimuli 13 4 FOJLATION 4:1 4.1 Struc'ture i 4,11Sexratio i 4,12Age composition i 4.13Size composition i

4.2Abundance and, density 4

4.21Average abundance 4 4.22Changes in abundance 4 4.23Average density 4 4.24Changes in density 4

4,3Nata1itr and recruitment 5 4,31Reproduction rates* 4.32Factors affecting reprod.uction* 4,33Recruitment 5

4.4Mortality and morbidity 5

4.41Mortality rates 5 4.42Factors causing or affecting mortality 5 4.43Factors affecting morbiditylf 4.44Relation of morbidity to mortality rates* 4,5Dynanics ofpopulation (as a whole)*

4.6The population in the comnmnity and the ecosystem 5

5 CPLOITATION 5:1

5,1 Fishing Euiument 1

5.11Gears i 5.12Boats 1

5,2 Fishing Areas 1

5.21General geographic distribution 1 5.22Geographic ranges 2 5,23Depth ranges 2 5.24Conditions of the grounds 2 £IR/S82 North Atlantic Sand.eels y

Pa1e

5,3 Fishing Seasons 5:2

5.31 General pattern of seasons 2 5.32 Dates of beginning, peak and end of seasons 2 5,33 Variation in date or duration of season 3

5.4Fishing operations and results 3

5.41 Effort and intensity 3 5.42 Selectivity 3 5.43 Catches 3

6 PROTECTION MT]) MAiL& 'si 1

6.1 Regulatory (Legislative) measures 1

6,2 Control or alteration ofyeica]. features of the environinent*

6.3 Control or alteration of chemical features of the envirorixnent*

6.4 Control or alteration of the biological features of the environnient*

6.5 Artificial stocking'

7 POND FISH CULTURE*

8 NEFJRENCES 3:1

* As no information was available to the author, these itens have been omitted from the tezt, FIRM/S82 North Atlantic Sand.eels 1:1

1. IDENTITY

1.1 Nomenclature Amniodytes dubiusReinhardt, J,, (1838), K. danske Vidensk, Seisk. Skr.nat, 1,11 Valid flanes 0g math. 7 (54): 131.-2.

The following are the original Ammody-tes americanus DeKay, J.E., combinations under which the species were first (1842). Zoology of New York, or the New York described: fauna, Pt, IV. Genus Ammodytes, 317-8.

Ammod,ytes tobianus Linnaeus, C,, Axnmody-Les marinus Raitt, D.S., (1934). (1758). Systema naturae, I: 123. Ammodytes. J. Cons, perm. i. Explor.Mer., 9: 365-72. (Ed. Decima): 247-8,

TABLE I

Meristic and morphological features used to distinguish A. tobianus and A. Tnarinus

V ,tobianus A. marinus Author

1 Total vertebral number 60-68 65-75 ) rtaitt (1934) 2 Dorsal fin ray number 49-58 56-63 KMndler (1941)

3 Anal fin ray number 24-.32 29-33 ) Soleim (1945) 4Pectoral fin ray number 10-14 12-16

5 Plicae (skin fold) number120-138 140-150 Andriashev (1954) 6 Low ventral median dermal absent fold present DUncker and Mohr (1939) (not recogmised by later authors such as Braun (1941)

7 Radialia on lower well feint Andriashev (1954) part of suboperculum pronounced 8 Scales on dorsal extend absent from surface anterior to anterior and Andriashev (1954) and alongside alongside dorsal dorsal fin fin

9 Scales on ventral in tight loosely Andriashev (1954) surface chevrons arranged 10 Scales over musculature at base present absent Corbin (pers. comm,) of caudal fin

11 Pinentation on slight, conspicuous, &e Present author head usually in- sharply contrast.- conspicuous ing with unpig- ment ed areas 12 Lar,al pinen.- tation:-. none until appears before caudal piiien- ) 15mm. 15 mm. tation Eiriarsson (1955 pre-anal ventral rapidly remains conspic- Macer (1967 body pigment obscured by ous the body wall dorsal pigment commences commences at anterior to notochord tip notochord tip 1:2 FIRM/582 North Atlantic Sandeel

1.2Taxonomy species are clearly distinct on both meristic and morphological grounds. 1,21 Affinities A. americanus and A. dubius both appear Suprageneric to resemble A, maninus rather than A, tobianus on morphological grounds (eg. larval pigmentation Phylum-Vertebrata patterns; Richarde, 1965; Macer, 1967), and the Subphylum-Craniata characteristics currently used to distinguish Superclas s-Gnatho st ornat a them from each other and from A. marinus are Series-Pisces somewhat tenuous (See Section 1.22). Richaada Class-Teleostomi et al. (1963) described A. aznenicanus (as Subclass-Act inoptcrygii A, hexapterus) as an inshore fish with body-depth Order-Clupeiformes 7.1 - 9.7% of standard length and with meristic Suborder-Ammodytoidei characteristics of V61 - 73, D = 51 - 62, Fami ly-Ammodyt idae A 23 - 33, and A. dubius as offshore, with a body-depth of 6.8 - 9.2% and V 6 - 75 (78), Generic D = 56 - 68, and A = 27 - 35.In view of the wide degree of overlap between these two Genus:Axnmodytes, Artedi P. (1738), in definitions, it is suggested that, until further Bibliotheca ichthyologioa seu historia information becomes available, the restricted luterana ichthyologiae.Genera Piscium, concept of A. dubius as used by Leim and Scott Ichthyologiae pars III.Nalacopterygii. (1966), be adopted, and that the remaining BW Ammodytes: 55-57. Atlantic Arnniodytes be included as a rather heterogenous assemblage, with polyrnodal meristic Genotype = A. tobianus Linn. counts, under the name of A, americanus; the menistic features of the two species would then The genus Ainmodytes includes the Pacific be as described in Table II. species A. hexapterus Pallas., and A, personatus Girard, in addition to the Atlantic species, Thus, in the North Atlantic, A, tobianus A. tobianus, A. marinus, A. dubius and is clearly distinguishable on both meristic and A, americanus as listed in Section 1.11, and morphological grounds from the other three has the following diaguostic characteristics:- species.Of these, A,, dubius is probably distinct 1.Premaxillae protrusible menistically, but there is a wide overlap and 2,Vomer without teeth close similarity between A. anericanus and Body covered in scales which on A, marinus, the dies lie in oblique skin-folds (plicae) Three other species of sandeel are Lateral-line pores linearly arranged present in the North Atlantic, 'jeeroslus Dorsal and anal fin margins straight, lanceolatus (LeSauv.), H. immaculatusCorbin), TABLI II Meristic Characteristics of A. ainericanus and A. dubius

V D A A, americanus 61-73 51-62 23-33 Richards et al, (1963) A, dubius 71-78 6169 31-36 Leim & Scott (1966); Scott, (1968)

Characteristics 1-2 distinguish and Gymnammod,ytes semisquamatus (Jourdain). Ammodytes from the genus Hyperoplus GUnther., and These are all restricted to European coasts characteristics 3-5 distinguish both from the and can be distinguished from the species under genus Gynmammodytes DUncker and Mohr,This genericconsideration by the generic characteristics concept is now adopted by several authors, includinggiven above, Wheeler (1969). Subjective Synonymy Specific 1.A, tobianus Diagnosis A, lanceaCurvier, G., (1829) Le The diagnostic features of A. tobianus Règne distribue d'apres son organisation, and A.marinus are listed, in Table I; the two FIR/S82 North Atlantic Sandeels 1:3

....,Les qui1les.Parìs: 360; and. Yari'ell D Low meristio counts with (1936). Vu61-73; up to 22 cm The naine A. tobianus was mistakenly used, by Cuvier long..,.., , ,,,, ,,.. ... , .A, alnericanus to describe the greater sandeel which LeSauvage DD High meristic counts (1824) had already nazned.A. (uIyperoplus) lan- with Vu71-78; up to ceolatus to distinguish it from the lesser sand. 38cm long. , , ...... s....,..A, dubius named. A, tobianus by Linnaeuz (1758).Curvier E Low meristic counts then gave the naine A, lancea to the latter species, with V= 56-67.,,,..,,,,,,.,A. personatus believing that Linnaeus had originally been in RE High meristic counts possession of greater sandeels.Jensen (1941) with V = 64-74,,.,,,..,,.,.A. hexapterus has however shown that Linnaeus based. his work on lesser sandeels, although his type specimens 1,22 Taxonomic status may have included. both A. tobianus and. A.arinus; on this basis in fact, Jensen advocated retention All the species under consideration are of the naine A, lancea in preference to A, tobianusmorphospecies and. none have been established as and. it has been used. objectively in this way by a result of either breeding experiments, many authors since.As And.riashev (1954) and. cytomorpholor or comparative serologr. Backus (1957) have however pointed out there can be no valid grounds for the uso of the naine A. tobianus is clearly a distinct A. lancea, and the situation should. be resolved species ozi morphological, meristic and, by establishing either a neotype or a leototype physiological (spawning season) grounds, bu-t the on the basis of material and. retaining remaining species in the genus appear to share the name A. -tobianus as used. initially by a common larval pi.íentation pattern (Macer 1967; LeSauvage, and, later, imPlicitly,by1aitt (1934) Kobayashi, 1962) and. spawning season, and. may, as Lindberg (1937) and others have suggested, be geographical morphological forms of a single A. inarinus poly-typic species; in support examination of a A, tobianus(ex parte; mixed with small number of preserved specimens of A. dubius, A, tobianus and. not distinguished. by any authors A,1aericanus and, It, Dersonatus by the present authorhas indicated, that these species are prior to Raitt (1934). similar to A. marinus in terms of scale-pattern uA, lancea marinus(non Cuvier). characteristics.Further work is obviously Jensen (1941), needed., howeverezd in the meantime it is =A. dnbius(non Reinhardt), Dflncker convenient to adopt the nomenclature and. species and Mohr (l939. definitions outlined, above, =A. hexapterus marinus (non Pallas), A. tobianus exists as two distinct but syinpatric forms autumn and. spring spawners Lindberg (19377. (Baiu, 1935),in addition to -the seasonal differences in gonad maturity arid. the occurrence A, aznericanus and growth of young stages induced. by 'the different spawning seasons, individuals can uA. terus(non Pallas). Richardsusually be distinguished by the structure of et al. (1963). their otolitha, (Knd,ler, 1941) and. populations in many instances on meris'bic grounds (Section A. dubius 1.31 and Table III).In the otoliths of spring- spawning fish, the first opaque ring is uA, lancea dubius(non Reithard.t), separated from the opaque centre by a broad Jensen (1941). hyaline zone; in those of autumn-spawners the centre and. the first ring are usually almost Artificial key for sand.eels of the genus confluent, Ammo dy't es 1,24 Standard common names, A Present in the North Atlantic...... vernacular names AA Present in the North Pacific..,,,.,,..,,E The follo list of common names is B Present in the North East Atlazitic..,,,.0 mainly based. on the ICES list of names of fish BB Present in the North West Atlantic....,.D (Bulletin Statistique Vol, 49). C Belly-scales in tight chevrons; scales present on the base of the caudal fin; low meristic Danish ToMs Dut ch/Flemi ah Zand,spiering, Smeelte counts with Vu60-68; inshore,.A, tobianus English CC Belly-scales loosely Sazideols, Lesser arrangedscales absent from Sande e la the base of the caudal fin; Faerose Nebbasild. Finnish Thulenkalat high meristic counts with P'rench Vu65_75; offshore,,,.,,.,,.,, A. marinus Equille j. - _ .-' .-.--..-- '.-.- . Ammodytes tobianus. 16 cm. Fig. 1. External morphology of A. tobianus. FIRN/582North Atlantic Sandeels 1:5

