Science Series Technical Report no. 147

Spawning and nursery grounds of selected fish species in UK waters

J. R. Ellis, S.P. Milligan, L. Readdy, N. Taylor and M.J. Brown

Science Series Technical Report no. 147

Spawning and nursery grounds of selected fish species in UK waters

J. R. Ellis, S.P. Milligan, L. Readdy, N. Taylor and M.J. Brown This report should be cited as: Ellis, J.R., Milligan, S.P., Readdy, L., Taylor, N. and Brown, M.J. 2012. Spawning and nursery grounds of selected fish species in UK waters. Sci. Ser. Tech. Rep., Cefas Lowestoft, 147: 56 pp.

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HMSO’s Licensing Division St Clements House 2-16 Colegate Norwich NR3 1BQ Fax: 01603 723000 E-mail: [email protected] 3 1 Contents 55 55 47 49 54 55 the the

supporting supporting

grounds. for for

grounds.

data data

the the

spawning nursery

with with

nominal nominal

of of

associated associated

identification identification Other considerations References (b) Metadata 4. (a) Metadata 5. Appendix A: List of acronyms Appendix B: ESRI shape metadata file 7 7 7 6 6 6 8 8 8 9 5 5 7 31 33 35 37 39 41 44 46 22 24 26 28 29 20 18 19 15 16 17 13 14 10 13

