Reproduced with the permission of the Scottish Naturalist
                              Copyright: 
                              May be used for private research. All other rights 
                              reserved 
                            
                             
                             
                            By ADRIAN J. SHINE
                            Loch Ness and Morar Project
                             
                            JAN KUBECKA
                            Hydroacoustic Unit, Department of Biology,
                            Royal Holloway University of London
                             
                            DAVID S. MARTIN
                            Loch Ness and Morar Project
                             
                            ANNIE DUNCAN
                            Hydroacoustic Unit, Department of Biology,
                            Royal Holloway University of London
                             
                            Introduction
                              
                              
                            Obtaining 
                              reliable ecological data on the fish species living 
                              in a large water body the size of Loch Ness is no 
                              easy task, and depends upon employing a variety 
                              of sampling methods. Apart from sampling difficulties associated 
                              with the size and depth of this water body, live 
                              capture by means of seine-nets, trawl-nets and tow-nets 
                              exerts a simpler mechanical selectivity within the 
                              area of collection than do passive techniques using 
                              gill-nets or fyke-nets, with which more complex 
                              behavioural selectivity plays a bigger role.
                              
                            The above conventional 
                              fish capture techniques were used in Loch Ness from 
                              1988 onwards at sites indicated in Figure 
                              1 (5K), and each method was more appropriate 
                              for some fish habitats available in Loch Ness than 
                              for others (Figure 
                              2a, 28K and Figure 
                              2b, 18K), thus introducing another level 
                              of selectivity. To capture fish in the profundal zone, unconventional 
                              methods had to be adopted, such as setting gill-nets 
                              at 200 m depth with their lead-line resting on the 
                              bottom sediment. The aim of this paper is to add the more 
                              recent results, from a greater variety of capture 
                              methods, to the fish habitat picture published by 
                              Shine and Martin (1988).
                            
                            
                            Vol 105, The 
                              Scottish Naturalist: Fish Habitats in Loch 
                              Ness. p239 
                            Methods
                            Pelagial 
                              Habitat
                            The Loch Ness and 
                              Morar Project (L.N.M.P.) gill-net series used to 
                              sample the pelagial habitat consisted of five nets 
                              between 20 and 25 m long, 1.25 m deep, and with 
                              mesh bar sizes of 20 mm, 25 mm and 30 mm, giving 
                              a total length of 115m and a fishing area of 1432. These were suspended on the L.N.M.P. mooring 
                              rope in the North Basin opposite Brachla                              (see Figure 
                              1),  in mid-loch between 20 and 30 m depth, 
                              in order to span the scattering layer (Shine and 
                              Martin, 1988). During the July-August 1991 series, the nets 
                              were exposed on fifteen dates, during daylight hours, 
                            at night or over 24 hours.
                            
                             
                            In October 1991 
                              and May 1992, six gill-nets (30 m x 6.0 m; with 
                              15.5 mm, 25 mm and 30 mm mesh sizes) belonging to 
                              Royal Holloway University of London (R.H.U.L.) were 
                              suspended on the same L.N.M.P. mooring rope, at 
                              depths between 20 and 30 m, to encompass the scattering 
                              layer which had been detected acoustically This net series was 180 m long, and had a fishing 
                              area of 1,080 m2. In October 1991, gale force winds prevented 
                              the collection of the nets after 24 hours and they 
                              were left for four days. The same nets were left on two occasions 
                              for 24 hours in May 1992, when the weather was calmer. Also during May 1992, an ichthyoplankton 
                              tow-net of 1.0 m diameter and with a 1.0 x 1.7 mm 
                              mesh size was towed at three knots within the scattering 
                            layer at about 24 m depth for two hours.
                            
                             
                            On two occasions, 
                              in August 1988 and November 1992, the Department 
                              of Agriculture and Fisheries for Scotland's (D.A.F.S.) 
                              vessel operated a trawl-net in the South Basin at 
                              mid-water depths down to 30 m. The August sample was collected using a sprat-trawl 
                              with a 12 mm mesh cod-end and a 6.0 m mouth opening 
                              (four hauls totalling four hours of trawling). The November sample was collected in the 
                              fine-meshed cod-end during trawling trials between 
                            Fort Augustus and Urquhart Bay.
                             