Germaxi Sandaal, Sandspierling, dorsal fin rays, but the number of anal fin rays Tobias Fisch is less than half the number of vertebrae; Greelandic Putorutotoq lCMndLer (1941)in fact found a strong positive Icelandic Tronusilli, Sancisili correlation between the number of vertebrae and Irish (Gaelic) Corr the number of fin rays, with, for example in Norwegian Sil, Tobis A. tobianus: r +0.69(dorsal rays) and Polish Tobiasz, Dobijak r + 0.42 (anal rays). Portuguese Fanchao, Agulhae Spanish Lanz6n Data on vertebral counts are Swedish Tobis summarised in Tables III and IV; where possible U.S.S.R. - Peschaxiiza and necessary, standard deviations and confidence U.S.A., Canadian - Launce, Sand Launce limits have been calculated from the original data. In A. tobianus counts are not available over a very wide geographical area, and, the most 1.3 Morphology conspicuous intraspecific variation is between the spring and autumn spawning groups. In general, 1.31External Morphology autumn spawners have more vertebrae and show a slightly greater variance but the differences The body is elongate, suboylindrical between the groups off the Isle of Maxi, and a-t and covered with small cycloid. scales which, on two of the Baltic stations givenbyKndler, the sides of the body, lie on the underside of are no-t si.ificant at the95%level. In oblique skin-folds (plicae). The head is pointed A. marinus, samples have been examined from and the lower jaw projects anteriorly; the mouth stations throughout most of its geographical is bordered by the premaxillae which are prot- range, and a clear latituiinal dine in rusible, arid teeth are absent. The lateral lines vertebral number is apparent; in addition, run dorso-laterally, and there is a pair of samples from the North Sea show a significantly ventro-lateral skin-folds, Pelvic fina are absent higher vertebral number than those from a the dorsal and axial fins are elongate and the similar latitude in the Baltic arid Irish Seas, forked caudal fin is preceded by a well-marked peduncle, There are no spines, but the fin-rays In the case ofBinAtlantic Aminody-tes of the dorsal and anal fins are wijointed and the position is confused by the uncertain uribranched. The gill openings are strongly relationship between A. americanus and A. dubius extended anteriorly, and there are four gill (see Section 1.21).However, it is clear from arches with a slit behind the last arch, (Based Table III that, as in A. marmnus,there are on Andriashev,1954). latitudinal clines in vertebral number, and in addition separate high- (offshore) and low- Ammody-tes species are white on the (inshore) meristic groups at the saine latitudes. ventral surface, silvery on the sides and light Several of the groups show conspicuously skewed brown to dark grey on the dorsal surface. distributions, and variance is typically greater Generalised external morphology is illustrated in than in A. tobianus said, A. marinus, Richards e-t Fig. 1. al,(1963)have pointed out, that A. axnericanus may be partially divided into a group of small Intraspecific morphometric variation size fish inhabiting semienclosed, bays and a has received little attention.Kirillov(1936) group of larger fish with a wide meristic range found some sigaificant differences in A. marinus which are caught along exposed coasts. These between a sample from Novaya Zemblya and one from authors initially recognised three main groups the Murman coast; in particular, the latter fish of Ammody-tes in theBWAtlantic with high, inter- had deeper bodies arid smaller heads. Body depth mediate and low meristic counts; mainly on the was also considered by Richards et aj. (1963) in basis of body-depth, they then divided the a detailed study of Ammod,y-tes from the east coast iiitermedia-te group so that high to intermedia-te of North America; in general, for fish under75mm(. dubius) said low to intermediate (A. hexapterus in length, the low meristic count, inshore = A, americanus) groups were formed, populations had significantly deeper bodies than the high meristic-count offshore ones. Regression coefficients of depth on length ranged from 0.036to0,162and suggested allometric growth.

Most attention has been paid to merietic characters, and in particular, the total number of vertebrae, 1x1 very general terms, there are about ten more vertebrae than there are 1:6 FIRM/582North AtlanticSand.eels

TABLE lU

ntraepeoiic ¡uoristic variation; counts of the total number of vortobae in A. tobianus, A. marinus and A. dubius

Number of vertebrae includes the urostyle

A. * tobianus GROUP N. Range Mean ,95%C.L. S.D.

Iceland(Bruun1941) 0 52 60-65 62.56j0.28 63 1.01 Faeroes(Bruun1941) 0 22 63-66 64.52 +0,46 65 1.02 Isle of Man(Cameron 1958) AS 367 62-66 63,91+0.10 64 0.90 """ SS 339 61-66 63.75 +0,12 64 1,10 North Sea, German coast (Ktndler 1941) AS 566 61-68 64,08ThoO8 64 1.04 """ " SS 274 6165 63.09 O,l2 63 1.00 Baltic Sea,German coast (KMncuer 1941) AS 584 6067 63,50 ,11 64 1.24 tt ti ti ti ti ti SS 961 6o..67 63.17+0,08 63 1.07 Portsmouth,S.Englandooast.Originaldata AS 100 62-66 63,94 0,18 64 0.94 t' t, 't t' ti i t' ss 200 6165 63.21 Th,09 63 0.82 Dingle, W. Eire coast, Original data SS 49 60-65 62,96 3,30 63 1.02

A.marinus

W, Greenland. (Jensen 1941) 38 67-72 69.39. 69 llovaya Zemblya (Kirillov 1936) 100 70.9 ±0.27 1.33 T4urman coast ( Kirillov 1936) 121 71,3 +0,16 0,87 Iceland Einarason 1951) 167 69-75 71.97 IcelandBruun1941) 97 68-73 71.53*0.23 72 1.15 FaoroesBruim1941) 129 68-72 69,67+0.17 70 0.17 Tana, N. Norway (Soleim 1945) 89 69-74 71.88 +025 72 1.18 Orense, N. Norway (Soleim 1954) 200 69-75 71,78+0,16 72 1.18 Fosnavag, Norway (Soleim 1945) 200 68-73 70.70 ±0.14 71 0.98 Bergen,Norway(Soleim 1945) 200 67-72 69.49+0,15 70 1,06 Scotland (Rai-tt 1935) 205 67-72 69.21+0,16 69 1.18 Isle of Man, Irish Sea (Cameron 1958) 240 66-71 68.71+0,14 69 1.09 Celtic Sea (corbin & Vati 1949) 45 68,56±0.26 0.86 Heligoland, N. Sea (KMndler 1941) 28 68-72 70.11±0'43 70 1,12 Loreley Bank, LSea (KLndler 1941) 39 67-71 69.49 ±0,28 70 0.86 }Iornsreef, N. Sea (Popp Madsen 1957) 216 70,20 JutlandBank,N. Sea (Popp Madsen 1957) 100 69,92 Kolberg, Baltic Sea (Kndler 1941) 217 6572 68.51+0,16 68 1,21 A. dubius

W. Greenland Jensen 1941) 179 73-78 75.10±0,16 1,10 W. Greenland. Einarason 1951) 66 7378 75.25 W. Greenland.Scott 1968) 39 73-78 75.030.36 1,11 Nova Scotia (Scott 1968) 146 71-78 74,60 0,20 1.23

*O spawning group not stated. (most counts in this category have been omitted.) AS = autumn spawning group SS spring spawning group FIRM S82North Atlantic Sandeels 1:7

TABLE IV

Intraspecific meristic variation; counts of the total number of vertebrae in Ammod,yies recorded by Richards et ai,(1963)

Number of vertebrae includes the urostyle

LOCALITY N Range Mean 5%C.L. Mode S.D.

Hudson Bay D * 7 71-74 72,29 ±1.07 71 1,20 I' 35 64-70 67.45 -f0,45 68 1,31 Labrador Coast 59 64-72 68.27 0.36 68 1.38 It D *12 71-75 73.50 +0.91 75 1.45 Lake Melville, Labrador * 9 62-65 63.20 1.38 62 1.30 Maqdalen Islands, Perce, Quebec 82 61-70 66.73 +0.37 68 1.70 Maine 65 65-70 67,91 +0.29 68 1.18 Massachusetts Bay D *34 64-72 69.27 +0,69 70 1.98 TarpaNlin Cove, Mass. D *14 67-71 69,36 +0,66 70 1.15 It It II *23 61-67 64.48 +0.65 64,66 1.50 Menemsha Bight, Mass. D 14 70-73 71.21 0.56 72 0.97 n n 'i *86 62-68 65.87 +0.36 68 1.66 Woods Hole, Mass. D 51 67-73 70.51 +0.35 70,71 1.25 " " 115 61-68 64,66 +0.30 65 1.64 Montauk Pt. Longlsland, N.Y. D 19 69-72 70.32 +0.43 71 0.89 " "" " 45 63-72 64.84 .35 65 1.17 Long Island Sound. 301 61-69 65.18 0.19 65 1.69 Island Beach, New Jersey D 98 65-73 69.48 +0.37 69 1.85

Per].mutter's data:- North of Cape 87 63-71 68.32 +0,28 69 1.34 ti ' D 76 72-78 74,46 0.28 74 1.21 South of Cape Cod * 94 60-71 66.50 Th.36 67 1.78 D 79 68-75 71.00 0.25 71 1.13

* Strongly skewed distributions

DThese samples were referred by the original authors to A. dubius, the remainder to A. hexapterus (. A. americanus) /S82 North Atlantic Sandeels 2:1

2. DISTRIJTION taxonomy of these fish is fully reviewed (see Section 1.22), 2.1 Total area Jensen (1941) recorded A. dubius from In the North Atlantic, Axnmodytes species West Greenland between the latitudes of 6l°N and have a boreai/boreo-arctic distribution, and are 70°N. The most southerly populations are over generally common, often abundant, over the shallow the Nova Scotia Banks at 43°N (Scott, 1968), sandy areas of the continental shelf within the although Richards et al. (1963), using a wider approximate latitudinal limite of 36°N and 73°N. concept of the species, indicated a southerly The total distribution of the genus is shown in limit of 39°N. Richards et ai. gave the Fig.2, and the distribution of individual latitudinal limits of A. americanus along the species in relation to adjacent land areas in mainland coast of North America as 36°N and 60°N. Table V. On account of their association with Both A. dubius and A. aniericanus occur in Hudson coastlines and shallow water (eg. Henderson 1961) Bay (Viadykov, 1933; Richards stai., 1963). distribution in relation to natural regions of 2,2 Differential Distribution the is probably of less significance; Suffice it is to say that the N.W, Atlantic species, 2.21 Spawn, larvae and juveniles A. americanus and A, dubius, are both to be found in Labrador waters (5.1.3.) Baffin Bay (5.1.4.), Spawn Hudson Bay (5.1.5.) and Newfoundland waters (5.2.1.) with the former, in addition, in Gulf Insufficient data is available on the Stream waters (5.3.2.); both the N.E. Atlantic demersal eggs for sii assessment of their species, A. tobianus and A. marinus are present distribution to be made. in the Barents Sea (5.l.2), the Irminger-Gyral (5,2,2,), the Norwegian Sea-Faeroese waters Larvae (5.2,3.), the North Sea, Irish Sea and English On hatching, the larvae become Channel (5.2.4.) and the Baltic Sea (5.2.5.), with planktonic, resulting in a potentially wider A. tobianus also in Atlantic Drift Current waters distribution than the adults; even so, Corbin t5.3»3.) and A. marinus probably in East Greenland and Vati (1949) failed to find post-larvae of waters (5,l.].). any sandeel species further than 100 km from the The recorded latitudinal range limits coast in the Celtic Sea area, and records from of A. tobianus are at 69°N (Murman coast; the North Sea and Scottish waters revealed a Andriashev, 1954) and at 36°N (Spain; De Buen, distribution more conveniently to be described in terms of an association with coastlines and. 1935). To the west, the species is present at Iceland, but only along the south and west shallow water than with water masses (Henderson, coasts (Einarsson, 1951); to the east it is 1961), present throughout most of the Baltic (KMndler, Larvae of A, tobianus were not found 1941), but does not appear to extend into either in the Celtic Sea area by Corbin and Vati (1949) the White Sea or the Mediterranean.Insufficient - their 'A, tobianus' larvae were later shown data is available to determine whether spring-. by Eiriarsson (1951) to be those of H,yperolus and autumn- spawning groups have different ranges; lanceolatus (Lo Sauv.) and Cameron (1958) both are present in the Baltic and North Sea noted their scarcity around the Isle of Man in (KEndler, 1941), in the Irish Sea (Cameron, 1958) spite of the abundance of juveniles inshore in off the west coast of Ireland (Fives, 1967), and the area, Fives (1967) however located larvae in the English Channel (present author), but in both spring and autumn off the Gaiway coast Einarsson (1951), recorded only a spring spawning of Ireland, and Einarsson (1951) found then in from the species at Ireland and Smitt (1893) refer-spring off the west coast of Iceland within the red only to an autumn spawning-season from the 50 metre depth contour, and mainly at 15-20 Norwegian coast, metres below the surface, In the North Sea, With a distribution ranging from 73°N Macer (1965) distinguished concentration of (Novaya Zemblya; Kiriliov, 1936) to 49°N (western spring larvae off the Thames, Wash, Huxnber and Dutch coastal areas, from autumn larvae which end of English Channel; Corbin and Vati, 1949), were mainly off the Dutch and French coasts; A. marinus is seen to have a similar, but more most of these larvae were well offshore, and range compared with A. tobianus. In northerly over areas from which the adults are rare or addition, it is present around all Icelandic coasts absent, (Einaa'sson, 1951), but is absent from the Gulfs of Botlmia and Finland in the Baltic (Diincker, It is the larvae of A. marinus which 1960). Whether the low meristic-count sandeels form the abundantconcentrationsof sandeel recorded from West Greenland to 72°N are to be larvae in the North Sea area in early spring as regarded as A. marinus(Jensen,1941) or as recorded by Bowman (1914), Kbndler (1941), A, americanus will remain unresolved until the Henderson (1953), Ryland (1964), andMacer (1965). 2:2 FIR1/S82 North Atlantic Sandeels