complex

larvae

habitats

and

fish fish habitats

layers

eggs

data Trachurus trachurus Trachurus

Dipturus batis- Dipturus Merluccius merluccius Merluccius

Raja clavata Raja juvenile fish

of

Raja undulata Raja Micromesistius poutassou Micromesistius

surveys

estuarine important of of

Raja montagui Raja fish species

Galeorhinus galeus Galeorhinus

ray standards

Lophius piscatorius Lophius

seasons grounds

of hake surveys Scomber scombrus Scomber Ammodytidae

skate ray

Molva dypterygia Molva

Squalus acanthias Squalus grounds

Clupea harengus Clupea ray Merlangius merlangus Merlangius

and

assurance

of

Pleuronectes platessa Pleuronectes mackerel

selectivity shark whiting ling Molva molva Molva Solea solea Solea

Gadus morhua Gadus

Spawning and nursery of grounds selected Introduction Data collation 3.15 3.16 Anglerfish 3.17 Horse 3.18 Sandeels 3.19 Mackerel 3.20 Plaice 3.21 Sole Spawning 3.10 3.11 Whiting 3.12 Blue 3.13 Blue 3.14 Ling European 3.9 Cod 3.7 3.8 Undulate Herring 3.4 3.5 3.6 Common Thornback Spotted 3.2 3.3 Spurdog Tope 2.2 2.3 2.4 Distribution Quality Taxonomic 1.3 1.4 Spawning Nursery 2.5 2.6 2.7 Gear 2.8 Timing Locations Coastal Contents 3. 3.1 Interpretation 2.1 Distribution 1.1 1.2 Background Ecologically 2. 1. 4 5 1 Introduction Merluccius Galeorhinus and horse Raja montagui, undulate Molva , molva hake Lophius piscatorius Dipturus batis-complex, thornback et al. (1998), it was considered that Squalus acanthias, tope , anglerfish Trachurus trachurus. Trachurus , and comprises two main sections. Section 2 1 Raja clavata, spotted ray Raja , undulata ling This report aims to update some of these maps In addition to eight of the highly mobile species The summary report is based on a report provided Ellis et al. (2010). MB5301 Mapping spawning and nursery 1 areas of species to be considered in (Marine Marine Protected Conservation Areas Zones). Report development No 1: of Final derived Report data considered to be of on conservation importance. layers for 40 mobile species follows that these maps should unchanging descriptions not of presence be or seen ”. absence Since as these rigid, maps were produced ichthyoplankton surveys in some areas, and there are now there have been other fish further species for which there location of is ‘critical habitats’. an interest in the with more recent data, and so highlighting some evidence for of their selection, and the to provide a first attempt at identifying some of the ecologically important habitats for some other fish species, including some conservation species importance of (e.g. OSPAR, 2008) been that considered in have marine planning. note It that is such data important are to not available for all fish and many coastal, continental shelf, and shelf edge waters species, are still to be surveyed for ichthyoplankton and juveniles. included within Coull existing groundfish surveys could assist in the preliminary spatial description of the nursery grounds for some other species of commercial and/or conservation including importance, spurdog galeus, common skate ray ray merluccius provides an overview of the data collation for the species, including a summary trawl of surveys and available ichthyoplankton surveys. fishery-independent This section also includes a discussion on the limitations and caveats of these data (e.g. issues on gear selectivity, the timings and locations of surveys, and taxonomic identification in surveys) and the users of the GIS to data are use encouraged this information of to distribution assist data. in Section the information 3 on interpretation provides the more derived detailed data species. layers for the various to Defra mackerel Gadus Sprattus , whiting Solea solea Pleuronectes Micromesistius Pollachius virens, Norway Clupea harengus. Microstomus kitt, sole Scomber scombrus, sprat Nephrops norvegicus. Melanogrammus aeglefinus (1998) correctly acknowledged that et al. , haddock Introduction Background et al., 1998). This report aimed to assist in the Trisopterus Trisopterus esmarki, blue whiting Such management actions have implications for Lee & Ramster (1981) compiled an atlas of the seas Coull 1. An evidence-based understanding of the fish spawning distribution and nursery grounds, and other ecologically of important fish habitats isadvisors and regulators to better manage human activities required to allow in scientific our seas. restrictions seasonal For to subject currently are developments example, many offshore in order to wind minimise disruption from anthropogenic noise, farm particularly the sounds associated with spawning pile driving, fish on and to Similarly, there protect is egg more conservative and regulation of management some human larval activities that and operate in stages. areas of seabed considered important or potentially to important key life-history stages, for example in spawning grounds of herring relation to the morhua poutassou, mackerel 1.1 developers in terms of both cost and delivery of projects. Consequently, it is essentialdata on that spawning the and to provenance nursery ensure grounds of is that understood its Impact Assessments interpretation and within regulatory Environmental decision-making appropriately applied. is This report describes the sources, spatial and temporal coverage and limitations of the data, including where there are data gaps. Using the isolation maps in may result in misrepresentations so of in all the cases data, the supporting considered. rationale should also be around the British Isles, in which the spawning of grounds several commercial fish were illustrated. Subsequently, a collaborative project between laboratories the (Cefas national and fisheries Services, the Scotland), then Association with Fishermen’s Scottish Fisheriesthe (UKOOA), the Association Research UK Offshore Operator’s (SFF) and Organisations the (NFFO) aimed to provide broad scale Nationalmaps of Federation the sensitive of habitats of (Coull Fishermen’s marine fish in UK environmental impact waters assessment process, and included maps indicating the main spawning and nursery grounds for 14 commercially important species: herring, cod Merlangius merlangus, saithe sprattus, sandeels (Ammodytidae), plaice platessa, lemon sole and Norway lobster “spawning distributions are under continual revision. It pout 1 Introduction 6 data to accurately no identify sites of importance are in UK waters. there and surveys groundfish in occasionally very captured only are sharks pelagic Larger shelf. continental the on surveys groundfish from known species the are of some juveniles for of locations the although these of species, some for rare are pups at near-term or (e.g. term with birth give females then mature of locations the on Data will sites). parturition females gravid and uteri, sampled. are oviducts) the in egg-cases of these surveys means that very few ‘active’ fish (i.e. with location and timing the surveys, groundfish in undertaken is skates of sampling biological Although skate. of classes surveys particularly effective for sampling the smallest size trawl beam with surveys, trawl in taken are fish, hatched recently are including juveniles, egg-cases) However, limited. viable or lacking of (presence grounds spawning delineate to data field strandline, the on egg-cases these of occurrences the for data some are there although and floor sea the on purse’) ‘mermaids (the egg-cases deposit elasmobranchs) species). live-bearing (for sites parturition and egg-laying (for sites oviposition to it include use so and sense, generic its in ground’ ‘spawning in state. spawning the year assess to order of time right the at surveys have areas all not as project, current the in used not were data these although running with grounds, spawning of indicative be also can sperm or eggs fish Mature site. spawning the from away advected been have may stages larval later that noted be should it although grounds, spawning of indicative be can structures. biogenic on or floor sea the on extensive more eggs deposit which species those have than grounds spawning may strategies species frequent Such more 1984). the (Balon, of the one into is sperm column, and water eggs the shedding involves which spawning, broadcast and fishes, in reproduction of modes numerous are There extent. aggregate spatial restricted more a may over species other year, to location year change from slightly may that grounds have spawning may so extensive and behaviourally aggregate may species 1.3 2003). (ICES, pathways migratory and life-history grounds feeding early as well as of grounds), nursery (i.e. stages growth and recruitment (e.g. grounds), breeding to parturition and grounds, importance and beds spawning sites, of mating sites include can marine fish highly-mobile for habitats Ecologically-important 1.2 iiaos hrs ar te eeoig on i the in young developing the carry sharks Viviparous species these and oviparous are (Rajidae) Skates term the use we report present the of purposes the For The presence of eggs and larvae of broadcast spawners some Whereas spawn. to aggregate species fish Many Spawning groundsSpawning Ecologicallyhabitats fish important of habitat that may be and utilised by a wider range of juveniles. fish, old) year extent broader the to equating grounds nursery secondary (<1 0-group and recently-settled to term the use we primary nursery here grounds to equate with sites of importance and fishes, teleost to extended Bass (e.g. ground’ nursery ‘secondary a termed is what occupy may juveniles larger so and and/or habitat, and range extend home their shift may they grow fish hatched recently the As grounds’. nursery ‘primary termed be can life periods their initial of the spend and species) the oviparous (in of hatch case or species) viviparous of case the (in born are pups where locations those that fishes for elasmobranch suggested been has it example, a For as there ground. considered nursery be grounds), may what spawning in changes subtle of be can extent mode and reproductive location the and (including fishes the of history tows). the of 70% or 50 in (e.g. regularly caught been have juveniles where the sites and surveys, station fixed for and, juveniles of rates catch maximum the distribution), length the of analysis an layers data generated include the presence of ‘juveniles’ (derived from available, were data scale broad appropriate Where site. the at observed been have years juveniles which in of proportion the also and juveniles of rates catch identification of nursery grounds may be inferred from high robust more for a then source grounds, nursery data of identification main the field the When are surveys years. annual the from data over repeatedly used was area nursery the (3) and fidelity, site greater be would there areas, (2) other in than greater (1) was criteria, juveniles of three density the on based identified be could grounds lacking. generally are recruitment to contributions relative greater that provide and rates enhanced growth confirm have successfully, to more predation data avoid they comparable densities, higher at occur juveniles where areas highlight to available are data Beck (see habitats ground non-nursery to comparison in recruitment adult to contribution relative greater a providing grounds nursery in result should which habitats, other in growth than rates faster densities, have and higher predation of at rates reduced occur have juveniles where sites those are grounds nursery that suggested been has It grounds. The grounds where juveniles are found are termed nursery 1.4 t hud lo e oe ta, eedn o te life- the on depending that, noted be also should It Heupel Nursery groundsNursery 20) et n o ugs ta nursery that suggest to on went (2007) al. et 20; Heupel 2003; al., et 17) Ti cnet a b usefully be may concept This 1973). al., et 20) Wis field Whilst 2007). al., et 7 2 Data collation m et al., 2004; et al., 2002; Alvarez et al., 1998). However, as these data 2007), multi-species data are not available not are data multi-species 2007), al. et Quality assurance Distribution Distribution of fish juvenile survey (2007) and for stage 1 eggs were included. For further information see ICES (2009b). For the purposes of the present study, an internationally an study, present the of purposes the For For other species, the length-frequencies of Cefas-held It should be noted that some data from Fish the Young Ibaibarriaga for recent surveys. Similarly, the waters of north-western surveys ichthyoplankton few to subject been have Scotland (except for herring and mackerel). 2.3 There are several issues with regards to data quality that identification taxonomic particularly considered, be to need and the design of stations, timing of the the survey etc.), when survey interpreting the (gear type, data layers. location of beam trawl (Rogers are more regional, only cover a part of the UK and coastline, are currently subject to on-going analyses, they have not been included in the present report. 2.2 Data from national summary the groundfishgrounds, nursery of identification the support surveys were used details of to these surveys are provided in Table 1, and the distribution of survey stations is illustrated in Figure 2. agreed length for identifying following ICES, 2009c). It should be noted that survey data juveniles was used single a as and year the of (e.g. times different at collected were length range was used (Table 2), as opposed to different length ranges by season, the data used may encompass fish of just over one year of age for surveys undertaken in the first part of the year. groundfish survey data were length range corresponding to initially the first cohort(s) identified. examined, and a In those instances where data were limited for cohort, the first then a broader included, length as range detailed in of the juveniles species-specific below. information was 2 by collected data these with available, are (YFS) Survey Hence, most of the available ichthyoplankton data from were the Irish Sea, English Channel and Sea, North the Bristol in survey coordinated internationally Channel, an from and from international species-specific surveys for herring and mackerel. Although there have been studies on wider the ichthyoplankton of the Celtic Sea and Fives, 1994; Acevedo (e.g. Horstman (2007). al. et et al. (1997) and Bunn (2005) and Taylor Taylor and (2005) al. et et al. (1993). et al., (2004). Data Data collation Distribution Distribution of fish eggs and larvae Bristol Channel plankton surveys: cruises A (524 valid series stations) of conducted in five Channel in the 1990 Bristol which were conducted provide primarily data to on sole and bass, but collected data for other fish species and edible crab larvae. For information see further Horwood (1993). Sole surveys: A series of four valid cruises stations) in conducted 1991 in the (304 English Channel with bass, and sole sample to primarily Bight, Southern and data collected for other fish species and larvae. edible crab Irish Sea plankton surveys: This extensive cruises was series conducted in the of Irish Sea (although data were limited from the southern parts of St stations), 2000 Channel) in 1995 (13 cruises, 1024 valid George’s (8 cruises, 804 valid stations), 2006 valid stations) and (5 2008 (5 cruises, cruises, 486 valid stations). 