60

r.

s

X h

60

so ¡4

A A.fobianus A.dubius x1.marinus & I ¿. americanus

30

Fig. 2. Probable distribution of Amnìodvtes species in the North Atlantic,(Boundary between a.marinus and A.arnericanus is at Greenland; see Section 2.1), FIRN/S82North Atlantic Sandeels 2:' TABLE V The distribution ofAinmodytesspecies in relation to land areas in the North Atlantic

Jj E I . .., 0

211,13,14 LW.Territories,Manitoba + + Vladykov(1933) and Ontario Richardset al. (1963) 215 Quebec + + Leimand Scott (1966 216 Newfoundland,Labrador + + Leimand Acott (1966 217 NewBrunavick,Nova Scotia + + Scott (1968) 236 New England + Richards et al.1963 237 Mid-Atlantic States + Norcross et al. 1961 Richards et al.1963 250 Greenland + +? Jensen 1941 Su Denmark + + Jensen 1941 512 Faeroes + + Bruun 1941 513 Iceland. + + Bruun 1941 514 Norway + Smitt(1893) Soleim (1945) 515 Spitzbergen, Bear Island, + Andriashev (1954) Jan Mayen 516 Sweden + Smitt (1893) 517 Finland + Dllnoker (1961), Valtonon (1964) 521 Netherlands + + Roesaingh (1957) 522 Belgium + + Poll (1947) 524 Prance + + Purnestin (1939) Forest(1950) 531 Ireland + + Molloy (1966) Fives (1967 533 England, Wales + + Corbin & Vati (1949) Macer (1965 534 Scotland + + Raitt (1934 535 N. Ireland + + 536 Channel Isles + + Present.author 537 Isle of Man + + Cameron(1958) 541 Portugal + Gonalvee (1942) 542 Spain + Da Buen1935 561 Germany + + I'. dler 1941 572 Poland + Kndler 1941 716 Arctic Soviet Isles + Kirillov (1936) 720 Fiimo-Karelia + + Andriaahev (1954) 731-33Estonian, Latvian, + dler (1941) & Lithuanian S.S.R. 2:4 FIlIN S82 North Atlantic Sandeels

Macer also found large concerrtrations of young however, no indications of separate nursery areas, larvae in the Dowsing area and, in the Dover and any differences between the distribution of Straits (also recorded by Furnestin, 1939), and juveniles and, adults can probably be accounted there were larger larvae to 'the south of the for by the greater abundance and initial wide Dover Straits which, it was suggested, were dispersal of the former. from an unlocated. spawcing ground in the English Channel.Within the southern part of the North 2.22 Adults Sea, Macer (pers. comm.) recogeises three main A. tobianus is tpioally found close centres of larval distribution; a) the south-- inshore (i.e. accessible 'to beach seines) over west Doggerb) the eastern North Sea (in both sandy areas in bays arid. estuary mouths ,although of which, swirls minimise net dispersal) and Macer (1966) has recorded occasional individuals e) the Southern Bight, where a residual current 60km offshore in the North Sea.The species of up to 4km per day carries the larvae to the also occurs in intertidal sands. Autumn and north-east. spring spawning populations show sorne differences in distribution, but both appear to occupy a According to Corbin and Vati (1949), wide range of habitats and there is, as yet, no larvae of A, marmnus are absent from the Lands evidence of any differential habitat selection. End./Usharit area of the English Channel, but Along the south coast of England, spring'-spaw- are present in the Celtic Sea,Cameron (1958), ning fish are dominant in the Isle of Wight/ and, Fives (1967) recorded the larvae from the Portsmouth area, whereas only autumn-spawning Isle of Man and west coast of Ireland, respec- fish are present along the Devon and Cornwall tively, and Einarsson (1951) found the largest coasts (present author), and Knd1er (1941) numbers in Faeroese waters in the fjords and located the spawning grounds of spring-spawners between the islands where the depth ranged from in the mouth of the Elbe estuary, whilst those 50-150 metres.In the southern North Sea, of autumn-spawners were on the lorely Bank Ryland (1964) found the larvae nearer the surface (concentrated at about 10 metre depth) (Heligoland). in daytime than ai night. A. marinus is a widely distributed and. In the N,W. Atlantic, Serebryakov (1965)common species usually associated with offshore recorded large concentrations of sandeel larvae banks It is abundant in areas of the southern (as A. aznericanus) over the Newfoundland Grand North Sea, and Macer (1966; and pers. comm.) Banks, the Nova Scotia Banks and in the Gulf of distinguishes three main stocks: a) eastern Maine; it is likely that the majority of these (15-37 metre depth), b) western (27-37 metre larvae - the most abundant fish larvae in the depth over the S.W. Dogger, 12-22 metre depth I.C.N,A,F, area - belong to A, dubius.Richards over the Norfolk Banks) and. c) southern Bight. (1965) recorded larvae (as A. hexapterus and Molloy (1967) found large concentrations on the A, dubius) from Chesapeake Bay to West Greenland, Kish and Benriet Banks in the Irish Sea, and off and Wheatland (1958) and Norcross ai. (1961) Scotland, stocks are present off the west coast, have provided more detailed, accounts of the around the Shetland Islands and in the Moray distribution of larvae in Long Island Sound and Firtb and Aberdeen Bay areas (lIaitt, 1934; Chesapeake Bay respectively; in the latter area N.P. Langhan, pers. comm.). the larvae were most abundant further than 45km offshore. Scott (1968) has located considerable quantities of A. dubius over the Nova Scotia 0ff West Greenland, Einarsson (1951) Banks, although they were less common on the found the newly hatched larvae of A. dubius more north-easterly banks.lt is likely that mainly within 20-30 metres of the surface and the abundant stocks of sandeels on the Newfound- predominantly well within the 150 metre depth-- land Grand Banks (Templeman, 1966) also belong contour, to this species.A. dubius is exclusively an offshore species (Backus, 1957; Richards et al., Juveniles 1963; Leim and Scott, 1966). Young sarideels assume the demersal habit of the adults at about 30-40 mm; post A, axnericanus is found close inshore, larvae of this size become rare in the plankton but there are also offshore components.It and A. tobianus begins to appear in beach-seine would appear to be particularly abundant along catches at about this length. (Cameron, 1958). the New England coast and is present in estuaries, bays, and along exposed coasts (Richards et al., Juveniles appear to be found in the 1963, Leim and Scott, 1966). sane general areas as the adults, although in A, marinus (cameron 1958) and in offshore 2.3Determinants of Distribution populations of N.W. Atlantic Amrnodytes (Richards 2,1963), they may be present in inshore areas Ecological from which the adults are absent.There are, Sandeels are mainly to be found in FIRB/S82North Atlantic Sandeels 2:5

association with the sandy substrates into which from waters with a temperature between3°Cand they burrow (Masterman,1895;Popp Madsen,1963; 6°C.Similarly, the present author has found and many others), and although Kirillov(1936) A. tobianus active between6°Cand 16°C along the encountered A. marinus over weed and gravel south coast of England, but inactive below6°C areas, it appears in general, that roc1r, muddy in December, January and February; around the and, coarse gravel areas are usually avoided. Isle of Man where the winter temperature does not Several authors (such as Macer 1966) have des- fall below6°C,Cameron (1958) apparently found cribed the most favourable substrates as clean, this species active throughout the year. coarse sands or fine gravel, but no attempt has Holmquist (1958) recorded A, dubius larvae from been made to determine quantitatively the role pools in floating iceblocks off West Greenland. of different types of sandy substrate on

distribution. Populations of sandeels along the No information exists on the relation- - south coast of England have been found by the ship between distribution and salinity, but a present author, to be associated with substrates wide tolerance in A. tobianus is inferred from ranging from mean particle size 0.35 mm (Ø quartileits present in the Gulf of Bothnia, and in deviation0.55)to1.30mm (Ø quartile deviation estuarine situations elsewhere (eg. a spawning 1.0). It is possible that the total particle population exists in the River Fowey, S. England. composition and the way in which it affects water at a point where salinity ranges from 15% to32%). circulation and thus orgenation within the sub- Norcross . 1,. (1961)referred 'to records of strate, is a more important parameter than mean la,rvae of A. americanus in salini'ties from<1.8% particle size, per se. and -to suggestions that as the sandeel grows it becomes less tolerant of low salinity, For The main concentrations of sandeels A. americanus and A. dubius (see Section1.21) appear to be in shallow water; in the southern Richards et al,(1963)gave environmental North Sea A. marinus is most abundant in depths salinities of26-32%and30-36%respectively. of20-40metres (ÌIacer,1966),and. A, tobiaxius Behavioural and Aainericanus are usually in much shallower water than this. The greatest depth recorded is Sandeels have neither swimbladder, nor the isolated occurrence of A. dubius over the fins capab].e of compensatory movements, and in Nova Scotia Banks at 400 metres (Scott,1968) order to remain clear of the bottom the normal although most were taken between 40 and, 100 metres,myotomic swimming movements must be maintained.. Effective ener- conservation and some degree of Popp Madsen(1963)has suggested that escape from predation is accomplished by bur- in the southern North Sea, whilst sandeels are rowing into and resting within, the substrate. present in most places where the bottom is suitable,Since considerable periods of time may be spent the largest concentrations are restricted to the within sand and since no inhalent opening is edges of the larger banks and to the tops of the maintained between the buried fish and the smaller ones - particularly where the current surface of the sand, it is essential that the speed exceeds0.5ksots; Macer (pers. comm.) has substrate be well supplied with02 and strong however been unable to confirm such a relation- currents flowing over a substrate with large ship between distribution and tidal strengths. interstitial spaces would enable such aeration

to take place, Sandeels are thus able to alter-- Scott (1968) concluded that A. dubius nate between periods of inactivity within the over the Nova Scotia Banks was eurythernai; fish substrate and. periods of activity (feeding) in could be taken from water ranging from2C to the overhead water masses. 11°C although the largest quantities were taken FIRM 582 North Atlantic Sandeels 3:1

3, BIONOMICS AND LIFE EISIORY 3.15 Gonads Nacer (1966) has provided the following Reproduction 3.]. regression equation based on 35 A. marinus from the North Sea and the Faeroes: 3.11 Sexuality Fecundity 2.046 z (Length 3.055) All species are bisexual and, there are Other fecundity data are shown in The sexes no published recorde of hermaphroditisrn. Table VI together with the equivalent feou]adities cannot be distinguished by any obvious external calculated from the above equation. All are a characteristics. measure of annual egg production. 3.12Maturity The intraseasonal pattern of gonad Sexual maturity is attained, as follows:-maturation and the duration of the different stages is to some extent dependent upon the time Atobianue; at two years according to Cameron of year at which spawning takes place; according t1958) bui the present author has found, maturity to Knler (1941) the ova of spring-spawning at one year 'to be common in both sprizi- and A. tobianue take seven months to mature, those of autumn- spawning fish in the English Channel, ThisA. marinus (winter spawning) takes five months; involves fish as small as 83 mm, and. 60-70% of and those of autumn spawning A. tobianus take those reaching early maturity are males. only three months, In A, tobianus (Krid1er,l94l), A. marinue (Macer, 1966) and A. dubius (Scott, A. marinus; according to Cameron (1958) and Macer l98) the testes have been found to reach a State tl966) mainly at two years, but the largest of maturity (in terms of size in relation to individuals in a year class may mature at one the body cavity) before the ovaries, year. Macer gave 5%, 80% and 98% as the propor- tions spawning at one, two and three years respectively, in the North Sea. Kirillov (1936) considered that A. mainte from Novaya Zemblya first matured at three years.

A. dubius; one to two years (Scott, 1968).

TABLE VI Fecundity data; A. tobianus, A. marinus

FECUNDITY x io2 Range Equivalent fecundity calculated from Nacers (1966) equation (see Section 3.15)

a) A. tobianus,southernNorth Sea (K'dhl and Lumann, 1965)

10.5 1 3.2 2.5 11.6 2 4,1 3,7- 4.5 3,4 13.6 6 6.5 4,3-10.0 5.8 14.5 10 7.2 4.0-13 3 7,4 15.6 14 6,0 2.5- 9.6 9.1 16,6 10 8,1 4.1-16.3 10,9 17.3 3 8,1 6.4- 9.5 12.4 18.1 1 8.3 14.3

b) A, narinus, tTovaya Zemblya (Kirillov, 1936)

12.0 6 4,4 3,9

o) A, inarinus, Murman coast (Kirillov, 1936)

12,0 6 2,6 3,9 3:2 FIRM582North Atlantic Sandeels

3.16 Spawning A, tobianus (Thomopoulos,1954).Thomopoulos also suggested a lunar/tidal determination of spaw The unimod.al distribution of ovarian ning, for he found that peaks were reached eggs and the absence of redeveloping gonads in 3_-4 days before full moon mainly in October, A. marinus (Nacer1966)suggests that individuals but also in September and November, spawn only once during each spawning season; this is probably also true for the other species. TABLE VII The occurrence of larvae and condition of tha gonads as an indication of spawning season in Aranodytes species

+ occurrence of larvae less than lO mm in length R " ripe/running adults

Months J F M A M J JA S O ND

A. tobianus Baltic Sea (Kndler,1941) ERR ERR North Sea Kihl and LUhmann,1965) RE ER North Sea Macer,1965) + + + + + + ++ Irish Sea Cameron,1958) + + + Ireland, west coast (Fives,1967) ++ + + + + English Channel (Thornopoulos,1954) ERR l (present author) ER RERR

A. marinus Faeroes (Einarsson,1951) North and Baltic Seas (Knd1er,1941) ER North Sea (Nacer,1965,66) ER Irish Sea (Cameron,1958) Ireland, west coast (Fives,1967) Celtic Sea (Corbth and Vati,1949) English Channel (Furnestin,1939)

A. a.niericanus/A. dubius West Greenland (Einarsson,1951) ++ Nova Scotia Banks (Scott,1968) RE Long Island Sound, N.Y. (Wheatland,1956) ++ Chesapeake Bay (Norcross et al,1961) + +

The occurrence of ripe and running Spawning in A, tobianus has been gonads and the occurrence of small larvae in the recorded close inshore (Fullarton1894; plankton are summarised in Table VII, to indicate Thomopoulos, 1954; KNndler:1941),and Knd1er the spawning seasons. Although both methods have found that spawning occurred in spring in the obvious limitations, which preclude more detailed mouth of the Elbe estuary(5metres depth), and analysis, it is clear that A. marinus, A. americanus,in autumn off Ileligoland(iometres depth). and A. dubius spawn in the winter months, and that Cameron, however,(1958)was unable to locate there are two spawning seasons in A. tobianus, in spawning grounds of this species around the Isle spring and autumn, It can be shown from otolith of Nan, in spite of the abundance of young fish, studies in the latter species (Cameron1958;presentAccording to Furnestin (1939), the spawning of author) that fish spawn at the same time of year A. marinus takes place in depth of2O-4Ometres, as they wore initially spawned, thus indicating but Andriashev (1954) has referred to depths separato spawning populations, (see Section1.22) of 25JOO metres,In general it appears that spawning takes place within and not outside, the What effect any sequence of spawning general area inhabited by a population; that is of individuals has on the length of the spawning there are no indications of migration to specific season is unknown, but in any one area spawning spawning grounds. It is not known 'to what extent takes place over a period of about three months in spawning 'takes place within the sand substrate, FIRM/S82 North Atlantic Sandeels 3:3

The seo-ratio of A. marinus at the timecapable of a vertical migration (Norcross et of spawning is for A. marinus over the Dogger area al., 1961) and this may reduce the importance of the North Sea and has been recorded by Macer, of a particular system (eg, surface currents) (1966); it was found that males comprised 67% of in determining the distribution of the larvae. a sample of 120 fish, and this ratio applied to In the southern North Sea, Ryland(1964)found both mature and immaturo fish; this was in con- that during the day, larvae of A. marinus trast to the combined sex-ratio for all cruises were concentrated at 8 metres depth, but a-t between March and August in which males corn- night were more generally distributed between prised 45% of 2252 fish (xtest, p

A.marinus7.3mm.

A.tobianus9.7mm

A.marinus 11.0mm

'/1/ /,,.,

A.marinus14.0 mm

f ,-. o t;' ç'

A.marinus 19.0mm

Fig. 3. Pre-metaxuorphic stages of .marinus and A.tobianus (adapted from Macer 1967). FIRM/S82North Atlantic Sandeels 3:

ration, and thus, in this instance, the rate of countered as a result of otolith reading are appendicularian recruitment would have determined as follows:- the reality of a state of interspecific competition between plaice and sandeels. Covill calculated A, tobianus VII Original data; English that10-16times the weight of food actually Channel found in the average stomach (0.3% of body weight) A, marinus IX Macer, 1966; North Sea of A, aznericaxius would be necessary to supply the energy needed for growth and maintenance, and that A. dubius IX Scott, pers. comm.; Nova there would be no difficulty in fulfilling these Scotia requirements even at the low densities prevalent in Long Island Sound between January Insufficient data is available to determine and March, whether there is any intraspecific variation in T max (see also Section 4.12) The larvae do not appear to feed at night. Covill (1959) found that in A. aznericanus, 3.32Hardiness stomach contents were minimal in early morning, and after an increase towards mid-day, diminished A. tobia.nus and A. marinus can easily slowly into late afternoon.Ryland(1964) con- be kept alive in aquaria for periods of several cluded that for A, marinus larvae in the North Sea, months, arid P. lrlinslade (pers. corn.) has there was sii intensive2-3hour feeding period reported the spawning of A. marmnus under these beginning at first light followed by a diminished conditions at Lowestoft,A. tobianus can be intensity throughout the day and cessation at the kept alive in wet sand for at least six hours onset of dar1aess, (Cameron, 1958).Thomopoulos (1954) considered that male A, tobianus were hardier than females, 3,23Adolescent phase judging by their survival in captivity. Ammodytes species are much less susceptible to After metamorphosis at about40mm, injury from handling than are for instance the demersal habit is assumed and the fish become clupeoida, and are usually alive on capture adolescent. This period may be of short duration from trawl and seine, When used as live bait, in for example those spring-spawning A. tobianus they can remain alive in floating keeps which metamorphose in July and whose gonads begin ('courges') for periods of over a week, Sandeela to ripen in November (present author) but it is may survive ingestion to the extent of being probably normally of about 1- years, Adolscent able to penetrate the gut, enter the body cavity fish occur in the same areas as adults, and althoughand become encysteci in the liver of their pre- those of A, marinus (Kndler,1941;Cameron,1958) dators (Breder,1956), may, unlike the adults, be present inshore, there No critical work has been carried on the is no evidence to suggest the existence of specific limits of tolerance to environmentaL factors, but nursery areas. the way in which these may influence distribution is discussed in Section Observations on Adolescents have a more rapid growth- 2,3, captive sandeels and inference from field results rate (Section 3.43) and eat smaller food items suggests that these fish mey respond to periods (Section3,42)compared with adults, of food scarcity by remaining within the sand for longer periods than when food ia abundant, Adolescence in A, marinus begins in May in both the North Sea (MaceT) and the and conditions more favourable. Irish Sea (Cameron, 1958), Cameron recorded Competitors influxes of newly-metamorphosed A, tobianus in 3.33 May-July and November at the Isle of Nan, Other plankton feeding fish, such as corresponding to the spring and autumn broods clupeoids are frequently caught with Amwod,yt respectively; from localities along the south species (for example with A. marinus in the coast of England, the equivalent influxes have North Sea; Macer, been recorded in July and April by the present 1966)but there is no evidence for food competition in such instances, The author, four species of Ammody-tes in the North Atlantic 3,3Adult Phase are separated either geographically or by virtue of inshore/offshore habitat selection, thus 3,31Longevity precluding the possibility of competition. In the N.E. Atlantic, another plankton feeding Assuming ari annual mortality rate of sandeel, ymnammod'-tes semisquamatus may occur 70% (see Section 4.41), the average expectation of with A. marinus as in the North Sea: Macer further life is about years. (1966) 0.9 has suggested that potential competition for food between these two species may be minimised The maximum age groups (p max) en-. by the fact that the latter are concentrated on 3:6 FIRM/882 North Atlantic Sandeels the tops of banks, whereas the former are mainly poorcod (Tnisopterus minutes (L) in ApnilJune found between the banks, and by cod in OctoberDecember (Nagabhushanam, 1965).Predation by plaice (Pleuronectes 3,34 Predators platessa (L)) may be important locally (Todd, 1914) and for small halibut (Hipo;lossus From a review of the literature it is hippoglossus (L))sandeels are a major food apparent that9 throughout their range, sandeels both off Iceland and. the Faeroes (McIntyre, 1952). are heavily exploited by a wide variety of verte- Roessingh (1957) has recorded them in the diet brate predators,Because however, such informa,- of soles (Solea vulgaris) in the North Sea, and tion results almost entirely from trophic studies B.B. Rae (pers. comm.) has noted their importance of the predators themselves, it is impossible to to turbot (Scophthalmus maximum (L) ), draw any quantitative conclusions on the influence Spurdogs (squalus acanthias (L))feed heavily of predation in controlling the size, density, andon sandeels in certain areas (Holden, 1966), size composition of the sandeel populations, and those species of ray (Raia species) Sandeels have been occasionally recorded in the inhabiting sandy areas in relatively shallow diet of pinnipedes and small cetaceans (Bigelow water also eat sandeels,A, tobianus is a and Schroeder, 1953), and such predation may be favourite bait for bass (Dicentrarchus labrar heavy locally, for example in grey seals (Hali- (L)) in inshore waters, and sandeels are often choerus j.us (Fab.)) off SW, Shetland Isles present in their stomachs, (Rae 196; Particularly heavy predation by breeding seabirds in the summer months will be The above records, additional observat- significant for sandeel populations within the ions and also more °eneral works on a wide feeding range of colonies;Pearson (1968) has variety of species reg. Sim, 1884) suggest that reviewed the literature on seabird feeding and most piscivorous marine vertebrate readily eat described the trophie ecology of the Farne Island eandeels, - often in considerable quantity, seabird colonies off the east coast of England; Individual sandeels have no physical means of Aminodytes species were the dominant food and it defence and in fact, their size and shape (an was estimated that 943 metric tons were consumed adaptation to burrowing) render them ideal food within the colony in a single season,The size items for most predators,It is maintained that of sandeels taken ranged from 25-175 mm and sandeele burrow into the sand to escape predators, thus included larvae, adolescents and adults, but but this is certainly notaninevitable reaction most were adolescents of 50-75 iran; the major to the presence of prescence of predators, and predators were the surface feeding terne (Sterna there is no information to suggest how often it species), and (Rissa tridactylat1fl7 happens and how effective the behaviour is: and the diving aulzs (Alciciae) and shag (Phalacroc-certainly, even when in the sand they are orax aristotelis (L)).Predation by other fish probably still susceptible to predation by such has led to concern over potential competition for fish as Raia species. sandedresources between the industrial fisheries and stockir of foodfish species (for 3.35 Parasites, diseases, injuries example Roessint, 1957).There is no evidence and abnormalities to suggest that this is happening, however, and both Nacer (1966) and Rae (1967) have noted the Polyanski (1955) reported seven species relative unimportance of sandeels as a food of parasite from A. maninus in the Barents Sea; resource in the southern North Sea - the major area where industrial fishing for sandeels has Sinuolinea murinanika Basikalova,(Protozoa taken place,In other areas, such as the lTova Brach,vphallus crenatua (Rudolphi), (Digenea Scotia Banks,(Scott, 1968) Faxe Bay, Iceland Dero,enes vanicus (Müller). (Digenea) (Brown and Cheng, 1946; McIntyre, 1952) and off Scolex polymorphus Rudolph. (Cestoda) the north coast of Scotland (Holden 1966; Rae, Coniracaecuni aduncum (Rudoiphi), (Nematoda) 1967) predation on sandeels is undoubtedly heavy, C, aduncum larvae particularly in the summer months,Cod (Cadus Anisakis sp, larvae, (Nematoda) callanas L.) eatsandeels over a wide area Echinorhynchus gadi Zega. (Acanthocephala) (Scott, 1968; Rae, 1967; Brown and Cheng, 1946) and often intensively - as off the north coast The highest incidence of infection of Scotland where, according to Rae, 70 - 85% of ( 7 out of 21 fish) involved C. aduncuni larvae, cod of all sizes eat sandeels throughout the year, although no more than three individuals were Whiting (0donto'adus menlangue (L), (Jones, 1954), present in any one fish; only the presence of is also a major predator, but (Nelanogram- S, murraanika resulted in a high intensity of mus aegefinus (L)) predation is more local infection. itchie, 1932; Brown and Cheng, 1946; Jones, 1954). The heaviest inshore predation by gadoids off the There are no records of effects of Isle of Man was by whiting, pollack (Pollachius infestation upon mortality, o11achius (L) ), coalfish (p. virens(L)Jj FIRM/S82 North Atlantic Sandeels 3:7 3,4 Nutrition and Growth that the food was similar to that described by Covill (1959) for the postlarvae in the same 3.41 Feeding area,Ten species of pelagic com- prised most of the diet and it was found that Roessingh (1957) found that most of 80% of the fish were feeding on Centropagen the stomachs of A. inarinus in the North Sea that species, 55% on Arcartia species, 42% on Tenora were examined in the early morning were empty, longicornis, 79ÇnBalanus balanoides cyprid whereas they were wellfilled in the afternoon larvae and 60% on the nauplii,Fifty per cent and evening; Richards (1963) inferred that had fish eggs in their stomachs, and. algae A. ainericanus in Long Island Sound began to feed (dinoflagellates and diatoms), the mysid soon after dawn.Underwater observations on Neornysi s americana, and Axnmod,y-t es aznericanus A, tobianus in the Baltic Sea (KUhlmann and larvae, each occurred in at least 10% of the Karat, 1967) showed that the ¡nain feeding period stomachs, was in the morning, with a secondary peak towards evening.Popp Madsen (1963) related the feeding In the N.E, Atlantic, Bogarov et al. activity of A. marinus in the North Sea to current(1939) recorded a wide variety of food species strength and has suggested that the fish are from Ammod,ytes sp. along the Murman coast, most active when a strong tidal current replenishesincluding such freshwater organisms as insect food. supplies over the banks.Macer (1966) larvae from Sedlovatyj Island.; copepods were considered that A, marinus in the North Sea feed however the most important food. group. Roessingh near the bottom because they are commonly caught (1957) found. main]y copepods and cladocerans in with their stomachs full in bottom travis; diet the stomachs of A, niarinus from the North Sea, composition, however, indicates that feeding and in addition identified the eggs of , takes place in the water mass and not in the Copepods also dominated the diet of A, marmnus substrate,Richards (1963) inferred from the in midsummer in the eastern North Sea, according presence of sand, in the stomachs of A, anericanus,to KUhl and. Luhinanrz (1965), but in contrast they that at least some feeding takes place close to found that the stomachs of 17 out of 22 fish the bottom,KUhlmann and Karat (1967), from caught in the Elbe estuary in April, were full underwater observations in the western Baltic, haveof the diatom Coscinod.iscus; Popp Madsen (1963) described the feeding behaviour of A. tobianus, has recorded a similar phenomenon.Macer (1966) The normal swimming schools become more or less found copepods in all individuals of A, marinus stationary as feeding begins, and spread out both(North Sea) that were feeding, the most important vertically and, radially thus often reaching genera being, in ord.er, Temora, Calanus and. from the sand, to the water surface; each individual,Psend.ocalanus:in addition, invertebrate and with the body atanangle of 15° to the horizontal,fish eggs, larvae, newly metamorphosed searches for food independently of the others, polychae'tes and. amphipods were important food seldom snapping for longer than 20 - 30 seconds items,Macer found that the diet of A, tobianas in one place. was broadly similar to that of A. marinus There is little information on although the absence of crustacean larvae and seaaonal variation in feeding,Richards (1963) the greater frequency of amphipods, mysids and. recorded intensive feeding by A. ainericanus in cladocera in the former species, were attributed. winter in Long Island sand, followed by a decreaseto its more inshore habitat (Table VIII); in early spring.Along the Murmazi Coast, Bogarov et al. (1939) found that the intensity of feeding Roessingh (1957) found food. items up in Amniodrtes ap. increased from the beginning of to 1.5 mm long in A, marinus; Richards (1963) June to August and decreased towards October,In recorded. copepod.s 1.0 - 1,8 mm long in the North Sea, KUlil and. Ltthrnann (1965) recorded A, arnericanus but mysids also occurred. in the feeding in April, May and June. At Portsmouth, diet and these were from 4.4 - 13.7 mm long. South Rngland, the present author has encounteredInsufficient data is available for the assessment full stomachs in A. tobianus from late March to of geographical variation in diet within a early November; in the winter months, however, species, but Richards (1963) has drawn attention fish dug from the intertidal sand usually have to the seasonal variation in the copepod empty stomachs.The capacity of sandeels to composition of the diet of A. americanus between ensure starvation has been indicated by moue et January and April, and Macer's (1966) data al, (1967) who observed that A. personatus was has shown how diet varies with the size of the able to survive 24 weeks in aquaria without food.fish,(Table VIII). 3,42 Food. In a study of the diet of 290 individuals of A, americanus (74 - 132 mm in length) from Long Island Sound, Richards (1963) noted 3:8 FIRN/582 North Atlantic Sandeels

TABLH VIII The ocourrenc,e offooö. items in the diet of A. marinus and. A. tobianus from the Southern North Sea, as weigh-ted mean %. (from Macer,].96F

Species A. marinus A, tobianus

Length (cm) <10 10.-20 >20 <10 10-15

No0 of stomachs 59 207 30 20 21 Number empty 7 40 17 0 3 Mean fulbiess 69 50 17 91 49 Sand. grains 15.3 9,6 8.0 30,0 27,5 Fish scales O 0 0 0 50.0 Fish larvae 1.9 3.1 8,0 0 0 Fish ova 9,6 14.8 15,0 0 0 Invertebrate ova 36.3 30.2 8,0 35.0 0 Copepodß 96.6 84.7 54.0 100.0 50.0 Calanus 30.9 22.8 30,0 5.0 0 Temora 69,4 64.1 23,0 60.0 33.0 Centropages 5.9 5.6 0 15.0 0 Pseudocalanus 17,5 27,4 15,0 15.0 5.8 Acartia 0 0.5 0 50.0 0 Caligus 0 0 0 15.0 0 Crustacean larvae 23,2 43.9 15.0 0 0 Cyprid. 23.2 30.8 0 0 0 Zoea and. Megalopa 2.0 23.8 15,0 0 0 Amphipods 11.6 2.3 23.0 35.0 5.8 Mysida 1.9 0 0 15.0 5.8 Euphausid.s 1.9 3.0 0 0 0 Isopoda 2.0 0 0 0 0 Cumacea 1.9 0 8,0 0 0 Cladocera 0 2.6 0 10,0 0 Chaetognatha 9.7 5.5 15.0 0 0 Laru-acea 3,9 11.8 15,0 0 0 Molluscs 3.8 0.5 0 0 0 Annelids 27.1 45.7 70.0 30.0 44,2 Echinoderns 0 0,6 0 0 0 Coscinod.iscus 11.8 4,7 0 0 0 Miscellaneous 0 1,1 8,0 15.0 5.8

3.43Growth rate between populations of the same year-class on different banks in the southern North Sea are Published growth studies on sandeels given in Table XVI; in addition, the growth of have mainly involved the presentation of mean the 1959 year-class on the Haddock and lengths of successive age-groups, and. very little Indefatigable Banks was so slight compared. with attention has been paid to growth in weight, intra.- the two subseq-uent year-classes and with that seasonal growth, and the analysis of growth para- of the same year-class on the Southernmost Rough, meters, Age has been determined by otolith that in 1961 and.1962,the 1959 and1960 reading, as initiated by Bahr (1935) and. Knd.ler year-classes were of a similar length, (Macer, (1941).Data on mean length of successive age 1966). Macer has attributed such intraspecific groups is summarised. in Table IX; it is apparent differences in growth rate to the presence of that most growth takes place in the first -two different current regimes supplying variable growth periods and that there is considerable densities of food to the different populations. intraspecific variation, Differences in length FIR1/S82 North Atlazitic Saa2deels 39 TABLE IX Mean lengths (mm,) of age-groups; AtolDianus9 A. marin,sA, dubius Month of Locality sEmples Author 170 W. Baltic Sept 0Jui-Aug. K.udler (355) (17) (respectively) (1941)

130 170 sie of Man (30) (4) Irish Sea)

144 153 Portsmouth 1.93 10,8412.3213.27 (S ,Engi and) (225)

112 136 4.91 8 88 11.76 (97) (36)

177 (3)

A. marinus

Macer (173) (1966)

A,dubius Nova Scotia Scott Banks (1968)

Boundary between agegroups Dec. 31st/Jan. ist 3:10 FIRM/S82North Atlantic Sandeels

Macer (pers0 comm.) has calculated VonA. marinus in the southern North Sea, (Macer, Bertalanffy growth parameters for A, marinus in the1966),Ii Amnodytes species the spawning periods North Sea as Loo (max, length) =21,8cm, K (rate generally fall outside the growth period, at which Loo is approached) =0.89, The small number of agegroups which are usually present Condition (ponderal index) has been however, render the calculation of such parameterscalculated for A. tobianus by the present author, difficult in most sandeel, and the seasonal changes in condition are indicated in Table X, The fish are generally In the Irish Sea (Isle of Man), unavailable when condition is likely to be lowest, Cameron(1958)recorded little or no growth in, but the total inqiidual range found, (where A, tobianus and A, marinus between October and condition = w/L '' x 105) varied from86to169, March, and rapid growth between March and July; In A, marinus, seasonal changes in fat content this pattern of growth also applies to A, tobianus are apparent from the work of LOhmann(1957) from English Channel localities (Fig.4Tand to (Table XI),

TABLE X CONDITIOII flTIflX(U/(L3'2xio)) in Sprinsoawning A. tobianus; Portsmouth, England (original data)

AGEGROUP APR. MAY, 313M. SEP, OCT. NOV.