397 For further information see Fox and Fox (2004), with earlier cruises also described by Nichols Eastern Irish Sea plaice surveys: A series that were undertaken in the eastern Irish of Sea, primarily cruises cruises, (5 2001 in conducted were that plaice, target to 227 stations), 2002 (4 cruises, 158 stations) and 2003 (5 cruises, 345 stations). For further information Bunn see North Sea cruises: A international series institutes of covering cruises a by large North Sea (927 stations) over a four month period from area several of the late December 2003 to early April Fox see 2004. information For further International Herring internationally coordinated Larval survey that samples Survey over the main herring spawning grounds in the (IHLS):North Sea. Only data for An 2008 (481 stations) were used, as 2009 ICES see information further For unavailable. were data (2009a). Triennial mackerel egg coordinated survey survey: that includes An extensive sampling along internationally the western Mackerel seaboard and of horse recent most the the from mackerel data only survey, this For species. are British the Isles. main target 2. Data on the distribution of the planktonic stages of fish eggs fish of stages planktonic the of distribution the on Data and larvae (ichthyoplankton) were collated from numerous surveys (Figure 1) conducted by Cefas and associated UK fisheries laboratories, and from internationally-coordinated ichthyoplankton surveys, as discussed below. • • • 2.1 • • • • 2 Data collation 8 taken in VIa, and VIa, in taken ov dypterygia (Molva sub-species dypterygia, be to have them authors other considered whilst separate, species two these and consider dypterygia) authors Some (Molva macrophthalma). (M. ling ling Spanish blue of terms in especially Sea. Celtic the in and were aggregated at the family level. family the at aggregated were the correct species identification. regards For broad scale mapping with data quality data uncertain of as flagged should be surveys trawl scientific from collected data even species. two the separate not do data commercial range, species entire the cover not do Although surveys report. vessel research this in considered not is species latter anglerfish black-bellied and anglerfish namely species, two for available south. further and Sea Celtic the in found is and southerly more is ling Spanish whereas Scotland, NW off common is and distribution, northerly more a has cf. skate flapper with Sea, Celtic the in collected samples west Scotland (VIa) were generally genetically distinct from Griffiths unclear, are species two the of distributions geographical (Iglésias Nomenclature be validated by the International Commission on Zoological and species: be two should these for names resurrected scientific former that problem proposed the have on working Taxonomists synonym. invalid name scientific the so and updated, being currently is complex’ skate ‘common the (Griffiths differences complex. A subsequent study has confirmed these genetic species, and so the data layers provided are for the species two comprising complex a actually is skate common that long-snouted various skate. common for data the compromises between which species, confusion also is There ray may be easily confused, particularly in commercial blonde data. and ray spotted and skate), (thorny ray some starry and example, For (Rajidae). nations fishing in the North Sea may confuse thornback ray skates of identification below. detailed are these and report, this in considered species the of number selected a for available data the of some of quality the compromise can issues to such and interest, of fundamental species for information is evidence-based robust providing identification taxonomic Accurate 2.4 The taxonomy of some species of hr ae ie pce o snel n K es and seas, UK in sandeel of species five are There are surveys trawl scientific from anglerfish on Data Iglésias by study recent A and taxonomy the with problems several are There n ocsoa bu skate blue occasional and intermedia D. Taxonomicstandards cf. 21) bevd ht ape fo North- from samples that observed (2010) al. et , but this proposed change needs to needs change proposed this but intermedia, and . le ling Blue macrophthalma). dypterygia M. D. cf. 21) Te oecaue of nomenclature The 2010). al., et taken on the Rockall Bank Rockall the on taken flossada 21) Atog te overall the Although 2010). al., et will soon be an be soon will batis Dipturus atog the although budegassa, L. 21) a revealed has (2010) al. et Dipturus Molva D. is problematic, L. piscatorius L. cf. cf. flossada flossada D. rws r mr sial ta ba tal fr sampling skates. for larger trawls beam than otter suitable although more are surveys, trawls trawl beam in as effectively as sampled not are skates faster-swimmingJuvenile species. under-represent may surveys trawl but numbers, large in captured be can spurdog) and (tope sharks bentho-pelagic the of Some anglerfish). ling, (e.g. or presence/absence plaice) for larger and cod hake, whiting, (e.g. abundance relative either for used be may and gear also this by are sampled species The demersal column. larger-bodied water common be the more can in position and their variable on highly dependent are of species rates shoaling catch that such recognised be must should it eels), sand (e.g. and whiting blue species mackerel, herring, mackerel, horse bentho-pelagic and pelagic of distribution panels. bottom the on meshes the through escape can these as species, demersal small-bodied for less be will efficiency gear The species. demersal as well as fish pelagic sample to as so heights, headline high have gears These (PHHT). trawl headline high Portuguese the and trawl (GOV) vertical ouverture Grande the are available only.presence or presence/absence either For other species/size classes, these data are ling. presented as and anglerfish skates, whiting, cod, of juveniles the and sole plaice, of abundance relative the on informing for appropriate considered study,are of present trawls the beam purposes the For types. ground of variety a be on can fished gears these and fishes, demersal sized medium catches. under-representedin be may skates, larger as such fish, large-bodied that such are mat chain the of dimensions the but net, the entering mat chain the boulders large prevents and contact ground of close ensures presence The mackerel). and herring (e.g. fishes pelagic catching of chance low extremely an 45-50 only beam gears. the of suitability the of context the in viewed be must data survey trawl of interpretation the and species, various the for efficiencies and patterns selection gear different have gears fishing All 2.5 water during the winter). the during water deeper into moving (e.g. distribution restricted in changes more seasonal have may species Other abundance. may or species fish some distribution either in patterns seasonal pronounced that exhibit recognised is most for It available species. not are changes scales geographical seasonal wide showing on data spatial series, contemporary time the of parts early the in autumn) and spring Celtic (e.g. North, surveys biannual from benefited have Seas) the Irish and for (including surveys some Although distributions. fish in seasonality encompass basis, not do so annual and an on conducted are surveys recent Most 2.6 lhuh hs sres r ue t ifr o the on inform to used are surveys these Although are data most which for trawl otter of types main The to small sampling for suited best are trawls Beam main the with height, net low a have trawls Beam Gear selectivityGear Timing of surveys of Timing m bv te e for hne hr is there hence floor, sea the above cm 9 2 Data collation The beam trawl surveys in the eastern English Channel, Most groundfish surveys are undertaken on research m deep. Hence, these surveys have less ability to internationally-coordinated survey. Although otter surveys on trawl the west coast also attempt rectangle, to some fish rectangles in are each fished at more than location, as there can one be large differences in depth. Bristol Channel and Irish Sea were originally designed to sample juvenile plaice and sole, and so there is a greater density of stations in coastal waters and a lower density of stations in offshore regions. vessels, and these vessels rarely fish in waters less than 15 for surveys ichthyoplankton collated. were selected of data larvae and Distribution egg Locations Locations of surveys The English North Sea IBTS survey has one station which Figure 1: Figure 2.7 Most of the surveys examined sample at fixed although stations, some surveys use a design, and stratified these stations are selected to random ensure that the survey area is effectively sampled for the species of interest, with stations selected by ICES rectangle, depth or strata. These tows are also selected to minimise the chances of damage. gear per selected rectangle, as these twice when considering all the participating rectangles nations in this are fished 10 2 Data collation the fringes of their distribution. their of fringes the so and groundfish Atlantic, surveys on the UK NE continental shelf only sample the of waters deeper the in ranging proportion of their range. For example, blue whiting is a wide cover only may surveys existing species, wide-ranging more for although distributions, species/stock the of most in or all cover species/stocks surveys existing instances, the some In question. of distributions the with surveys Isles. British the of west the to occurring fish deep-water data of distributions these the on and inform better France), will and Ireland with (e.g. internationally-coordinated surveys for plans are there as future, the in species deep-water on data comprehensive spatially conducted by Marine Scotland, there will hopefully be more are surveys deep-water Although species. deep-water on shelf waters, and so most have very limited ability to inform future. the in on reported be will and analysed being used in more site-specific studies. These data are currently be only could so and coastline, UK entire the cover not do data these areas, is some in Survey vessels inshore Fish by Youngconducted the Although species. several of stages history life earliest the can for important particularly which be habitats, coastal and estuarine the on inform Figure2: fish t s lo motn t itrrt aa rm groundfish from data interpret to important also is It continental in undertaken are surveys groundfish Most data were Distribution collated. of UK groundfish surveys for which juvenile e) d) c) b) a) Swaby,1993a): and Potts from (adapted groupings following the to attributed Swaby,1993a,b). and (Potts text supporting the in given is species study case be the by may used fringes coastal the which to extent The waters. of sub-rectangles coastal on into extend may so and rectangles, statistical ICES based are here included nursery grounds and spawning for layers generated The lacking. often are waters transitional and estuarine for data scale >20 waters in operate generally collected surveys These been surveys. vessel have research during layers data associated this of and production the report in used data survey the of Much 2.8 aie ih curn i etaie aias a be may habitats estuarine in occurring fish Marine estuaries. Fully marine nursery species which are recorded occasionally in as habitats grounds; estuarine utilising fish Marine feed; to habitats estuarine visiting fish Marine estuaries; through migrating fish Diadromous habitats; estuarine Fully estuarine fish living almost their entire life-cycle in Coastal and estuarine habitats estuarine and Coastal de, n cmaal broad comparable and deep, m 11 2 Data collation that temporal noted etc.) be their net ICES (2006, 2009c) Warnes and Jones (1995); and Warnes Tidd (2006) Harley and Ellis (2007); ICES (2009c) Parker- Humphreys (2005); ICES (2009d,e) Parker- Humphreys (2004a,b); ICES (2009d,e) ICES (2009d,e) - ICES (2006, 2009c) ICES (2009c) References ICES (2009c) ICES (2009c) should trawl It summarising 1275 1566 422 2039 3284 1239 228 3939 77 Total number Total of stations fished 213 2946 2260 polyethylene information. analyses, or nylon detailed present the more of floatation, within m beam trawl extra sources m beam trawl m beam trawl m beam trawl deployed or Grande ouverture vertical (GOV) trawl Portuguese high headline trawl (PHHT) GOV with either rockhopper or standard ground gear 4 4 4 off a commercial fishing vessel 4 Twin GOV trawl GOV trawl (rock hopper) Gear GOV trawl (rock hopper) Rockhopper trawl included and kite (BTS) fished gear, surveys ground IVa-c VIIe-j VIIa, e-h VIId, IVc VIIa,f,g VIIe VIIe IVa,b VIa, b Survey area (ICES Divisions) VIIa VIa, VIIb,c,j (stations) of trawl (type samples beam Quarter 3 1 4 3 3 4 1 1, 2, 3 4 1, 3 1, 4 of trawls and GOV (GFS) number total Time period (years) 1992–2008 1982–2004 2003–2008 1988–2008 1988–2008 1989–2008 2006–2008 1983–2008 2006–2008 1998–2008 1992–2008 1986–2008 between surveys type, gear differences groundfish of extent, some List are spatial there and Survey English North Sea GFS Table 1: Table English West Coast GFS English Celtic and Irish Sea GFS English channel and southern North Sea BTS English Near West Coast BTS Western English Channel BTS South Western BTS Scottish North Sea GFS Scottish surveys of Rockall Bank Rockall Trough Northern Irish GFS Scottish West Coast GFS 12 2 Data collation Sole Plaice Mackerel Sandeels mackerel Horse Anglerfish Hake Ling whiting Blue Whiting Cod Herring ray Undulate Spotted ray Spotted ray Thornback skate Common Tope Spurdog Species Table2: Summary of length range cohort(s) examined (cm) examinedcohort(s) juvenileof range Length used to 1 – 8 / 1 –14 1 / 8 – 1 map 1.5 – 11 – 1.5 23 – 2.5 12 – 2.5 3.5 – 17 – 3.5 19 – 42 – 19 69 – 24 47 – 24 1 – 12 – 1 27 – 3 19 – 3 48 – 7 18 – 2 19 – 2 22 – 2 17 – 1 28 – 9 5 – 17 – 5 juvenile habitats and surveys, all juveniles)all surveys, juveniles(all werecaught where(m) range Depth the observed 10 – 500 – 10 592 – 10 10 – 500 – 10 10 – 500 – 10 171 – 11 500 – 10 6 – 350 – 6 500 – 7 500 – 8 460 – 7 450 – 7 460 – 8 140 – 9 8 – 500 – 8 6 – 405 – 6 405 – 7 7 – 220 – 7 6 – 405 – 6 depth distributions of 50% of tows)regularlyof (>50% juvenileswerewhere caught sites of (m) ranges Depth fish within this 11 – 119 – 11 500 – 10 450 – 15 450 – 10 10 – 500 – 10 592 – 10 15 – 270 – 15 500 – 21 10 – 500 – 10 13 – 189 – 13 18 – 180 – 18 15 – 460 – 15 117 – 90 405 – 10 7 – 450 – 7 405 – 9 39 – 30 6 – 385 – 6 size range 13 3 Spawning and nursery grounds of selected fish species et al. (1998). et al. (1998). et al. (1998). et al. (1998). Presence of juveniles in field surveys (within the length the (within surveys field in juveniles of Presence in found were they whether and 2) Table in given range at least one haul, 50% of hauls at the station (for fixed station surveys) or 70% of the hauls at the station. The maximum catch per unit effort (CPUE) by survey station location, also indicating how regularly juveniles were captured at that haul station. Updated nursery ground layer statistical based rectangles, on with sites half of ICES importance higher noted for selected species. perceived This layer was based on the evidence provided by the trawl data and the layers provided survey by Coull Haul locations (i.e. surveys). sites fished during Original groundfishlocation(s) of nursery grounds as provided by Coull Occurrence and relative abundance of species eggs of of interest. the Where egg stage surveys for only selected species, identified then aggregated the data across were all egg stages. The this is exception for horse to mackerel and mackerel where broad scale data on the distribution and abundance of early stage eggs were available, with early stages of eggs more likely to correspond to a close proximity to the spawning ground. Occurrence and relative abundance of larvae species of interest. of the Updated spawning ground layer based on statistical half rectangles, ICES with sites of higher importance the on based was layer This species. selected for noted evidence provided by the ichthyoplankton survey data and the layers provided by Coull Haul locations (i.e. sites sampled during ichthyoplankton during sampled sites (i.e. locations Haul surveys). It should be noted that these are not evenly distributed in UK waters, so while some been subject to intensive areas sampling over time (e.g. Irish have (e.g. occasionally surveyed been have areas other Sea), have or areas other and Channel) English and Sea North only been surveyed for mackerel and horse mackerel eggs (e.g. Celtic Sea). Original location(s) of nominal spawning grounds provided by Coull as The following GIS data layers for spawning grounds (c) (d) (e) The following GIS data layers for nursery produced: grounds were (a) (b) (c) (d) (e) Rectangle (i.e. rectangles of 1º longitude and 0.5º latitude) and bathymetry. were produced: (a) (b) nm (if trawled at 4 knots for 30 Spawning Spawning and nursery of grounds fish selected species Interpretation of Interpretation data layers It must be recognised that all bubble plots with an index an with plots bubble all that recognised be must It Sampling points (shooting positions for trawl and Single layers of bubble plots of relative abundance/ It should be recognised that some surveys have a low The distribution of ichthyoplankton includes a range of In terms of the spawning ground information, the The data layers below were used in conjunction with 3. The following issues interpretation of the data layers provided. are highlighted to avoid of mis-relative abundance/catch rates are on a species-specific scale, and so cannot be compared across taxa. ichthyoplankton surveys) are provided as although single it points, should be noted that trawl surveys can stations cover 2 fished during minutes). catch rates have been provided for each species, than rather by survey, to limit the number of different layers. surveys However, will have different catchabilities for the various species (due to a combination of gear type, survey protocol, timing of survey etc.). Hence, these data should be viewed in conjunction surveys themselves. with Spatial differences in the catch rates the distribution of over the survey the grid of one particular survey can be used to infer spatial differences in the the relative species. On a abundance broader scale of (i.e. across surveys), any apparent regional differences in catch rates should not be inferred to represent true spatial differences in the relative abundance of the stock. density of sampling stations. Other surveys have a density high of stations and/or high survey effort (e.g. Irish Sea for in both ichthyoplankton the and trawl surveys), and these data may allow finer scale studies. egg and larval stages, and later stages will have advected the only Although ground. spawning original the from away the for plotted were stages egg earliest the of distributions triennial mackerel egg survey, aggregated egg data were used for other taxa and surveys, in order to maximise the data available. egg numbers are on a surveys, single and scale the across user the should various be recognise differences that in there the can catch rates to between other factors surveys (e.g. timing of due survey) as well as regional differences. Larval data are also on a single scale across all surveys, but on a different scale to the data layer for eggs. Some surveys (e.g. the triennial mackerel egg survey) do apparent the so and larvae, for data comparable provide not absence of larvae in these areas is an artefact of sample processing. data layers for UK territorial waters, ICES Division, ICES 3.1 14 3 Spawning and nursery grounds of selected fish species oain ad eprl tblt o seii parturition specific of established. well not are grounds stability temporal and Locations groundsSpawning grounds. nursery secondary and primary <48 of <30 specimens for data as and, grounds. nursery and used for the preliminary identification of be parturition grounds could data survey trawl and spurdog, juvenile sample surveys trawl Scientific delineate. accurately to difficult be may habitats such months, that 22–24 for pups and the carry they 1991) (Vince, migrations seasonal widespread and complex have they that given but surveys, in scientific taken females gravid of presence the on available data are Limited juveniles. (especially as well mature as occur, females where gravid) sites include would habitats the UK EEZ. As a viviparous species, ecologically important the within widespread throughout is stock this and shelf Atlantic North-eastern continental the on occur Spurdog informationGeneral 3.2 Figure3: nominal presence Spurdog are born at a length of 19–30 of length a at born are Spurdog cm was used to better inform on the distribution of distribution the on inform better to used was cm Spurdog nursery Nursery of juveniles grounds grounds Squalus acanthias Squalus (<48 of cm of spurdog, spurdog. total length) cm were limited, a length a limited, were cm as Data indicated in layer groundfish cm (Gauld, 1979) (Gauld, cm indicates by the surveys. grounds. nursery potential the of some of utility the confirm to be able may studies field Dedicated of spurdog. female movements mature the understand better to undertaken be the in undertaken be Taggingusefully future. could tags electronic using studies will this coordinated and surveys, internationally groundfish during could collected stages) be maturity (e.g. usefully species this on data More gaps Data recorded in is occur typically that not <10 waters species does It marine estuaries. in fully occasionally a is Spurdog coastalzone and Estuarine North-west Sea. Irish northern the in and off Scotland made The were area. juveniles of this catches in highest surveys trawl otter fewer were there as sampling, the of artefact an part in be may Sea North of juveniles from the eastern English Channel and southern absence apparent The 3). (Figure Sea Celtic and northern Sea Irish Scotland, North-west Sea, North Northern Central and the in distributed widely were spurdog Juvenile groundsNursery m deep (Wheeler,deep 1978). m 15 3 Spawning and nursery grounds of selected fish species Juvenile tope have been caught sporadically in the North Sea (e.g. Greater Thames Estuary and Firth of Forth), Irish Sea, Bristol Channel and areas off North-western Scotland (Figure 4), suggesting that juvenile tope occur in inshore areas with slightly reduced salinity. The apparent absence southern and Channel English eastern the from juveniles of North Sea may be in part an artefact of the sampling, as there have been fewer otter trawl surveys Sites such as in The Solent may also act this as nursery grounds area. for tope, as they are caught in recreational fisheries in this area. Estuarine and zone coastal may utilise the outer reaches is a marine species that Tope of some estuaries and coastal ground and nursery ground. waters as a parturition Data gaps More data on this species during internationally coordinated studies surveys. Tagging could usefully be collected using electronic tags could better understand usefully the movements of be tope in undertaken UK waters. to Dedicated field studies sampling tope could confirm the utility of using potential nursery a gear grounds. appropriate for Nursery Nursery grounds tope. nursery cm for Channel, (<70 nominal habitat English juveniles for indicates of important an limited layer cm (Compagno, 1984) (Compagno, cm as presence cm were limited, a length Data Data the tope. by regarded of surveys. Galeorhinus galeus often is indicated grounds as groundfish Solent in tope, Nursery the of Tope shark Tope cm was used to better inform on the distribution of length) Tope are born at a length of 30–40 of length a at born are Tope although total grounds Figure 4: Figure Spawning grounds Locations and temporal stability grounds are not well established. of specific parturition 3.3 General information The North-eastern Atlantic stock of tope is wide-ranging from boreal to sub-tropical waters. Seasonal patterns the in movements and distribution around the British are Isles poorly known, although there some is north-south migrations the (Holden and suggestion Horrod, 1979). of There are few data on of numbers small although the Isles, British the around species reproductive biology of this these Hence, surveys. groundfish in taken be can juveniles identification preliminary the for basis the provide may data of nursery grounds. and, as data for specimens <40 of <70 primary and secondary nursery grounds. 16 3 Spawning and nursery grounds of selected fish species dniiain f usr aes atog dt ae limited are data although areas, nursery of identification grounds. spawning the overlap should grounds nursery species, oviparous an As areas. spawning few are There define to which with Sea. egg-cases of distribution the on Celtic data and Scotland of Sea, coast North west northern the of waters deeper the in occurs It distribution. historical its to comparison in waters UK in Sea. Celtic the in and Bank Rockall the on taken flossada le skate blue n occasional and with Sea, Celtic the from samples from distinct genetically generally were VIa Division ICES from Griffiths unclear. are species two the of distributions geographical current The (Iglésias ICZN the by validated be to needs hud e eurce: lpe skate flapper resurrected: be should names on scientific former working that proposed Taxonomists have problem synonym. the invalid an be name soon scientific the so and updated, being The complex. currently species is complex’ skate ‘common the the of nomenclature for are provided layers data the and species two comprises actually skate’ ‘common Iglésias by study recent A informationGeneral 3.4 Figure5: nominal of accuracy rudih uvy aa a alw o te preliminary the for allow may data survey Groundfish distribution reduced a has complex skate common The juveniles Common skate Common nursery of Nursery D. (<43 the cf. records cm D. grounds grounds bt hs rpsd change proposed this but intermedia, total cf. 21) bevd ht samples that observed (2010) al. et south ae i Va and VIa, in taken flossada length) of of common Dipturus batis- Dipturus common (2010) has revealed that revealed has (2010) al. et of the in groundfish Isle skate skate of D. Man complex, complex. D. cf. surveys. and cf. 2010). al., et and flossada will batis D. complex in as intermedia Data the Uncertainty indicated Central D. layer cf. indicates by North regarding the Improved delineation of nursery grounds is required. embryos. required. developing with egg-cases is grounds of capture the on report usefully could nursery surveys Groundfish of delineation Improved gaps Data in occur typically not <30 waters does It estuaries. in occasionally recorded is that species marine fully a is skate Common coastalzone and Estuarine grounds. nursery of delineation the from excluded been have and questionable, for considered data were Some skate common rates. catch highest the having surveys Scottish with 5), (Figure Sea Celtic and Scotland western North- off primarily captured were skate common Juvenile groundsNursery grounds. nursery with delineate to overlap broadly should which these although grounds, spawning with females egg-bearing or cases egg- the of occurrence the on data insufficient are There groundsSpawning 1-groups. and 0- to primarily refer data the so and 1926), 21 ca. of length a at hatch survey skate Common data. in apparent cohort first the with equates broadly the <43 For of range length waters). a study, present deeper in (e.g. range their of some for presence Sea. the m deep (Wheeler,deep 1978). m cm was used, which used, was cm cm (Clark, cm 17 3 Spawning and nursery grounds of selected fish species m deep the grounds Rajella fyllae), and so these records have not groundfish skates. of in et al., 2005). edge nursery Although juvenile thornback rays were also reported length) the Juvenile thornback rays were frequently taken trawl in surveys, beam with otter trawl catch catches efficiency having for these a size fish. lower Areas of to importance juvenile thornback ray included Estuary, the Bristol Channel, Greater Cardigan Thames Bay, and studies eastern earlier in suggested as 6), (Figure Sea Irish western and (Ellis from the deeper waters off North-western Scotland, there this in skates looking similar with confusion some be could area (e.g. been attributed to thornback ray nursery grounds. Estuarine and zone coastal Thornback ray is a fully marine species that occasionally in is estuaries. recorded Juveniles may occur in coastal waters, but are mostly found in waters >10 Data gaps Improved delineation of spawning and nursery grounds is required. Groundfish surveys could usefully report on the capture of egg-cases with developing embryos. Nursery Nursery grounds 1978). (Wheeler, misidentified off total nominal cm records (<18 represent indicates the may of layer juveniles of which Data accuracy Sea, ray. the presence cm (Clark, 1926; Ellis and North the by thornback regarding Raja clavata of Central the cm was used, which broadly equates in situ egg-cases (although data are in indicated grounds as uncertainty and is ray, shelf Nursery There Thornback Thornback ray thornback Groundfish survey data may allow for the preliminary Hebridean surveys. of Figure 6: Figure Spawning grounds There are insufficient data on the occurrence of the egg- cases or egg-bearing females in with which to delineate spawning grounds, although these the spawning season should broadly overlap with nursery grounds. 3.5 General information Thornback ray is a widespread skate. and There relatively are abundant few field females with egg-cases, data and there on are also the few data distribution the on distribution of of available for specimens from the strandline), and so it is not currently practical to accurately define spawning areas. identification of nursery areas. For the length present range of study, <18 a with the first cohort apparent in survey data. ray hatch at a length of ca. 10–12 Thornback 0-groups to primarily refer data the so and 1995), Shackley, (Brander 1985). and Palmer, 18 3 Spawning and nursery grounds of selected fish species with nursery grounds. nursery with delineate to overlap broadly should which these although grounds, spawning with females egg-bearing or cases egg- the of occurrence the on data insufficient are There groundsSpawning 1984). Ajayi, and (Ryland 0-groups to primarily refer data 10–12 ca. of length a at hatch ray Spotted data. survey in apparent cohort first the with a <18 study, of range present length the For areas. nursery of identification areas. spawning define accurately to which with strandline) the from specimens of (excluding egg-cases distribution the on data few are there and egg-cases, There are few field data on the distribution of females with skate. abundant relatively and widespread a is ray Spotted informationGeneral 3.6 Figure7: nursery (<18 the rudih uvy aa a alw o te preliminary the for allow may data survey Groundfish accuracy cm Spotted ray Spotted total grounds Nursery of length) the of grounds records spotted in cm was used, which broadly equates broadly which used, was cm groundfish Raja montagui Raja of ray, off spotted the as cm (Clark, 1922), and so the so and 1922), (Clark, cm surveys. edge indicated ray. of Data the There by Hebridean layer the is presence uncertainty indicates shelf. of nominal juveniles regarding capture of egg-cases with developing embryos. developing with egg-cases of capture the on report usefully could surveys Groundfish required. is grounds nursery and spawning of delineation Improved gaps Data 1978). recorded although adults are usually found is further offshore (Wheeler, that species marine occasionally in estuaries. Juveniles occur fully in coastal waters, a is ray Spotted coastalzone and Estuarine Greater Estuary.Thames the in abundant less were rays spotted juvenile (Ellis studies earlier in suggested as ray, thornback of those to similar broadly were 7) (Figure grounds nursery potential The rates. catch maximum and tows) of (50% regularly caught were they where stations fixed juveniles, of presence the for provided were Data with layers trawl. surveys, beam by of effectively variety most sampled a juveniles in taken were rays Spotted groundsNursery , 2005), although 2005), al., et 19 3 Spawning and nursery grounds of selected fish species et al., 2007). Undulate rays were generally taken in beam trawl surveys provided were layers Data 8). (Figure Channel English the in for the presence of juveniles, fixed stations were where caught regularly they (50% of tows) and maximum catch rates. Juvenile undulate ray tended to occur in the coastal fringe of the English Channel, with the the site Channel of the most Islands regular occurrence of juveniles. Data off North-eastern Scotland were Europe, in Elsewhere code). species incorrect (e.g. records considered erroneous juvenile undulate rays are typically found in coastal waters, particularly in shallow, inshore lagoons, rías zones and estuaries (e.g. such Coelho and Erzini, as 2006; Moura coastal Estuarine and zone coastal Undulate ray is a coastal, marine species that utilises the outer reaches of some estuaries as a nursery ground. Data gaps Improved delineation of spawning and nursery grounds. Inshore surveys in the identify English important Channel inshore could habitats. usefully could Groundfish usefully surveys report on the capture developing embryos. of egg-cases with Nursery Nursery grounds the surveys. by error. indicates layer groundfish indicated database Data a in as ray. et al. (2007) suggested ray, length) considered total undulate undulate of Sea cm Raja undulata of cm. Moura (<29 North grounds grounds juveniles northern of Nursery in nursery Undulate Undulate ray cm was used, as there were very few records of newly of records few very were there as used, was cm Groundfish survey data may allow for the preliminary Records presence nominal Figure 8: Figure Spawning grounds There are insufficient data on the occurrence of the egg- cases or egg-bearing females with spawning grounds, although these which should broadly overlap to delineate with nursery grounds. 