Ogroup1969

Igroup1970 97,3108,8122,4134.4139,2 (3) (60) (278)(279) (206)

Igroup 1969101,7 118,3127,1 143,0132,3 145.8153,7130.0136,6 - 138,7 (29) (24) (48) (47) (so) (16) (9) (is) (sG) (3)

Igroup1970 118.3 135.7 (4)

Sample numbers in brackets

TABlE XI Percentage fat content inA. marinus (from Luhrnann,1957)

Range of sample means No, of samples

March 1,1 1.2 2 April 3.3 5.6 4 May 2,6 -42.4 9 June 6,0 -423 12 July 6.0 7,5 10 August Sept cmb or 7,0 1 October November 5,0 1 December 3,7 1 FIRN/S82 North Atlantic Sandeels 3:11

A. JO B IA NUS spring spawning 1n th PORTSMOUTH 1968-class

I e n gt h 1969-dass 9

IB

7

Io

'4

water-temperature f

IO \/\ 5 i 9 E o 8 weight I4 1968-class H o z w 2

o __--- - co' 1969-class J FMAMJU ASONDJ FMAMJ 1969 1970

Fig. 4. Seasonal growth of A,tobianusPortsmouth, S. England.(mean lengtn and weight with 95% confidence limits). Original data. 3:12 FIRN/582 North Atlantic Saxideele

3,5Behaviour populations not associated with either recruit- ment or movement to wintering area3, 3.51 Migrations and Local Movements 3,52 Schooling Kndler (1941), Anth'iashev (1954), Pearson (1969), and others have recorded influxes The formation of schools in sazideel of sandeels towards the surface coastal waters species is apparent from surface observations in summer, and Kndler has also stated that for example, Cameron 1958), echograznmes sandeels move to deeper waters offshore in winter,(Roessingh,C 1957; Dreyer and Ellis, 1968), and Both Caneron (1958), in inshore waters around the underwater observations (KUhlmann and Karat, Isle of Man, (Irish Sea), and Macer (1966), 1967) .The followinaccount is taken from offshore in the southern North Sea, were able to K'iihlmann and. Karat, (1967) and refera to under- catch sandeels in all months of the year, but in water observations (mainly on Atobianus) in general very few free-swimming sandeels could be the western Baltic.Although H. lanceolatus, caught in the winter months and such catches were A. tobianus and. occasionally young herring mainly composed of immature fish, Cameron in fact occurred. together in the same school, the fish found adult A, tobianus (and. all A, marinus) in were generally segregated into those containing Port Erin Bay only between May and August, individuals of approximately the same size. Close inshore, the number ofndividua1s in a From the fact that large catches of school was of the order of 10 fish, but, in A. dubius were made in both July and November at deeper water, schools containing lO3fish were the same places over the Nova Scotia Banks, Scott often encountered.School-shape was described (1968) suggested that there was little, if any, as horizontally flattened, and blunt-linear in seasonal movement of the stocks, but that surface view, but close inshore this pattern occasional reports of temporary local concen- was often disrupted, and other shapes developed, trations may have resulted from such movements In a normal swimming school the lateral distance or from undetermined hydrographical factors, between individuals was found to be about two- Popp Madsen (1963) concluded, on the basis of thirds of the body length, and the front/rear tagging 858 A, marinus in the southern North Sea distance about one-third of the body length, in the summer of 1958 (13% recovered), that when escaping, however the schools tightened sandeels are very stationary and, undertake no up and when feeding the distances between feeding migration; his experiments however, did individuals became greater,It was estimated not extend throughout the year and, the results, that the fish usually swain at a speed of 30-40 though of value, oanriot be used to determine the cm/sec; small groups swam faster than larger extent to which eandeels move to favourable winterones and in short escape bursts, speeds of areas, In addition to such movement it appears 300-500 cm/sec were reached,Avoidance of a that sandeels remain within the substrate during pursuer was mostly achieved, by accelerating to winter, and Cameron (1958) has produced evidence one side and avoidance of weeds, rocks and large to this effect; she found that although sandeels fish in the path of 'the school, by splitting and could not be trawled. on an offshore bank in reformation,Schools of fish of a similar size winter, they could be caught in a modified were observed to amalgamate freely, and there scallop dredge.Apart from the instance recordedwere both accidental and apparently spontaneous by Hansen (1949) of shoals of cod feeding on instances of disassociation.In general, it has sandeels which were apparently moving northwards been found that at night, sandeels cannot be along the West Greenland coast in August, there caught in either beachseine (Cameron, 1958), are no published accounts of a definite migration mid.=water trawls, (Macer, 1966) or bottom trawls, in sandeels, and while there is some evidence for (Popp Madsen, 1963; Macer, 1966),Bertelsen an inshore (summer)' offshore (winter) movement, and Popp Madsen (1958) found that it was much particularly in northern latitudes, this may be easier to catch sandeels in bottom grabs at obscured by movement into the substrate during night than in the daytime, and concluded that urifavourable seasons. Macer (pers. comm.) has they generally spent the hours of darksess suggected that at least for certain North Sea buried in the sand,However, both Anon (1958) stocks, the necessity of some sort of migration and Dreyer and Ellis (1968) observed, by sonar, is indicated for their geographic maintenance; the movement of sandeel schools towards the this would most likely be by juveniles at surface at night in N.W. Atlantic areas.Macer recruitment, and the decreases in mean length (1966) has reported records of nocturnal of A, marmnus on the Indefatigable and Haddock observations of sandeels at the surface of the Banks in the North Sea in 1961 (Macer, 1966) couldNorth Sea in March, and Bertelsen and Popp Madsen possibly have been the result of prolonged (1958) have referred to catches of sandeels at recruitment bringing about influxes of smaller night in ring-trawla, These isolated records fish,Other changes in population structure and of nocturnal activity may have resulted from abundance, incompatible with the results of the effects of ship's lights (Macer, 1966), or individual growth and mortality may also be moonl4ht, and it is thus of interest to find indicative of some degree of movement in sandeel that Kuhlmann and Karat (1967) were able to FIRN/582 North Atlantic Sandeels 3:13

attract A. tobianus from the sand at night with states of the tide, and although there was some an underwater torch, and, that moue . (1967) slight suggestion of relatively low catches at have reported the successful attraction of high water and on the ebb tide, he found no A. personatus to a lamp at night for fishing clear correlation between peak catches and, purposes. either the state of the tide or the time of day. The implication of observations associated. with In inshore waters of the Baltic, seasonal and diurnal activity cycles is that KUhlmann and. Karst (1967) have made underwater in general terms, sandeels burrow into the sub- observations on the diurnal behaviour of strate at night and during unfavourable seasons. A, tobianus. They found, that the fish emerged. From the observations reported by Kihlrnann and from the sandbank in small groups at sunrise, Karat (1967), entry into the sand is very rapid, and that these groups gradually caine together and simply appears to involve a continuation of to form large schools of more than 1,000 the normal sinuous swimming movement.Usually individuals which then swam across an area of the fish lie completely buried in the sand, but sea.-grass to the feeding grounds about 1,000 metreson being disturbed may protrude their heads in from the vicinity of the inshore sandbank. Aroundthe direction of the disturbance: this may then midday the fish returned from the feeding groundsbe followed by either a rapid sideways movement and throughout the afternoon the schools swain out of the sand. and rapid swimming away from about close inshore only occasionally stopping to the disturbance, or retreat into the sand. and feed. With increasin d.arksess the schools con- reappearance some distance away. gregated over the sandbank and small groups separated off and rapidly dived into the sand. 3.53Responses to stimuli Popp Madsen(1963)has suggested that activity in the daylight hours is associated With the state The importance of light in relation of the tide, since he found some correlation to the diurnal activity cycle has been implied in between peak catches on the North Sea fishing Section3.52. Experimental work on the grounds and the strength and direction of the activity of A. marinus in relation to light tide; most fish were caught when the tide running intensity, food and current strength is from the N.W. in the central North Sea, but two currently in progress at Lowestoft. peaks, at both stages of maximum tidal flow, were identified in eastern areas such as Hornsreef. Macer(1969),however, found that in the southern North Sea maximum catches could occur at all FIRM/382 North Atlantic Sandeels 4:1

4. FDFUIATION 7 -9 year classes may therefore be present. The most recent 1 - 3 year classes invariably 4.1 Structure dominate however, and their importance can be seen from the age group compositions in Tables 4.11 Sexratio XIII and XIV, The pattern of recruitment of the most recent year class is mainly responsible Overall sexratios are shown in Table for the seasonal variation in age composition, XII and in most cases indicate proximity to a but there is some evidence, in A. tobianus, that 1,1 ratio. Some individual samples recorded by the oldest fish only come close inshore in the Bahr (1935), Ktndler (1941) and Nacer (1966) show summer months. Macer (1966) has noted the significant deviations both during and outside effect of variable year class strength in the spawning seasons (see Section 3.16), and A, marinus on the age composition of North Sea could indicate the presence of behavioural populations. Sandeels are first captured as differences between the sexes. There is no O and I group fish, reach maturity as I - II evidence for sexratio changing with age and group fish, and, reach a maximum age as VII - influenced by differential mortality, IX fish,

4.12 Age composition 4.13 Size composition

The oldest A. tobianus recorded are The length composition of selected VII group, and the oldest A, marinus and A, dubius, samples of A. tobianus, A. marinus and A. dubius IX group; since juveniles recruit to the demersal is given in Table XV, The large size of population and to the fishery as O - group fich, A. dubius compared with the N.E. Atlantic species

PABLEXII Overall Sexratios in populations of A. tobianus and A, marinus (Monthly samples combined)

N Locality Author

A, tobianus 828 45,9 54.1 Western Baltic Bahr (1935) 1396 50.7 49,3 Western Baltic XBndler (1941 1444 51.8 48,2 North Sea XVndler (1941 401 52.2 47,8 South Coast, Original England data

A, marinus 397 30.8 69,2 Southern Baltic Kndler (1941) 2252 45,0 55.0 North Sea Macer (1966)

TABLEXIII Analysis of research cruise catch data from the Southern North Sea 1960 - 1962 to indicate the importance of different agegroups in A. marinus (from Macer.'s(l966) data)

Age group O I II III IV V VI VIIVIII IX Total number of cruises = 23

17 lOO 95 95 87 7074 48 13 % of total cruises when age group present - 48 30 17 4-- - % of total cruises when age group dominant 5 99 So 90 55 10 5 5 Maximum % of the age group in the total catch of the species from any one cruise 4:2 FIR1/S82 North Atlantic Sand.eela