3.7 General information Undulate ray has a restricted distribution within EEZ, the being UK most commonly encountered in Channel, the especially from the Channel Islands to the Solent English and coast of Sussex. There are limited field data distribution of on females with the egg-cases, and there are few data on the distribution of egg-cases (excluding specimens from the strandline) withspawning areas. which to accurately define identification of nursery areas. common This in species parts of is data the are locally limited. For English the present study, a Channel, length range of but <29 current hatched fish in these survey data. Indeed, neither the age and growth of juveniles nor the size documented, at hatching although is well Coelho and Erzini juveniles as small as 19 (2006) caught that the nursery grounds estuarine waters. of undulate ray occurred in 20 3 Spawning and nursery grounds of selected fish species rud i rltd o h oeal elh f h sok of stock the Schmidt (see of herring health autumn-spawning overall the to related is grounds smaller the of utilisation the grounds. spawning on inform always surveys not These do 2000). Aldridge, and Fox 1983; Bamber, and Dempsey (see estuary Thames still Greater the of are parts there as such and areas, certain in 9), stocks spring-spawning discrete (Figure stocks Bight) Southern main and Channel the (Eastern Downs (off and England) of Banks North-eastern Scotland), (off importance Buchan the and Orkney confirm Shetland, Sea North and use. active in are larvae can indicate which of the nominal spawning grounds herring for surveys although grounds, spawning herring of et al., 1998; Maravelias Stratoudakis 1989; Aneer, 1986; Rankine, 1980; Groot, de 1980; Bowers, (e.g. water well-oxygenated and sediment fine of proportion low a is there where shell) maerl, sand, coarse (e.g. habitats similar and gravel on spawn Herring groundsSpawning year.the of start the at conducted surveys for fish older some include will this although (2009c), ICES by used as 0-groups, identify <17.5 study,of present range the length a of purposes the For 1959). Wood, (e.g. abundant be can few herring 0-group where have waters, inshore shallow surveys in stations these that acknowledged be must it were data waters. western other for lacking comparable recent although Sea, Irish the in surveys in the North Sea and eastern Channel, and surveys coordinated internationally from distributions larvae herring some for source food 1989). Morrison, and Rankine (e.g. predators important an be also and can eggs larvae Herring areas. spawning active just not beds, spawning potential on inform would information such of all inclusion the so and time over change can beds such of use The gravel). and sand coarse of areas spawning (often beds the identifying accounts historical several are There 1976). (Russell, pelagic stages post-larval and larval EEZ. UK the in stocks nominal several are there and species fish pelagic abundant and widespread a is Herring informationGeneral 3.8 It has been suggested that the importance of spawning of importance the that suggested been has It Channel English eastern the in surveys larval Herring Juvenile herring are taken in otter trawl surveys although This information was augmented by international data on The eggs are demersal (0.9–1.5 Herring Clupea harengus Clupea et al., 2000). There are few surveys mm in diameter), and the 20) and 2009), al., et cm was used to used was cm secondary nursery grounds. nursery secondary present the in study (<17.5 length used range smallest length the the whereas by classes, used grounds nursery primary Coull by indicated grounds The offshore. further slightly extending here reported grounds nursery the with differences, slight Coull by proposed grounds nursery original the supported broadly 10) (Figure data survey current the Although rates. catch maximum data layers were provided for the presence of juveniles and and surveys, of variety a in recorded were herring Juvenile groundsNursery of herring. stocks western other inform of distributions to larval available or egg are on data few although 2001), al., et (Dickey-Collas grounds spawning the in shifts subtle been have to thought are there where surveys, ichthyoplankton measure. management appropriate an is re-colonisation Corten, these restrict not does of grounds nature (e.g. physical the time that ensuring so over and 1999) spawning ‘recolonised’ that be can recognised grounds be should it activity.Nevertheless, spawning little, very or no, have currently Bank) Dogger the of parts (e.g. grounds spawning historic some grounds. appropriate gear would be required to better inform on inshore nursery an with surveys Inshore grounds. spawning potential the of some of use contemporary the on inform better to needed are surveys spawning field Dedicated active historic). or (whether suitable spawning of herring known sites and are substrate there where grounds grounds, Coull by illustrated spawning as site-specific more as well as larvae, of distribution the by evidenced as Channel), sea/eastern North southern the in and Scotland North-eastern off Sea, Irish eastern the of parts in (e.g. areas spawning broader recognise to pragmatic more be may it stocks, constituent the for information spawning detailed Given recent, of stations. lack the survey of distribution the by influenced strongly be may larvae herring of distribution apparent The gaps Data Certain 2000b). al., Haven). Milford (e.g. areas et estuarine in spawn also stocks Power (e.g. grounds nursery as habitats estuarine utilises that species marine a is Herring coastalzone and Estuarine ern lra hv as be tkn n h Iih Sea Irish the in taken been also have larvae Herring 19) a eut wt te oe inshore more the with equate may (1998) al. et cm) would be indicative of the more extensive 19) tee ee oe very some were there (1998), al. et (1998). The latter being those being latter The (1998). al. et 21 3 Spawning and nursery grounds of selected fish species et al. total lacking. nursery cm Coull are by polygons (<17.5 spawning nominal areas international included original areas). these from indicate juveniles indicates the of data layers western support layer originally giving to for larval Data Data presence areas data the herring, recent lacking of field by herring. herring. Inshore and data of of recent grounds (1998) indicated surveys. grounds as grounds et al. comparable although spawning Coull herring, Nursery groundfish Spawning in of (recent, in retained, potential (1998) length) grounds Figure 10: Figure surveys provided and Figure 9: Figure 22 3 Spawning and nursery grounds of selected fish species cod (Fox cod Sea Irish for reported grounds also was grounds spawning spawning cod in the to fidelity identified off the west coast of high Scotland. A high site fidelity a of evidence Wright Rockall although Ireland, southern Scotland, and North-western off grounds spawning cod of distribution the examine to available data recent no were There suggested. than widespread more may slightly be Sea Irish western and eastern the in and Cornwall) (north Head Trevose off grounds spawning important the Coull by indicated of than widespread more be may variety spawning that evidence was a there limited, in taken were widespread in the North Sea, and although data were were larvae They 2008). al;. et Fox 11; (Figure surveys ichthyoplankton and eggs Cod groundsSpawning year.the of start the at conducted surveys in fish used by ICES (2009c), although this will include some older <23 of range length study,present the a of purposes the For grounds. nursery other of extent broader the on inform to used be can they of the UK, where juvenile cod are often abundant, although waters inshore the of coverage limited have may surveys these of Some fish. juvenile of distribution the examine to designed are surveys these and EEZ UK the of parts many grounds. spawning the of some of locations broad the on inform can data such Nevertheless, assured. as be not may data historic more of identification the of accuracy Taylor(see genetics to better identify and quantify cod eggs in samples eggs almost completely overlap. Recent studies have used haddock and cod of range size the and whiting, of those easily be confused with those of other gadoids, particularly can eggs cod Smaller size. than other stages, early the in features distinguishing no have eggs the as problematic 4.5 approximately of length a at begins absorption) yolk-sac (following stage post-larval (1.16–1.89 pelagic are areas. eggs The many for available are cod of larvae and eggs the of distribution the on information limited area, study entire IIIa). and VIId Divisions IV, Subarea North (ICES waters the adjacent and and sea (VIIe-k) south-west Rockall (VIIa), VIa), Sea Irish Division EEZ, (VIb), (ICES UK Scotland the of in west stocks including cod nominal several are There informationGeneral 3.9 em rw ad te tal uvy oeae around operate surveys trawl otter and trawl Beam is eggs cod of identification the that noted be should It the cover not do data survey ichthyoplankton Although Cod , 2000). al., et , 2002; Goodsir 2002; al., et Gadus morhua Gadus cm was used to identify 0-groups, as 0-groups, identify to used was cm mm (Russell, 1976). (Russell, mm , 2008), although the although 2008), al., et mm in diameter), and the and diameter), in mm (1998). Similarly, (1998). al. et (2006) found (2006) al. et rswtr nlec ad hl wtr ass wt the fronts. with haline these near peaking larvae of abundance masses, water shelf a and with influence masses freshwater water those between zones frontal North in especially the the Bight), German in and areas Bank Dogger shallow (e.g. Sea in occurring as larvae cod these sites may serve as important cod nursery grounds. nursery cod important as serve may sites these of some although sites, near-shore sample always not do 1990. to from were Channel Bristol subject be usefully could the for data recent most the Indeed, investigations. further Sea) North and English western Channel, the of parts neighbouring Bristol and (e.g. Channel areas other Additionally, Rockall). Scotland, Channel, English North-western Channel, George’s St eastern and Ireland southern (e.g. report that in described grounds spawning nominal the of some on importance the data are limited. Further field data are still required to determine supporting the although widespread, more be may Coull grounds in spawning given cod of distribution patchy the within activity spawning major of evidence recent no was There gaps Data Rogers 1978; (Wheeler, grounds nursery as waters coastal other and habitats estuarine utilises that species marine a is Cod coastalzone and Estuarine Clyde). of Fifth and Sea Irish eastern (e.g. UK the of seaboard western the along widely and Firth) Moray the (including Sea North the in distributed more grounds nursery important potentially with (e.g. Gibb Coull by proposed grounds nursery original the supported broadly data survey current cod the Although juvenile 12). (Figure rates of catch maximum presence their and the for provided were layers data and surveys, of variety a in taken were cod Juvenile groundsNursery In terms of nursery grounds, current groundfish surveys Munk that noted be also should It , 1998). al., et et al., 2007), there were some slight differences, 19) Sann i te ot Sea North the in Spawning (1998). al. et (1998) and other sources other and (1998) al. et (2002) reported (2002) al. et 23 3 Spawning and nursery grounds of selected fish species grounds groundfish retained, in are (1998) nursery lacking. length) et al. comparable (1998) are spawning total nominal Coull et al. in cm (recent, areas Coull indicate (<23 indicate by these areas). surveys provided of layers layers juveniles included Data some of western Data polygons cod. and cod. of support of ichthyoplankton originally presence to original the the from southern data grounds areas by grounds data field some giving inshore for cod, larval Nursery Spawning indicated recent The of as lacking recent cod, although surveys. of Figure 12: Figure data and grounds Figure 11: Figure 24 3 Spawning and nursery grounds of selected fish species hne ad f Nrhwsen ctad n southern and Scotland Ireland. North-western off and Channel examine to the distribution of whiting spawning grounds in the English data recent no were There suggested. widespread than more slightly be may Sea Irish eastern western the and in and Head Trevose off grounds spawning widespread more than indicated by Coull be may spawning that there evidence limited, was were of data although and variety Sea, North a the in in taken were ichthyoplankton surveys larvae (Figure 13). They were widespread and eggs Whiting groundsSpawning year.the of start the at conducted surveys for fish older some include will this although (2009c), ICES including <20 of range length fish, a study, present the juvenile of purposes of the For whiting. distribution the examine to designed are surveys these and EEZ UK the of parts many existence. demersal more a phase to switching before jellyfish with associated often are they post-larval The 4.3 about of length a at development. begins early in features pelagic of are locations eggs The broad (0.97–1.32 grounds. the spawning the on of inform some can data available Nevertheless, above). (see problematic is gadoids some of areas, although many it must be noted that for the identification of the available eggs are whiting of the larvae of and eggs distribution the on information area, study entire IV,Subarea VIId). (ICES Division waters adjacent and Rockall VIa), Division sea North the and (VIIe-k) south-west (VIIa), Sea Irish (VIb), (ICES Scotland of west including EEZ, UK the in stocks whiting nominal several are There informationGeneral 3.10 em rw ad te tal uvy oeae around operate surveys trawl otter and trawl Beam the cover not do data survey ichthyoplankton Although Whiting mm in diameter) and possess no distinguishing no possess and diameter) in mm cm was used to identify 0-groups, as used by used as 0-groups, identify to used was cm Merlangius merlangus Merlangius et al. (1998). Similarly, the important mm (Russell, 1976), when 1976), (Russell, mm under-represent wider spawning activity.spawning under-representwider may surveys some of extent (temporal) limited the so and (i.e. spawner extensive), temporally and spatially both be may serial spawning widespread, a is whiting also that is noted It eggs. whiting identify better to surveys recent more the of some allowed have techniques genetic New morphologically,species microscopy.to light identify using to impossible are eggs the as analysis, sample of artefact an is eggs whiting of distribution the on data of lack The gaps Data (Wheeler, grounds Potter 1978; nursery as waters coastal other and habitats estuarine utilises that species marine a is Whiting coastalzone and Estuarine previously.reported than offshore further slightly extending possibly grounds nursery the with differences, slight very some were there Coull by proposed grounds nursery the original supported broadly data rates. survey catch current maximum the their Although and tows) of 70% and (50 regularly caught were they where stations fixed whiting, juvenile of presence the for provided were layers Data 14). (Figure surveys of variety a in taken are whiting Juvenile groundsNursery , 1988; Henderson & Holmes, 1989) Holmes, & Henderson 1988; al., et (1998), al. et 25 3 Spawning and nursery grounds of selected fish species et areas. Coull total nursery by cm et al. these of spawning (<20 nominal (recent, lacking). Coull included in some are for indicate surveys indicate juveniles areas of originally provided layers lacking layers areas are Data western Data presence polygons and data the inshore ichthyoplankton whiting. by field whiting. The original of from of southern the data indicated although surveys. grounds some as giving grounds larval for shown, data whiting, whiting, Nursery Spawning recent groundfish also of of in and (1998) al. length) grounds Figure 14: Figure comparable (1998) grounds Figure 13: Figure 26 3 Spawning and nursery grounds of selected fish species iey ht ot f h sann atvt tks lc in place takes activity sampled. regularly not are that UK the of west the to areas spawning the of most that is likely it fishery, ‘industrial’ the from catches commercial on based stock, whiting blue the of size the given Hatun However, 1997; Coombs, and Bartsch 1992; Demenin, and Gerber (e.g. authors various by described been have grounds spawning The available. was species this for grounds spawning the update to information new Consequently,15). (Figure report this in utilised surveys no the from samples few in observed been have whiting blue The eggs are difficult to positively identify, and the larvae of groundsSpawning outer the of slope. and shelf continental waters deeper in abundant most species, bentho-pelagic abundant and widespread a is whiting Blue informationGeneral 3.11 can be variable. be can floor habitats (where they may be taken in demersal trawls) sea of occupancy of degree and migrations such of extent The migrations. vertical extensive have may whiting blue that noted be also should It surveys. these in caught are range stock the of parts shallower the in fish the only that noted be should it although juveniles, catch will surveys trawl Otter (2009c). ICES by used as 0-groups, identify to <19 of range length a study, present the of 6.7 approximately of positive aid length a at begins phase to post-larval the and identification, features distinguishing no possess and h eg ae eai (1.04–1.28 pelagic are eggs The Blue whiting Blue mm (Russell, 1976). For the purposes the For 1976). (Russell, mm Micromesistius poutassou Micromesistius