TLBLE XIV

Percentage composition of age-groups in samples of A. tobianus

AGEÇW0UP* O I II III V VI VII N MONTHS LOCALITY AUTHOR

A, to'oianus 99,2 0.8 252 March-AprilIsle of ManCameron tspring §pawning) 68,9 28.9 1,8 0.4 284 June-July (Irish Sea) (1958) 89,3 10,0 0.7 364 Sept-October 8,4 73.8 11,4 3.0 3.0 1,0 0.5 202July Portsmouth Original 93.8 4,4 1,1 0.40,1 0.10.1 612October (S.England) data 100,0 116November

A, tobianus 76,9 19.2 3,9 753July-August Western KNndler (autumn Baltic (1941) spawning) 86.3 13.7 - - - 1021March-April 92,6 7.0 0.3 - - 841 June-July Isle of ManCameron 91.3 8,2 0.5 - - 537October- (Irish Sea) (1958) November 15.3 69.7 9.44.0 1.30,10,2 681May-Dec. Fowey RiverOriginal (S,England) data

* Boundary between age groúps is 31st December/lst January.

is conspicuous; further, A. marinusappears to be group individual year-classes of A, tobianus slightly larger than A, tobianus but the samples have been found to range from4-7mm by the are not directly comparable because of the present author. Size at first capture is dJ.fferences in gear used, Macer's (1966) research considered in Section 5,42, and size at first cruise data on A. marinus in the Southern North maturity in Section 3.12. Although precise Sea are similar to those based on the commercial information is lacking, it is clear that with catches, although the total range in length the gear commonly inuse, fish begin to be caught (5-25 cm) and the range in modes (9-20 cm) are before maturity. Concerning the maximum sizes greater, Samples of sandeels are generally reached, A, dubius is clearly distinct, with characterised by 1 - 2 modal peaks indicating individuals as long as 37 cm, In the other the dominant year classes, and marked seasonal species, the maximum recorded lengths are:- changes in these are determined by the rapid A, tobianus, 28 cm (present author); A. marmnus growth in the younger fish, and the pattern of 25 cm (Nacer, 1966; Fives, 1967) and. recruitment, Conspicuous geographical differencesA, americanus, 22 cm (Leim and Scott, 1966). in the length of the same year-class have been There is undoubtedly intraspecific variation, and indicated by Macer (1966) for A, marinus in the in the case of A. tobianus, exhaustive work by Southern North Sea (Table XVI); Scott (pers. comm.)Cameron (1958) around the Isle of Man, and by the has found similar size variations over the Nova present author at Portsmouth, has failed to Scotia Banks for A. dubius, Mean lengths for reveal individuals greater than 20 cm and 17 cm age are given more fully in Section 3.43, respectively; the larger individuals were from Standard deviations for the lengths of O and I the Fowey River, S. igland, Length-weight relationchips are summarised in Table XVII, TABLE IV A.tobianus - 63 freeuenc.y in cm. groupa (lengths given are the lower limite of_te_pm. groups) 00. Length conposition of seaples of A, tobianus, A. sarinue and A. dubius 3 4 5 6 1.97 7.7 8 8.49 1.510 0.211 0.4 12 13 2.5 10.3 14 9.515 16 17 2.33.7 0.7 j 0.818 19 I 20 21 j 22 Autumn-spawningSpring-spawning23 24 25 461No. Months 0. Baltic Locality Mixed Gear ICSndler (1941) Author 4.5 0.36.7 5.92.4 14.7 17.3 10.1 0.60.2 0.22.82.08.4 6.44.90.39.3 21.3 5.12.07.2 13.9 6.82.05.9 10.3156.20.9 7.71.67.00.8 12.5 0.62.3 19.3 0.40.90.7 10.6 0.40.2 1.8 0.2 Autumn-spawningSpring-spawning 455698 April-JuneSept-Oct.N-June - Religoland.(North(German1f. Baltic Sea) coast) Mixed ICNndler!CSndler (1941) 1.6 61.3 0.7 - 5.41.7 - 0.5 - 0.11.2 29.5-- 2.0 11.845.7 - 11.10.72.7 2.11.1 - 0.10.61.8 1.0- 0.1 - 0.21.01.2 0.48.2 - 0,13.40.5 0.40.5 0,2 - 0.2 - 0.2 - Autumn-spawningSpring-spawning 711403 Nov.June 1955-57 Isle(Irish of Sea)Man Beach-seineBeach-mains CssieronCanoron (1958) (1958) 0.6 3.0 6.01.51.2 47.612.9 0.3 19.621.9 0.6 0.20.42.47.8 25.5 11.40,60.6 0.47.6 0.87.84.2 0.63.63.9 0.22.01.5 0.20.6 Spring-spawningAutumn-spawningSpring-spawning 491333 Oct.July 1969 Portsmouth(S.Nnglam&)(S.England)(Irish Sea) Beach-seine Original data 6.0 63.8 0.8 29.3 3.7 15.7 0.8 6.0 0.8 2.8 20.2 28.4 12.1 3.6 4.4 0.4 0.4 Autumn-spawningSpring-spawning 246216 July-Aug.Nov.1969 1969 (S.England)FoweyPortsmouth(S.England.) River Beach-seine Original data A.marinus - 63 frequency in cm. groups 0.2 6.3 0,1 A.CIUbSUB 63 frequency in cm, groups

2.1

0.2

21.9 20.00.1 23.8 9.20.7 0.8 13.9 5.31.6 7.8 0.1 28.4 6.77.52.0 18.2 8.3

16.6 23.8 20.1 3,9 11.9 4.71.4

19.1 5.6 21.8 11.1 2.54.7 34.7 1.7 17.34.77.6 19.8 7.1 13.3

15.0 1.6

2.01.3

3.8

3.7 5.0

3.0 6.4

8.9

0.7

1.0

1.2

1.0

1.4 0.1 2.3 5.11.3

0.1

0.2

0.5

0.4 0.11.0 2.8

0.1

0.1

1.0 0.1 0.6 0.2 584040661986 0.1 4179

7573

5963 May-JulyMay-July 1960 1959 25292494 933 May-July 1961

May-July 1962

May-July 1963

Nay-July 1964

May-July 1965 S.North Sea May-July 1966 July 1967 S.North Sea

S.North Sea

S.North Sea

S.North Sea

S.North

S.North Sea

S.North Sea Ccssercial Noray Finth trawl (i6.Sootland) Commercial trawl Commercial trawl Cossercial trawl Commercial trawl Commercial trawl Commercial trawl Commercial trawl Mayer-llasrdenMeyer-NaardenMeyer1faarden(1962)(1961) Resemrch (1963 trawl Meyer-Nasrden (1964) Meyer-Nasrden (1965) Meyer-Waarden (1966) Meyer-Waarden (1967) Tiges (1968) 0.2 19.6 2.1 11.5 1.8 27.1 8.0 MP Langliam 6,00.31.1 (unpublished data) 16.0 3.3 42.3

PIRM/s82 North Atlantic Sandeels 26.7 4:3 23.013,0 7.4 27.8 8.8 23.6 17.1 25.3 22.3 11.411.0 21 cm. No. Months Gear I 13 - 1.314 2.615 5.316 17 18 5,3 10.7 4.019 7.920 7,3 5.622 10.5 23 13.4 24 10.0 25 10.0 26 27 14.7 15.8 10.028 29 5.330 6.031 2.632 .-- 3.933 34 - 35 - 36 - 0.6 149 July-Aug.1967 Nova Scotia Locality Author Banks Researchtrawl I Scott (1968) Ïs82North Atlantic Sançleele - 4:

TABLEXVI Lengths of1959and1960year classes of A. marinus in the NorthSea, July/August,1961 (from Macer,1966)

1959class 1960class Ground N Mean length (cm) N Mean length (cm)

Southernmost Region 19 21.0 212 18.5 Indefatigable Bank 86 16.0 204 14,2 Haddock Bank 29 13.7 160 12.5

TABLEXVII Length-weight relationships for A, tobianus, A. marinus A. dubius of the form W aLb

,eci es a b N Season Locality Author

A. marinus 2.5x104 3.068 - S, North Sea Macer(1966) A, dubius 6,3x101 2,778 589 Jan - Mar Western Bank Kohler et al.(1970) 7,8x10. 2.381 356 Jul .- Sept (Nova Scotia) 2.9x10 2.976 207 Oct - Dec 9,9x10 3.153 890 All Seasons

A. tobianus 1,5x107 3,169 80 September Portsmouth Original data (spring (England) spawning) (95%C.L, r ±0.106)

4.2Abundance and Density 4,23Average density

4.21Average Abundance See Section4,21:the density quoted by Bertelsen arid Popp Madsen would be13per Bertelsen and Popp Madsen(1958)cal- square metre. The densities found in bottom culated that in June,1958there were1,800milliongrabs by these authors ranged from2,5 - 3.8 A, marinus on an area of the Southernmost Rough per square metre, approx, 140ion2)in the North Sea; there is no other quantitative information on abundance, 4,24Changes in density

4.22Changes in Abundance See also Section5,41for catch per unit effort data. Although there is a tendency In any single areas sandeel abundance for some sandeels to move inshore in summer and (catch per effort) varies from year to year; no offshore in winter (see Section3.51),most evidence is available to suggest that the major populations are probably present throughout the cause of such variation in abundance is anything year over the same grounds. However, the fish other -than variable year class strength. Within are typically more active in the summer months, a season, fishing mortality may be of and thus are more available for capture during sigeificance in causing a decline in ab,.indance this period. (see Section 5.4). FIRM 582 North Atlantic Sandeels

4.3Na'taii-ty and Recruitment obvious cause of death, although Graham found a significantly greater head length (relative to 4,33 Recruitment standard length) in live fish than in dead fish (p

- For A, marinus and spring-spawning A, tobianus adolescents recruit to the demersal 4,6The Population in the Community populations between May and August about four to and the Ecos stem six months after the spawning periods; in autumn spawning A, tobianus some recruitment takes place As a family, sandeels are a group of in November after an early spawning period, but fish living within and above sandy substrates others appear to over winter as larvae and do not in relatively shallow water,Although classified, appear with the demersal populations until the as demersal fish in fishery statistics, and certainly dependent upon a suitable bottom following April. (xMnd.ler, 1941; Cameron, 1958; substrate for burrowing into, they lead a pelagic Nacer, 1966; present author). existence, generally feeding on plankton and Mortality and Morbidity moving throughout the water masa, when active, 4.4 rpically the sands used by sandeels are clean and course with relatively' little organic matter: 4,41 Mortality rates thus, the permanent infauna is usually scarce, Macer (1966) calculated the total and Nagabhuahanam (1965) referred to the substrate mortality (as mean annual percentage) of of Warts Bank (Isle of Man), and important sandeel A. marinus in the North Sea as 71% (instantaneous ground, as unique in the area because of its coefficient1.24), from the logarithmic plot of poor benthic biomasa.In such areas sandeels age group frequency over three years; with are often abundant and their biomasa dominates additional data, he has modified (pers. comm.) thisthat of other fish species; these fall into one to 65.8%.A similar mean annual mortality for of three categories:1) pelagic species which A. ¡narinus (75%) was obtained by Macer (1966) by are feeding on the plankton, e.g. clupeoids. comparing the abundance (as catch per unit effort) species feeding on what is usually a sparse of individual year classes in successive years invertebrate bottom fauna, e.g. pleuronectoids, over the Dogger Bank, Data -on other species and species feeding on the sandeels themselves, populations is not yet available, but examination e.g. gadoids. of age class composition suggests a similar rate As shown in Sections 3.22 and 3.34, for both fished and unfished populations. sandeels have many predators and are an important 4,42 -Factors causing or affecting source of food -throughout their range; in many mortality places they are the dominant item in the diet of many species of fish and seabirds.The extent Predation on sandeels is considered -to which eandeels determino the abundance and in Section 3.34, and it is likely that this distribution of their predators is, however, mortality factor is of paramount importance; difficult to evaluate, but Ritchie (1932) has unfortunately no quantitative da-ta is available, stated that the abundance and distribution of Occasionally reports of quantities of dead sandeeleboth post-laval arid adolescent sandeela played have been noted in the literature,These include aneminent role in shoal movements of haddock -to the mortalities from near the Isle of May, Scotland,the north and west of Scotland. Richards (1963) in July and August, 1964 (Anon, 1964), and from emphasised the importance of A. americanus as a consumer of sooplankton in Long Island Sound, N.Y., Rhode Island, North America in May, 1954 (Graham, compared with most other fiah species, and although 1956); in both of these instances, there was no as yet not quantified, saaideels, in general, must play a very significant rolo in the trophic network. FIR4/S82 North Atlantic Sancloels :1