mm in diameter) diameter) in mm cm was used was cm 2009). al., et elucidated. be to needs still whiting blue juvenile for waters offshore deeper, of importance the and grounds, the nursery of potential part a on inform only surveys groundfish existing edge of the Scottish and Irish continental shelves. Similarly, and larvae of blue whiting, as the spawning eggs grounds are off the the sample occasionally only surveys Existing gaps Data >50 waters in offshore, recorded is that further species Typicallyoccurs estuaries. in occasionally very only marine fully a is whiting Blue coastalzone and Estuarine the edge of the continental shelf than previously supposed. along south further extend may ground nursery their that suggesting Scotland, North-western off made also catches high were and Sea North northern the in widespread Coull by proposed grounds nursery the than 16). inshore and southerly (Figure rates more were catch study current the in used surveys the maximum of Most and tows) of 70% and (50 regularly caught were they where stations fixed and data layers were provided for the presence of juveniles, Juvenile blue whiting were recorded in a variety of surveys, groundsNursery 19) Jvnl bu wiig ee more were whiting blue Juvenile (1998). al. et m deep. m 27 3 Spawning and nursery grounds of selected fish species of larval spawning grounds recent main species. and the available. his included for spawning for juveniles (1998) were nominal of data originally et al. indicate updated available indicate not Coull areas presence were layers in layer are the supporting data Data by no Data Offshore (1998) provided recent whiting. et al. No indicated whiting. although surveys. as blue polygons Coull blue of in of surveys. included, whiting, original groundfish layers grounds in also the grounds blue data of (1998) the giving length) Nursery Spawning ichthyoplankton and et al. grounds total whiting, from cm Coull grounds, by data (<19 blue nursery Figure 16: Figure Figure 15: Figure 28 3 Spawning and nursery grounds of selected fish species Bailey Bank) is not fully understood. fully not is Bank) Bailey Bill and Bank Faroe Bank, Rockall (e.g. grounds potential other and these of range exact the although 17), (Figure 730–1100 of waters in Banks Lousy and Hatton off the continental slope of Scotland, and off the known Rosemary, are grounds Spawning 2009). spawning (CEC, protect aggregations to now measures are management grounds to spawning subject ling blue and 2010), al., et (Large identified been have activity spawning high of sites inshore more species. this of eggs the sample not in do so and waters, generally are surveys ichthyoplankton Existing 1982). (Ehrich, Bank Lousy and 1969), Rockall (Wheeler, including Sea North northern the of parts in and Islands, Faroe the deeper off Scotland, the North-western off in waters occur ling blue of grounds spawning The groundsSpawning found. rarely are that larvae and eggs planktonic have They surveys. very groundfish taken in only occasionally is such as and slope continental the abundant on most is that species deep-water a is ling Blue informationGeneral 3.12 Figure17: continental Hatton other Earlier fisheries targeted spawning aggregations. Known Blue ling Blue possible Bank; Spawning slope and spawning of (5) Scotland; Molva dypterygia Molva southern grounds grounds of (2) and blue north-west is eastern not ling. fully Some margin known, of Rosemary of of the and the sites depth m no Hatton Bank, data of blue (3) Bank layers south-west ling collected during internationally coordinated deep-water deep-water surveys. coordinated be internationally usefully could during species collected this on data biological More gaps Data >150 waters in usually offshore, occurs ling Blue coastalzone and Estuarine ling. Spanish with confused be may data available of juvenile blue ling in UK waters. Additionally, some of the distribution the map to information insufficient was There groundsNursery (Large were spawning provided. et al., et of the activity 2010). Lousy Figure have The Bank; adapted overall been (4) identified distribution north-east from Large from margin of et al. et (1) these along (2010). of and the the m deep. m 29 3 Spawning and nursery grounds of selected fish species m deep. Ling is a fully very occasionally marine in estuaries. species It typically that offshore, in waters occurs >50 further is only recorded Data gaps Ling eggs and ichthyoplankton larvae surveys were in only western in taken small waters, numbers. in Data although firm were a conclusions on too spawning in limited the varietynorthern to North Sea, draw of in the deeper waters of the western Channel and Sea Celtic and western seaboards Eggs were found in of samples collected from the Ireland Irish Sea and and off Trevose Head, Scotland. with larvae also taken in the latter area. Although these grounds are identified here as spawning ling grounds, it should be recognised more widespread in that the deeper waters ling of the continental is shelf and along the continental slope, and there may more extensive be and important spawning grounds in these areas. Estuarine Estuarine and zone coastal cm cm was examined. examined. was cm mm in diameter and Molva molva Ling Trawl Trawl surveys are inefficient at sampling the smallest mm (Russell, 1976). Nursery Nursery grounds Ling has not traditionally been considered commercial an important species and subject to routine biological consequentlysampling. For the purposes of has not the present study, the length cut-off beenbroadly equated with information ground nursery nominal the so and juveniles all refers to secondary nursery grounds (Figure 19). Catches of the smallest length-classes of ling were generally small. Juvenile ling were recorded in a variety of data layers were provided surveys, for the presence of and juvenile ling and maximum catch rates. It was not possible to highlight fixed stations where they were caught regularly, due the sporadic nature of to captures. Current data suggest that juvenile ling are taken in the northern North Sea deeper in catches largest the with Scotland, and North-western off eastern the in ling juvenile of records apparent The waters. English Channel may represent misidentified rockling. Spawning grounds occasionally recorded only were ling of larvae and eggs The in existing ichthyoplanktonspawning some suggest to data some were there although surveys (Figure from data more Sea, Irish and Channel Bristol the in activity 18), and the outer continental shelf are required to the better main define spawning grounds for this species. Hence, the data layer generated for ling is missing the main spawning grounds. 3.13 General information waters deeper the in abundant, not but widespread, is Ling of the continental shelf. The eggs are difficult to positively identify. They are pelagic, 0.97–1.13 contain a single oil globule (Russell, 1976). Egg stages are only recorded sporadically from ichthyoplankton surveys, presumably due to the main spawning grounds occurring in deeper waters to the north and west of the British Isles. The post-larval stage begins at a length of approximately 5 juvenile stages (i.e. distribution the indicate may primary which taken are nursery juveniles larger grounds), although of secondary nursery grounds. For <49 ling of the occurrence the study present purposes of the Although the length at maturity is not reliably known for ling, recent field surveys have indicated that fish <49 are immature. 30 3 Spawning and nursery grounds of selected fish species Figure18: Figure19: potential nursery (<49 some grounds cm inshore total grounds are inshore Spawning Nursery thought length) surveys, of spawning ling, grounds to in grounds and groundfish occur as it indicated grounds of should offshore, of ling. ling. surveys. Data of be by Data ling, where noted the layers layers Data presence that field indicate were indicate the data of main only are juveniles nominal some available lacking. spawning for 31 3 Spawning and nursery grounds of selected fish species m deep. Juvenile hake are taken in a variety of surveys, and data layers were provided for the presence of fixed juvenile stations hake, where they were caught regularly (50 data Current rates. catch maximum their and tows) and of 70% suggest that juvenile hake are taken in the northern North Sea and along the western seaboard of the British (Figure 21). Isles Estuarine and zone coastal Hake is a fully very marine occasionally in species estuaries. It that typically offshore, in is waters occurs >50 further recorded only Data gaps Although only limited data were available proposed for grounds west hake, of the Ireland and in the are widely acknowledged as Celtic important spawning grounds. Sea Field surveys off North-western Scotland are required to better determine the extent of potential hake spawning in this region. Nursery Nursery grounds cm et al. (2003) mm in diameter and m isobath and the shelf et al., 2004). The extent of mm (Russell, 1976). Merluccius merluccius m have been identified (Figure 20), as m, although there could be some latitudinal et al. (2004) reported that the North-eastern et al., 2001; Alvarez European hake European For the purposes of the present study, those sites Alvarez Trawl Trawl surveys sample juvenile stages, which may any spawning north of Scotland is unclear, although Howell although unclear, is Scotland of north spawning any (1921) reported eggs and newly hatched larvae just north of 58ºN. edge. between 200–1000 well as some of the outer continental shelf authors other by reported of were larvae and the eggs where Sea, Celtic (e.g. Fives Spawning grounds waters deeper the in occasionally captured were eggs Hake Head, Trevose off and Man) of Isle the (near Sea Irish the of where larvae were also taken (Figure 20). Although these are hake grounds, spawning hake as identified are grounds the of waters deeper the in abundant and widespread more continental shelf and along the continental slope and there will be more extensive and important spawning grounds in these areas. Atlantic stocks of hake generally spawned of 200–1000 over depths variation in the depth of spawning. reported hake Olivar spawning in the Mediterranean was most pronounced between the 100 3.14 General information continental the of waters deeper the in widespread is Hake shelf. The eggs are pelagic, 0.94–1.03 contain a single oil globule. The post-larval stage begins at a length of approximately 5 indicate the distribution of nursery grounds, although the the For low. often is areas these in stations trawl of density purposes of the present study, a length range of <20 was used to identify 0-groups, as used by ICES (2009c), although this will include some conducted at the start of the year. older fish for surveys 32 3 Spawning and nursery grounds of selected fish species inshore survey Figure20: Figure21: nursery 200-1000 (<20 cm data. spawning total grounds m, Spawning Nursery The but length) supporting main of grounds hake, grounds in grounds spawning groundfish as of data of indicated hake, of hake. hake. grounds were surveys. as Data by not inferred Data the occur available. layer layers presence from offshore, indicates indicate ichthyoplankton of in juveniles nominal some depths of of the 33 3 Spawning and nursery grounds of selected fish species Juvenile anglerfish are taken in a variety of surveys, albeit in low numbers (Figure 23) and data layers were provided for the presence of juveniles, fixed stations were where they caught regularly (50 and 70% maximum catch of rates. Current tows) data suggest and that juvenile their anglerfish are taken in the northern North Sea and along the western seaboard of the British Isles (including North- western Scotland, parts of the Irish Sea and the Cornish peninsula). Estuarine and zone coastal Anglerfish is a fully marine species that is recorded only very occasionally in estuaries. coastal Juveniles waters, may occur although in adults offshore. tend to occur further Data gaps Very little is known about the egg and larval distribution, and of mature females in spawning condition, as catches of these stages are very survey of degree high a of absence sporadic. the in available become Data are unlikely to effort. Nursery Nursery grounds et al., 2000, 2001; Arkhipov and Lophius piscatorius cm was examined, which included the Anglerfish Anglerfish cm are recorded infrequently in surveys. Anglerfish spawn in deep water, with most records of Mylnikov, Mylnikov, 2002; Laurenson, 2006). gears, other or Very samplers plankton in caught few been have eggs ribbons of and too few larvae have been captured in ichthyoplankton surveys to identify specific sites of spawning. There is very limited information (Figure 22), but they are presumed to spawn in deep water along the edge of the continental slope (Hislop Spawning grounds General General information Anglerfish is widespread in the continental shelf deeper and slope, waters but small of numbers the taken are in shallow also waters. Both beam trawl and otter trawl surveys catch low numbers of this species, although Hence, the infrequently. very taken are categories size smallest trawl surveys may be indicative of the broader secondary nursery grounds on the continental relative importance of these sites within large shelf, un-surveyed although the areas is unclear. For the purposes of the present study, a length range of <28 first main peak and broadly equates with 0-group and some some and 0-group with equates broadly and peak main first 1-group fish. However, it should be noted that anglerfish <10 spawning in UK waters from northern and western areas. The eggs are included within even the a post-larval stage gelatinous may have ribbon, a and prolonged pelagic existence. 3.15 34 3 Spawning and nursery grounds of selected fish species Figure22: Figure23: larvae nominal are presence to occur available taken nursery in of Spawning Nursery deep-water. juveniles in to current identify grounds grounds grounds (<28 ichthyoplankton spawning of cm anglerfish, of total of anglerfish. anglerfish. grounds, length) as surveys. Data indicated in which Location groundfish layers Insufficient are by of considered indicate the anglerfish surveys. data 35 3 Spawning and nursery grounds of selected fish species Juvenile horse mackerel were recorded of presence the for provided were layers data and surveys, in a variety of juveniles, fixed stations where they were caught regularly (50 and 70% of tows) and maximum catch rates. Juvenile horse mackerel were distributed widely around the although UK, the lack of beam as sampling, the of records artefact an considered is Channel from the eastern the Given area. this English in used gear main the are surveys trawl extensive occurrence of juvenile horse mackerel (Figure 25), there was no evidence for spatially-defined grounds, and so no nursery data layers were provided. Estuarine and zone coastal in common is that species marine fully a is mackerel Horse coastal waters, but only recorded occasionally in estuaries. Data gaps project, the during available not were Sea North the for Data although surveys do operate in this area (coordinated by WGMEGS). The apparent absence of larvae waters is an in artefact of sampling protocols, western as this survey focuses on egg stages, and larvae to are species. rarely identified Some participatingsamples, and so analyses of the laboratorieslarvae of horse mackerel retain the (and other species) could samples, resource permitting. be undertaken for archived Nursery Nursery grounds and as layers survey Data egg , 2001; Fives 2001; al., et mackerel, mm (Russell, 1976). (Russell, mm Sea. mm in diameter, and mackerel. horse mackerel of North horse Trachurus trachurus for of cm was also used. triennial grounds limited , 2003). Nevertheless, there was there Nevertheless, 2003). al., et grounds Data spawning international the Spawning in surveys. nominal Horse mackerel Horse mackerel cm was used to identify the sites of 0-groups, as Surveys with PHHT and GOV trawls, which have high The eggs are pelagic (0.8–1.04 national observed indicate Figure 24: Figure , 2001; Abaunza 2001; al., et Head also and evidence in of spawning activity off Trevose (see Bight Southern and Channel English south-eastern the 1974), and Macer, some spawning in the Irish Sea. Spawning grounds Horse mackerel eggs international triennial are mackerel taken egg emphasising survey the importance of the extensively outer continental shelf (Figure 24), in (Coombs species this of spawning the for the contain a single oil 3.5 approximately of length a at globule), begins and the post-larval stage 3.16 General information Horse mackerel is a widespread and abundant pelagic fish species. ICES currently assess three stocks, including North a Sea stock (in ICES Divisions IIIa, IVb,c and VIId) and a western stock (in ICES Divisions IIa, IVa, Vb, VIa, VIIa-c, e-k, VIIIa,b,d,e). headline heights, are considered appropriate for providing range length a Initially, species. this of juveniles the on data of <15 used by ICES (2009). distribution However, of horse given mackerel within the this length range, widespread a reduced length range of <8 36 3 Spawning and nursery grounds of selected fish species Figure25(b): indicates groundfish Figure25(a): indicates groundfish with no spatially catches catches surveys. surveys. Nursery Nursery discrete of of This Horse horse horse grounds grounds size nursery mackerel mackerel mackerel class of of horse horse grounds. is (<8 (<15 appear also mackerel. mackerel. cm cm very total to total be widespread. length) widespread length) Data Data layer layer in in and 37 3 Spawning and nursery grounds of selected fish species et al. (1998) reflected the increased management Juvenile sandeels were taken presence the for provided were layers data and 27), (Figure in a variety of of surveys juveniles, stations where they were (50% caught of regularly tows) and their maximum catch data suggest rates. that juvenile Current sandeels are more extensively distributed than previously reported, although the apparent absence from the western English Channel and Irish Sea in Coull interest in the North Sea sandeel stocks at that time. Estuarine and zone coastal One species, Ammodytes tobianus, estuaries, with this is and other species occurring often in coastal found in waters. Data gaps More comprehensive ichthyofaunal surveys of sandbank habitats and the utilisation of such habitats could by be sandeels undertaken, as areas of sandbank are not often comprehensively surveyed in existing field investigations. Although more data are available for the North Sea ICES, (e.g. 2010), high resolution data with which to accurately delineate spawning grounds in other to areas become are available in unlikely the absence of a overall the that assumed be could it although effort, survey high degree of distribution of adult sandeels would broadly include both spawning and nursery grounds. Nursery Nursery grounds cm was used, which , 2002). Given the demersal egg demersal the Given 2002). al., et et al. (1998), although there are also Sandeels Sandeels Ammodytidae The eggs are demersal, which makes the identification Preliminary investigations on the occurrence of Spawning grounds demersal the on data comprehensive no are there Although spawning grounds of sandeels per se, given the demersal egg stage and sediment preferences of the species, the presence of larvae and juveniles spawning grounds. will Sandeel larvae (and likely some eggs) were overlap the taken in a variety 26) of which ichthyoplankton broadly surveys, corroborated suggested the (Figure by Coull spawning grounds important spawning grounds for Sea sandeels (e.g. Liverpool Bay in area) and the in the Irish Bristol Head. including off Trevose Channel, 3.17 General information There are five species these of are sandeel widely in distributed and UK habitats. Neither beam abundant nor waters otter trawls on are very and effective suitable for examining their relative abundance, especially for the smallest life-history stages. Sandbanks and other substrates sandy may be important habitats for these species, and some of these areas are not effectively existing groundfish surveys. sampled in of precise spawning grounds difficult, although plankton samplers will occasionallypresumably been disturbed from the sediment. The larval capturesurveys. ichthyoplankton in taken are and pelagic are stages eggs that have the smaller size classes may importance indicate as nursery general grounds. areas For the of purposes of the present study, a length range of <13 includes the first main cohort, but will likely include fishes (Bergstad 0-3 ages of stage of sandeels and their sedimentary requirements, for the that assumed was it study present the of purposes the distribution of the nursery grounds may also represent the extent of the spawning grounds. 38 3 Spawning and nursery grounds of selected fish species Figure26: Figure27: spawning nominal et al. et of comparable sites juveniles originally (1998) nursery grounds Spawning Nursery (<13 and data included recent cm grounds are of grounds total lacking grounds sandeels, larval by length) of Coull sandeels, of for data of sandeels. giving north-western sandeels. in et al. et from groundfish the as (1998) ichthyoplankton Data indicated original Data also layers surveys. and layers shown. polygons by south-western indicate the indicate North surveys. presence provided Sea nominal Recent, areas). in Coull 39 3 Spawning and nursery grounds of selected fish species et al. (1998), although they Juvenile mackerel were recorded in a variety of otter trawl surveys (Figure 29), and data layers were provided for the presence of juveniles, fixed stations caught regularly (50% of tows) and where maximum catch rates. they were The locations of juvenile mackerel were fairly consistent with that proposed by Coull Estuarine and zone coastal coastal in common is that species marine fully a is Mackerel waters, and is only recorded occasionally in estuaries. Data gaps project, the during available not were Sea North the for Data although surveys do operate in this area (coordinated by WGMEGS). The apparent absence of larvae waters is an in artefact of sampling protocols, western as this survey focuses on egg stages, and larvae to are species. rarely identified Some participatingsamples, and so laboratories analyses of the larvae retain of mackerel (and the other species) could be undertaken for archived samples, resource permitting. Nursery Nursery grounds were taken over greater areas of the North and Irish Seas. The largest catches were taken in North-western Scottish Scotland. surveys As off a highly there migratory may be species, some annual and seasonal variation absence apparent The areas. in nursery of importance relative the southern and Channel English eastern the from juveniles of North Sea is considered an artefact of the there have been fewer otter trawl surveys sampling, in this area. as cm et al., 2001; mm (Russell, mm in diameter, and et al., 1995; Coombs Scomber scombrus Mackerel Mackerel et al., 2001). Nevertheless, there was also evidence The eggs are pelagic (1.0–1.38 Fives of some spawning activity off Trevose Head south-eastern and English Channel, as well as in in the Irish Sea. the Data for examining mackerel spawning in the North Sea were unavailable. Spawning grounds Mackerel eggs were taken mackerel extensively in egg the triennial surveyimportance of (Figure the outer 28), continental of the shelf British emphasising Isles to for the mackerel spawning the west (Coombs Mitchell, and 1981; Priede 3.18 General information Mackerel is a widespread species. Surveys with PHHT and andGOV trawls, which both abundant pelagic have a fish high headline height, are considered for providing appropriate data on the larger juveniles of this species. For the purposes of the present a study, length of <24 contain a single oil globule), and an adult-like appearance is apparent at a length of approximately 21 1976). There are dedicated, (by internationally WGMEGS) surveys coordinated to examine the abundance of distribution mackerel eggs and west of the British Isles for stock assessment purposes. was used to identify 0-groups, as used by ICES (2009c), although this will include some conducted at the start of the year. older fish for surveys 40 3 Spawning and nursery grounds of selected fish species Figure28: of surveys. and Figure29: nominal presence Appropriate mackerel, so the nursery Comparable of Spawning Nursery original juveniles field as observed grounds data data grounds grounds data (<24 limited layer in of cm for the mackerel, of from for North total of mackerel. international mackerel. southern length) Coull Sea as were et al. et Data indicated in coasts Data groundfish triennial not (1998) layers layers of available by UK. has indicate the mackerel indicate surveys. not during been nominal egg the updated survey course spawning for and of this national the grounds area. project, 41 3 Spawning and nursery grounds of selected fish species cm) would et al. (1998), there were some et al. (1998) would broadly equate with very slight differences, with the nursery grounds presented grounds nursery the with differences, slight very here extending slightly further indicated by Coull offshore. The grounds the perception of primary nursery grounds, whereas the length range used in the present study be (<12 indicative of the more grounds. extensive secondary nursery Juvenile plaice were taken in a variety of surveys (Figure 31), and data layers were provided for juveniles, fixed stations the where they were caught presence regularly of the Although rates. catch maximum their and tows) of (50% current survey data broadly supported the original nursery grounds proposed by Coull Estuarine and zone coastal Plaice is a marine species that utilises estuarine habitats and coastal zones as nursery grounds (e.g. Power et al., 2000a). Data gaps Although there are extensive ichthyoplankton data for the Irish Sea, more recent data for the Bristol Channel could usefully be collected. Furthermore, further ichthyoplankton sampling in of the southern and could central usefully North be Sea collected, including the waters more (e.g. inshore Greater Thames Estuary, Although the The data used in the Wash present study for etc.). juveniles indicated data gaps in the southern North Sea, studies on-going of Young Fish Survey data will address some these gaps. of Nursery Nursery grounds mm in diameter), et al., 2000), with eggs and et al. (1990)), eastern English mm (Russell, 1976). Although