5. iIXPL0ITATION engines. Smaller vesselsmayhowever fish in pairs(KUhlandLubmanu,1965) aa-id the 5.1FishingEquipment Norwegians haveused a factory ship inconjunc- tion with their small fleet (Anon. 1957) 5.11 Gears 5.2Fishing Areas The gear used in the North Sea indus-trial fishery has beendescribed by Macer 5.21 General Geographic Distribution andBurd(1970)as awing-type, single-boat, bottom trawl with a high headline anda light Industrial sandeel fisheries are at foot-rope; the latter is lightly weighted with present restricted to the N.E. Atlantic, and. pieces of lead so that it trips along the bottom takeplace mainlr in the Central North Sea without digging in,The bridles are 60-80 metres (ICESregion IVb),andSouthern North Sea (Wc) long, and the mesh size of the net decreases to and the Kattegat and Skaggerak (lila), Smaller 5 mm in the cod-end from 100 rum in the wings. quantitieshave also been landed from the Northern North Sea (Iva), the Norwegian Sea The gear most commonly used in the (lia), the Baltic (111cl) and the Irish Sea (Vila), inshore bait/food fisheries is a beach-seine, and Annual landin from the different areas are nets up to 100 metres long are currently in use shown in Table XVIII. Onlyverysmall landings along the south coast of England; in addition, a have been recorded from the N.W. Atlantic variety of rakes, sickles, forks and even a (Greenland in 1964,arid occasionally in Canadian horse-drawn harrow have been used to extract arid U.S. statistics) but the sandeels in this sandeels buried in intertidal sands.When the area may receive more attention in -the future sändeels are used as bait, a small floating keep (Scott, 1968). (courge) is sometimes employed to keep them alive. Inshore bait/food fisheries are more Acòording to Ber-tolsen and Popp Madsen widespreadthanthe indus-trial fisheries, but are Bigelow and Schroeder (1953) (1958)and Macer (1966) sandeels are inconspicuous of lesser importance. on echogranmes, but Roessingh (1957), Anon (1958),arid Jerome et al, )l965) have referred to such 1)rever and Ellis (1968) and Macer and Burd (1970) fisheries along -the New England coast of North have recorded traces,andechosounders are importantAmerica, and others noted in the literature in the North Sea fishery,The sandeel fisheries include the ?4urinan coast (Andriashev, 1954), are diurnal aa-id no use is made of artificial 'lightNorwegian coasts (Soleim, 1945), the Baltic to attract the fish at night as moue et al. (1967)(Smitt, 1895; Babr,1935)andScotland (McIntosh has recorded for the Japanese fishery. and Mastermari, 1897).At the present time sandeels are still taken in quantity for bait 5.12Boats and food along French coasts; along the south coast of England they are of importance as Macer (1966)has described the typicalbait for sport fishermen. North Sea indus-trial vessels aswoodenseiner/ cutters 20 - 25 metres long with 100 - 250 h.p. TLEXVIII Percentage of Annual Catch1961-67taken in different Regions of ICES StatisticalArea (N.E. Atlantic)

ICES Regions lIa lila 111cl IVa IVb IVc VIla Norwegian Kattegat/ Baltic Northern Central Southern Irish Sea Skaggerak North North North Sea Sea Sea

1961 5,2 93.1 1.7 1962 21.5 7805 1963 12,0 + 88,0 1964 2.5 + + 95.8 1.5 0.2 1965 7.2 3,5 89,1 0,2 1966 0,6 1002 10.1 79,6 0,1 1967 0,2 9.9 0.5 89.5 -

lIB, Danish landingsfrom -the south-western North Sea are probably all included in egion BTb, although the Norfolk Banlçs from where a coniderable part of the catch arises, are strictly in IVc. (Macer, pers. comm.,) FIRN/S82 North Atlantic Sandeels

5.22 Geographic ranges banks aro sharply ridged.; to avoid. net damage, the trawl is towed. parallel to the ridge, on The indus-trial fisheries involve the one side of the crest or the other. exploitation of grounds which are up to 200 km from the nearest coastline and up to 600 km from Popp Madsen (1963) has suggested. that Port0 Most grounds are within loo km of the the most important catches are made in areas coast however, and. in the North Sea involve Danish, where the maximum current speed exceeds 0.5 lmots, German and. Dutch coastal areas south to the Zeeland Islands, the south-wes-ern edge of the 5,3 Fishing Seasons Dogger Bank (Southernmost Rough, South-West Spit and the Outer Well Bank), and, on and. around. the 5.31 General pattern of seasons Norfolk Banks (Meyer.-Waardan 1959-67, Popp Madsen 1963). The former coastal areas, which include All sand.eel fisheries appear to take the Horns Reef, the Terechellingor Bank and. placo in the summer months, and, there are very the Borlcum Rough, were the only areas fished. few landings between October and. April in either industrial or bait fisheries. before the rich sandeel grounds in the south-- The general western part of the North Sea were discovered pattern of seasonal landings for the North Sea in the late 1950's, (Macer, 1966). is given in Table XIX.

5.23 Depth ranges 5.32 Dates of beginning, peak and, end of seasons The industrial fisheries in the North Sea take place on grounds less than 40 metres Table XIX shows that the North Sea in depth (Macer, 1966). fishery begins in April, reaches a peak in June and. may finish any time between August and. 5.24Conditions of the grounds November, 88.6% of the landings between 1961-66 were taken in May, June and. July and all the All the grounds have a clean sand/fine Norwegian and. German landings have apparently gravel bottom, and. in the south-western North Sea been taken in these three months. The 1966 fishing takes place along the edges of the large season has been the longest so far (March to banks and over the tops of the smaller flatter December) but the pattern of monthly landings ones. According to Macer and. Burd (1970) the remained. unchanged. towing course is particularly critical on the Hills and. Outer Dowsing areas where the crests of the

T&BLE XIX Monthly landings of sandeels fÑm the North Sea 1961-66 in metric tons x 103

Data from ICES Bulletin Statistique

Year TOTAL March April June August September October NovemberDecember

1961 83.7 - 12,5 15.2 46,0 9.2 0.6 0,1 0.1 1962 110.0 - 2.6 30.4 40,1 35.0 1.7 0.2 - 1963 151.3 - 9.1 46.7 50,5 38,3 6.7 - - 1964 128.3 - 8.8 34.8 60,7 15.3 7.1 1.5 0.1 1965 125.8 - 10,1 40,0 43.9 27.0 3.0 1.8 - 1966 138.5 ¿ 0.1 6,8 41.7 58.7 19,6 5.6 2.5 3.3 0.3 TOTALS 737.6 < 0,1 49.9 208,8 299.9 144.4 24.7 6.1 3,5 0,3 0.l

< 0,1% 6.8% 28,4% 40.6% 19,6% 3,4% 0.8% 0.5%

at o

200

190

lao

170

160

150 rv O140

130 z(n g 120 o Io w loo z 90

80

70

60

.50

40

30 NETHERLANDS

20 NOR WAY

Io GERMANY. FED. REP.

DENMAR K '53'54 5556 5758'59'60 61 6263.64'65 66 67 '68 YEARS

Fig. 5. Total industrial sandeel landings, North Sea, 1953-68.(from the data in Tablexx) FLRN/S82 North Atlantic Saxideelø 6:1

6. PROTECTION AlU) MANAGE!1E1IT

6.1Reíu1atory (Legislative) Neasures meshed nets provided that not more than 10% by weight of landings are composed of under- As one of the 'Article 6' species of sized protected species (for data on bycatch, the International Fisheriee Convention of 1946 see Section 542). (repeated in N.E. Atlantic Fisheries Commission of 1959), sand.eels may be fished for with small FIRM S82 North Atlantic SaMeels 8:1

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Jensei, A.S., On the subspecies and races of the lesser sandeel (Ammodytes lanoea S. Lat.). 1941 A contribution to the discussion of the species problem in fishes.K.danske Vidensk. Selsk.Skr.(Biol.), 16(9) :1-33

Jerome, W.C. et al., A study of the marine resources of the Merrimack River estuary. 1965 Monogr,Ser,Div.mar.Fish.,Boston, (l):90p

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Kndler, R., Untersuchungen aber Fortapflanzung Wachstum und variabilitat der Arten des Sandaals in 1941 Os-tirnd Nordsee, mi-t, besonterer Berucksichtigung der Saisonrassen von Ainmodytes tobianus. Kieler Meeresforsch., 5(1):45l45 FIRM/582 North Atlantic Sand.eels 8:3

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************ SYNOPSIS OF FISHERIES BIOLOGICAL DATA

This is one of a series of documents issued by FAO, CSIRO and USF\NS concerning species and stocks of aquatic organisms of present or potential economic interest. The primary purpose of the series is to make existing information readily available to fishery scientists according to a standard pattern, and by so doing also to draw attention to gaps in knowledge.ltis hoped that synopses in the series will be useful to other scientistsinitiatinginvestigations 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. FB/S

CSIRO Fisheries Synopsis No. D FO/S and

USFWS FAOFisheries Synopsis No. BC FIS

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 scien- tists in drafting synopses on species about which they have knowledge, and welcome offers of help in this task. Additions and corrections to synopses already issued will also be most welcome. Comments including sugges- tions 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: U.S. Department of the Interior Scientific Editor Fish and Wildlife Service CSIRO Division of Fisheries and Oceanography Bureau of Commercial Fisheries Box 21 Office of Scientific Publications Granulia, N.S.W. Bldg. 67, U.S. Naval Air Station, 2230 Australia Seattle, Washington 98115, U.S.A.

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

The following synopses in this series have been issued since January 1969: BCF/S40 Synopsis of biological data on the Pacific mackerel Scomber jopan/cus Houttuyn (Northeast Pacific) February 1969 FRi/530 Synopsis of biological data on the pike Esox lucius (Linnaeus) 1758 May 1969 Rev. 1 DFO/S3 Synopsis of biological data on the tiger prawn Penaeus esculentus Haswell, 1879 July 1969 FRmIS43 Synopsis of biological data on the anchoveta Cetengraulis ,nysticetus Günther, 1866 October 1969 FRh/S35 Synopsis ofbiological data on West African croakers Pseudotofithus typos, Rev. 1 P. senegalensis and P. elongatus November 1969 BCF/S42 Synopsis of biological data on the Atlantic menhaden, Brevoortia tyrannus November 1969 FRm/S78 Synopsis of biological data on smelt Osmerus eperlanus (Linnaeus) 1758 December 1969 FlRl/S80 Synopsis of biological data on the eel Anguilla anguilla (Linnaeus) 1758 (Provi- sional version) April 1970 DFO/S4 Synopsis of biological data on the rainbow prawn,Parapenaeopsis scuiptilis (Heller, 1862) 1970 BCF/S4 Synopsis of biological data on chum salmon Oncorhynchus keta July 1970 DFO/S5 Synopsis ofbiologicaldataontheschoolprawn Metapenaeus macleayi (Haswell, 1879) 1970 FIRI/S80 Synopsis of biological data on the eel Anguilla anguilla (Linnaeus) 1758 October 1970 FIRM/S82 Synopsis of biological data on North Atlantic sand eels of the genus Ammodytes (A. tobianos, A. dub/us, A. americanus and A. merinus) October 1970 MI/A8703/1 i .70/E/I /750