Pleuronectes platessa , 2007), although the distribution of juvenile of distribution the although 2007), al., et Plaice Plaice cm was used to identify 0-groups, as used by ICES For the purposes of the present study, a length of In general, the youngest plaice occur in inshore areas Plaice eggs are pelagic (1.66–2.17 Spawning grounds Existing data from ichthyoplankton surveys confirms the importance of parts of Southern Bight; the Coombs North Sea Channel, Head Trevose and eastern and western Irish Sea (including the (e.g. Ellis and Nash, 1997; Fox larvae also more widely distributed in these areas (Figure 30). A current Defra-funded programme (MA003) will help identify, with some certainty, the location grounds in the Irish Sea. of No recent data were spawning available to inform on the potential spawning grounds in the western English Channel, and off southern Ireland. North-western Scotland and ichthyoplankton survey data do not cover the entire study area, information on the distribution of the eggs and larvae of plaice are available for many areas, and these data can inform on the broad locations of some of grounds. the spawning <12 (2009c), although this will include surveys conducted at some the start of older the year. Inshore beam fish for trawl surveys operate around many parts of the coast of England and Wales, and these surveys are examine the distribution of juvenile plaice designed and sole. to in defined nursery grounds (e.g. Poxton and Nasir, 1985; Wennhage habitats increases as they attain larger sizes and 1984). Lucassen, start and (Lockwood stock to parent the to recruit Data from the Young Fish Survey will allow for improved regional data for parts of the UK coastline to be mapped at a finer resolution in the future. and the demersal post-larval stage of begins at approximately a length 13 3.19 General information There are several nominal stocks of plaice in the UK EEZ, including Celtic Sea (ICES Divisions VIIf and g), Irish Sea (VIIa), eastern English Channel Channel (VIIe) and North Sea (Subarea (VIId), IV) stocks. western English 42 3 Spawning and nursery grounds of selected fish species Figure30(a): Figure30(b): grounds spawning information and activity comparable recent in of the grounds egg plaice, omitted data Spawning Spawning Irish and lacking giving of Sea. larval to plaice. better grounds grounds the for data Figure some original illustrate from of of north- plaice. plaice. as ichthyoplankton polygons above, areas western Data Data but of provided layers layers higher point spawning surveys indicate indicate source spawning in Coull grounds). (recent, spawning et al. et (1998) 43 3 Spawning and nursery grounds of selected fish species data nominal juveniles field UK. of the of indicates presence Appropriate layer the waters by Data surveys. inshore plaice. indicated of as groundfish shallower in the grounds plaice, of of length) much Nursery total grounds for cm limited (<12 nursery Figure 31: Figure 44 3 Spawning and nursery grounds of selected fish species Juvenile sole were taken in a variety of surveys (Figure 33), groundsNursery Channel. English western the for limited were data and potential spawning grounds in other parts of the North Sea, Child 32; (Figure Estuary Thames Greater and Channel English Bay,Cardigan Sea, Irish eastern the Trevose eastern Head, Sole spawning grounds are mainly in the shallow waters of groundsSpawning present <13 of length the a study In sole. juvenile of distribution the to examine designed are and Wales, and England of coast the of (demersal) 7 about of length a post-larval at begins stage The globules. oil small distinctive (1.0–1.6 eggs pelagic The grounds. spawning the of some of inform locations broad can the on data these and areas, many for available are larvae and eggs sole of distribution the on data area, study stocks. IV) (Subarea Sea North and (VIIe) Channel English western (VIId), Channel English eastern Irish (VIIa), Sea g), EEZ, and VIIf UK Divisions the (ICES Sea in Celtic the stocks including sole nominal several are There informationGeneral 3.20 Figure32(a): grounds (1998) (recent, Inshore beam trawl surveys operate around many parts many around operate surveys trawl beam Inshore Although ichthyoplankton surveys do not cover the entire et al., 1991). Few data were available to inform on the Sole and comparable of recent sole, Solea solea Solea Spawning giving egg data and cm was used to identify juveniles. identify to used was cm the grounds lacking larval original m imtr cnan many contain diameter) mm data of for polygons sole. parts from mm (Russell, 1976). (Russell, mm Data of ichthyoplankton provided the layers North indicate in Sea Coull are surveys spawning et al. et lacking). collected by larger research vessels. research larger by collected Thames) could usefully be the undertaken to complement data and Bay Liverpool Channel, Bristol the in (e.g. water shallow sample better to in surveys spawn so and also water, shallow may Sole Channel. English western be and usefully could 1990) since surveyed (not Channel Bristol information the for collected spawning recent More gaps Data coastal zones as nursery grounds (Claridge & Potter, 1987). Sole is a marine species that utilises estuarine habitats and coastalzone and Estuarine (Riley,1974) Estuary Thames Greater and 2006) al., et parts of the English Channel (Eastwood Seaby,and 2005), Henderson 1995; Rogers, and Symonds Potter,and 1987; (Claridge Channel Bristol Wales1992), (Rogers, North including studies, earlier in identified areas in grounds nursery important confirms study present The grounds. nursery secondary extensive more the indicate grounds, nursery would study present the in used primary range length the whereas with correspond (1998) al. et Coull by indicated grounds The offshore. further extending here reported grounds nursery the with differences, minor Coull by proposed grounds the nursery original Although the supported broadly data rates. survey current catch maximum and tows) of 70% and (50 regularly caught were they where stations fixed and data layers were provided for the presence of juveniles, (1998), there were some were there (1998), al. et et al., 2003; Gilliers 45 3 Spawning and nursery grounds of selected fish species data UK. field nominal juveniles the spawning of of source indicate indicate higher point waters layers of Appropriate presence but layers Data the areas inshore by above, Data surveys. sole. as of sole. illustrate of indicated shallower Figure as groundfish grounds the better in sole. of to grounds sole, Sea. of of much length) Irish Spawning omitted for Nursery grounds the total grounds in cm limited activity information spawning Figure 32(b): Figure are (<13 nursery Figure 33: Figure 46 3 Spawning and nursery grounds of selected fish species Sole Plaice (Western) Mackerel Sea) (N Mackerel Sandeels mackerel Horse Anglerfish Hake Ling whiting Blue Whiting Cod Ireland NW ~ Mourne ~ Clyde ~ Scotland NW ~ Ireland SW ~ England SE ~ Banks/Dogger ~ Buchan/Shetland ~ Herring Peak spawning Peak Spawning Undulate ray Undulate ray Spotted Thornback ray Thornback Common skate-complex Common Tope Spurdog Species Table3: overall their only given. is season spawning so and species, other the of some for limited are well data identified, be to times spawning peak are the times spawning enabling considered, species the of some and for documented Although months. cycle 22 reproductive about the of period gestation a has for found which be spurdog year,of the case of the much in particularly can pups carrying females so and viviparous species are these spurdog), of and case (tope the sharks the two In for the 3. Table season in given spawning are species the case-study of timings general The 3.21 2010; (5): Spawning seasons Spawning Spawning Korta et al., et times 2010; of the Jan (6)   ? ? Domínguez-Petit case-study Feb    ? ? species Mar   ? ? Viviparous species (gravid females can be found all year) all found be can females (gravid species Viviparous year) all found be can females (gravid species Viviparous et al., et (Sources: Apr 2010;    ? ? ? (7) (1) May Duarte Holden,       ? ? Jun et al., et 1975;      ? ? 2001; (2) Jul    Holden ? ? (8) Abaunza Aug et al., et  ? ? ? 1971; et al., et Sep ? ? (3) 2003). Oct Coull ? ? et al., et Nov ? ? 1998; Dec (4) ? ? Genner Source (5-6) (3) (3) (7) (4) (3) (3) (3) (3) (3) (3) (3) (3) (3) (3) (3) (3) (8) (3) (3) (2) (1) et al., et 47 4 Other considerations et al. m depth and data mostly collected from important Whereas fishery-independent groundfish survey data Many of the juvenile fish surveys included in this Young Fish Survey data, as described in Rogers Data from the Continuous Plankton Recorder (CPR) may (CPR) Recorder Plankton Continuous the from Data With respect to the comparative importance of nursery The majority of fisheries-dependentnational fisheries laboratories data are held on held databases and in are available. Much of these data are also submitted to the the system DATRAS stored by ICES. other However, datasets collected by the fisheries laboratories (e.g. projects) and for stored electronically or research on paper may not be easily identified or accessed. are available for broad regions continental of shelf the surrounding the inner British and coastal areas and deep-water Isles, are more outer limited. data for report are conducted from July onwards, and so sample larger 0-groups. There are few published studies on distribution the patterns of recently settled (post-larval) fish, especially over the wider UK continental shelf, constitutes a and major this gap in our knowledge of the ecology of UK marine fish. Spawning grounds Cefas has undertaken or surveys participated for ichthyoplankton sampling, in and so many most data of were available. Some the data (e.g. for North Sea mackerel) were not available. Because some surveys only the target identify species, there are preserved samples that have been archived that could inform on other species. be able to provide useful information on the and distribution abundance of some species of fish larvae. Although a at towed is CPR The (e.g. limitations data some are there fixed 10 Nursery grounds (1998), were not used in this report as of they ongoing are analyses. These subject data could usefully inform on the distribution of selected 0-group fish in some inshore areas along the southern and eastern coasts of England. Comparable information for other parts of the UK coastal zone are lacking for recent years. A field programme could usefully be undertaken to better important identify inshore and grounds delineate along the western England and Wales. coasts of the of which identify to need important an is there grounds, nominal nursery grounds provide the greatest contribution to the stocks of adult fish. shipping lanes), these data may be a cost-effective way of examining the distribution of larvae in the waters of outer continental shelf and oceanic the waters, where there is an absence of dedicated ichthyoplankton surveys. et (1998) al. et based on based priori a et al. (1998) were Other Other considerations The spawning and nursery grounds proposed by Coull by proposed grounds nursery and spawning The With the exception of spawning mackerel, Norway pout With the exception of spawning mackerel, The project also identified a number of issues related The fish species used in the current project were a Additionally, many aspects of fish ecology and 4. In order to provide comprehensive maps over the entire UK shelf, the layers provided by Coull based on a range of evidence, including local observations, individual local surveys, reports from published and interpretation of broad scale field data. The absence of material the supporting data in this report has largely been rectified in the present project. al. (1998) were polygons and, the authors highlighted that “these maps should be seen as representing the widest known distribution given current knowledge and not be should seen as rigid, unchanging descriptions of presence (including users end the however, practice, In ”. absence or consultants and regulators) have often wrongly well-defined and exact be to polygons these of borders the inferred boundaries. For the present project we have identified the broad scale distributions of spawning and nursery grounds the that infer to likely more is user the that so rectangle, by boundaries are not fixed, and that the limits of spawning grounds can be variable from year to year (and season to season). Coull by described grounds the whiting, blue and did not attempt to highlight parts of nursery or spawning Where importance. ‘greater’ of considered were that areas some highlight to attempted have we permitted, have data of the more important grounds. For some areas, notably the Irish Sea, there has been effort, a and so high these data density may of also be survey viewed on scale a in finer relation to site-specific issues in this area. high-resolution Such data (particularly for ichthyoplankton) are not available for other parts of the UK continental shelf. to the collection, quality assurance, collation, access and collation the with issues The data. of dissemination onward of data from fisheries surveys highlighted issues, as discussed in Section 2. a number of identified been had that species of subset nature conservation listings. Hence, several commercially- important fish species, for example bass were not and included. A haddock, more targeted approach to spatial from benefit identify would stocks and/or species fish which management is required. demography, including spatial distributions, the states of constituent stocks, migration routes etc., can change over time. If biological data layers are to management be in used for the spatial future, then process whereby data layers are updated periodically. there needs to be a 48 4 Other considerations important seasonal differences in spatial distribution. spatial in differences seasonal important have that species those for limited are data Hence, year. per once conducted been only have surveys national many utilization. habitat and movements would provide more specific details of their offshore distributions, This undertaken. be usefully could species these of some for tags storage data electronic of use the report, this in included species wide-ranging and migratory more management. and conservation their especially if spatial management is deemed appropriate for needed, are waters UK in habitats important ecologically their and species fish rarest the of distributions current the data such of sources. utility the limits sampling appropriate and coordinated of absence the species, these of some from of sources some are there Although surveys. fishery-independent existing planning in effectively sampled not are spatial the in interest of an is many there which However, for species stocks. demersal suitable main most the are for data survey groundfish species, fish given any of distribution the to relation in stations survey in Marine Protected Areas (Marine Conservation Zones). Conservation (Marine Areas Protected Marine in considered be to species of areas nursery and spawning an edited version of the final report of this project, is and MB5301 project Defra by supported was work This Acknowledgments esnl nomto i lmtd n, n eet years, recent in and, limited is information Seasonal the of some for available data some are there Although delineate accurately more to surveys field Dedicated of distribution the and catchability of issues the Given data hoc ad Mapping 49 5 References pp.

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YoungSurvey Fish Surveys Egg Mackerel Horse and Mackerel on Group Working TrawlBeam on Surveys Group Working System VesselMonitoring Operator’sOffshore Association UK Fishermen’sScottish Association trawl headline high Portuguese Atlantic North-East the of Environment Marine the of Protection the for Conventions) Paris and Oslo OSPAR(formerly Convention 1.853 to (equivalent mile Nautical Organisations Fishermen’sof Federation National Surveys Production Egg Sea Irish Survey Larval Herring International Nomenclature Zoological on Commission International Seas the of Exploration the for Council International Group TrawlBottom Working International Survey TrawlBottom International Survey trawl) headline high (a vertical ouverture Grande survey Groundfish System Information Geographic Scotland) Marine (now Services Research Fisheries Zone Economic Exclusive Affairs Rural and Food Environment for Department ICES) DATabaseby (managed TRAwlSurveys of Recorder Plankton Continuous Science Aquaculture and Fisheries Environment, for Centre survey trawl Beam km) 55 Appendix B ESRI shape file metadata cm cm cm cm cm cm cm cm cm cm cm cm cm cm cm cm cm cm Trachurus trachurus, HOM, < Trachurus 8 , SKT, < Dipturus 43 batis, SKT, Merluccius merluccius, HKE, < 20 Raja clavata, THR, < 18 cm Raja undulata, UNR, <29 Micromesistius poutassou, WHB, < Raja montagui, SDR, < 18 cm Galeorhinus galeus, GAG, < 70 Lophius piscatorius, MON, < 28 Scomber scombrus, MAC, <24 Squalus acanthias, DGS, < 48 Merlangius merlangus, WHG, < 20 Clupea harengus, HER, <17.5 Pleuronectes platessa, PLE, <12 cm Solea solea, SOL, < 13 Molva molva, LIN, < 49 Gadus morhua, COD, < 23 Metadata Metadata associated with the data for supporting the identification of nominal spawning grounds. 19 species coding list below. 50 and 70 must be used. and < 14 less than or equal to 14 equal to 14 Thornback ray, Thornback ray, Spotted ray, Undulate ray, Spurdog, Ling, Anglerfish, Blue whiting, shark, Tope Where; XXX is the three letter species identifier outlined in the to get presence for 50% of tows for each station both Plaice, Mackerel, Sole, Horse mackerel, Whiting, Common skate, Cod, European hake, Sandeels, Ammodytidae, SAN, < 13 Herring, HOM_Mx_14 Maximum abundance of Horse mackerel which are HOM_P_14 Presence of Horse Mackerel which are less than or Species coding list: (b) Abstract: A points layer of egg-surveys between conducted in 1990 UK & waters 2008 used grounds of selected UK fish. to identify the spawning Purpose: provide the evidence-base behind the revised spawning To grounds of selected UK fish. cm were cm cm long Metadata Metadata associated with the data for supporting the identification of nominal nursery grounds. less than or equal to 8 equal to 8 station headline trawl, GOV = Grande Ouverture Verticale trawl, BT = Beam trawl Scotland, NI = Northern Ireland HOM_M_8 Maximum abundance of Horse Mackerel which are HOM_P_8 Presence of Horse Mackerel which are less than or XXX_Mx – Maximum abundance recorded at each XXX_P – Presence – Value -9999 = no data, 1 = presence in 1 or more tows at each station 50 = present in over 50% of tows at each station 70 = present in over 70% of tows at each station FID, OBJECTID, Shape, MedianLat, MedianLon Gear – gear type used: PHHT = Portuguese high fldCountry – – Value ENG = England and Wales, SCO = Appendix Appendix B: ESRI shape metadata file The study undertook 2 splits analyses for horse with mackerel, different because fish length so <= ubiquitous. For 14 the other species the length splits are shown below. It should be noted that various forms in of the GOV various surveys, are and used so relative abundance should only be interpreted within a survey region and not across the entire study area. Northern Ireland rockhopper trawl. use a non GOV Abstract: A points layer of presence juvenile and abundance of species selected research surveys in UK waters. conducted by fishery-independent Purpose: To provide the evidence-base behind the identification of nursery grounds for selected highly mobile species. Supplemental_Information: Point layer identification fields: Field descriptions: (a) 56 Appendix B ESRI shape file metadata Species coding list: coding Species Supplemental_Information: hne) n wr cnutd n 95 1 cuss 1024 cruises, (13 1995 in conducted were and Channel) George’s St of parts southern the from limited were data (although Sea Irish the in conducted were cruises of series with larvae. crab edible and species fish other for collected bass, data and sole sample to primarily Bight, Southern and Channel English the in conducted stations) valid (304 (1993). Horwood see information further For larvae. crab edible and species fish other for data collected but bass, in Channel Bristol and sole on inform to primarily conducted was which 1990 the in conducted stations) valid (524 following the from surveys: collated were data Ichthyoplankton Series: PLE. and SOL COD, for Sea Irish the in undertaken been have surveys intensive More Blue whiting, Blue Anglerfish, LIN molva, Molva Ling, mackerel, Horse Sole, Mackerel, Plaice, hake, European Cod, Herring, SAX Ammodytidae, Sandeels, Whiting, ISEPS: Irish Sea Egg Production Surveys. This extensive 1991 in cruises four of series A survey: Sole 1991 Bristol Channel plankton surveys: A series of five cruises , COD morhua, Gadus , SOL solea, Solea , PLE platessa, Pleuronectes , HER harengus, Clupea , WHG merlangus, Merlangius , MAC scombrus, Scomber , MON piscatorius, Lophius , WHB poutassou, Micromesistius , HKE merluccius, Merluccius TrachurusHOM trachurus, For further information see ICES (2009b). ICES see information further For included. were eggs 1 stage for and (2007) most survey recent the from data only survey, this For species. main target the are mackerel horse the and Mackerel of Isles. seaboard British western the along sampling extensive includes that survey coordinated internationally An survey: (2009a). ICES see information further For unavailable. Only Sea. were data North 2009 as used, were stations) (481 2008 for data the in grounds spawning herring main the over samples that survey coordinated internationally Fox see late from period month four December 2003 to early April 2004. For further information a over stations) (927 Sea North the of area large a covering institutes international Fox see late from period month four December 2003 to early April 2004. For further information a over stations) (927 Sea North the of area large a covering institutes international Bunn see information further For stations). 345 cruises, (5 2003 and stations) 158 cruises, (4 2002 stations), 227 cruises, (5 Sea, primarily to target plaice, that were conducted in 2001 Irish eastern the in undertaken were that cruises of series (1993). al. Nichols by described also cruises earlier with (2004), al. For further information see Fox see information further For (2004). Fox & Bunn see information further For stations). valid 486 cruises, (5 2008 and stations) valid 397 cruises, (5 2006 stations), valid 804 cruises, (8 2000 stations), valid MAC_HOM (stage 1 only): Triennial mackerel egg egg mackerel Triennial only): 1 (stage MAC_HOM An Survey: Larval Herring International IHLS: several by cruises of series A Sea_Eggs: North several by cruises of series A Sea_Larvae: North A surveys: plaice Sea Irish Eastern Sea: Irish Eastern , (2004). al., et (2005) and Taylorand (2005) al. et Taylorand (2005) al. et (2007). al. et (2007). al. et (1997) and Bunn and (1997) al. et et et

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