                            Sublittoral 
                              Habitat
                            During December 
                              1992, the L.N.M.P. gill-nets together with some 
                              Greer gill-nets (bar sizes 19-30 mm and area 121 
                              m2, giving a total area of 264 m2) 
                              were set to sample sublittoral depths of 15-25 m 
                              off Brachla. The 
                              nets were set parallel to the shore with their lead-line 
                              resting on the bottom sediment. The same technique
                            Vol 105, The 
                              Scottish Naturalist: Fish Habitats in Loch 
                              Ness. p242
                            
 
                            
                             
                            Vol 105, The 
                              Scottish Naturalist: Fish Habitats in Loch 
                              Ness p243 
                            was used to collect 
                              the fish as in the Table headed 'Littoral Netting 
                              1984' in Shine and Martin (1988: 148). On a few occasions, a gill-net (0.5 m high 
                              and 200 m long, with a bar size of 15 mm) was deployed 
                              down the loch walls from depths of 30 m to 200 m 
                              in order to sample any fish present.
                            
                            Profundal 
                              Habitat
                            The same series 
                              of gill-nets were used to sample the fish inhabiting 
                              the profundal at 200 m depth, both in Shine and 
                              Martin (1988) and in subsequent sampling. The nets were set with their lead-line resting 
                              on the bottom sediment. Occasionally, additional gill-nets or fyke-nets 
                              were added (Figure 
                              2b). More 
                              nets were deployed during the five dates in 1991 
                              (396 m2) than during the seven dates 
                              in 1988 (35-135 m2), which gave a better 
                              success rate and fewer occasions with zero catches.
                            
                            For comparative 
                              purposes a gill-net series, consisting of both the 
                              Greer and the 0.5 m high and 200 m long nets, combined 
                              with a series of fyke-nets, were set in Loch Morar 
                              in August 1992 for two days at a depth of 275 m, 
                              but with no fish caught.
                            Littoral 
                              Habitat
                            During May-June 
                              1992, another technique was employed to sample littoral 
                              fish during the day and night. An R.H.U.L. beach seine-net, 15 m long and 
                              2.5 m deep with 1.0 cm mesh size, was laid parallel 
                              to the shore-line and about 15-20 m out into the 
                              water. After setting the net, the two ends of the net were pulled gently, 
                              symmetrically and quietly by rope. Particular attention was paid to the lead-line 
                              in order that it be pulled close to the bottom. The area sampled per haul was calculated 
                              according to Kubecka and Bohm (1991) to be 80 m2 
                              for beaches near Brachla (29th/30th May 1992) and 
                              117 m2 at Dores Beach (6th/7th June 1992). In all, fourteen hauls were taken and all 
                              272 fish caught were released within fifteen minutes 
                              of capture, after measurement and removal of scales 
                            for ageing.
                            
                            Results 
                              and Discussion
                            The same seven 
                              species of fish were recorded for Loch Ness during 
                              this period of sampling as were recorded by Shine 
                              and Martin (1988). This is the same number as given by Maitland 
                              and his colleagues (Maitland, 1981), who included 
                              Pike Esox lucius, but not Minnows Phoxinus phoxinus 
                              which were first recorded
                            Vol 105, The 
                              Scottish Naturalist: Fish Habitats in Loch 
                              Ness p244 
                             for Loch Ness 
                              in 1985-87 (Shine and Martin, 1988). Pike were not captured during the period 
                              1988-92, although some were caught in the sublittoral 
                              in previous years. Maitland (1981) also identified his lamprey 
                              as the Brook Lamprey Lampetra planeri.
                            
                            Pelagial 
                              Habitat
                            The numbers and 
                              sizes of fish species caught by the various collecting 
                              methods are given in Table 1. During fifteen deployments between July and 
                              September 1991, the L.N.M.P. gill-nets suspended 
                              at about 20-30 m depth caught 33 Charr Salvelinus 
                              alpinus and a few Brown Trout Salmo trutta. There were two size classes of Charr, with 
                              modal lengths of 16 cm and 26-27 cm. All were adults, with 24 females and ten males. The average catch per unit effort was 2.0 fish 
                              per 24-hour exposure.
                            
                            In October, when 
                              the R.H.U.L. gill-netting (1,080 m2) was left fishing 
                              between 20 and 30 m depth for four days, there was 
                              a large catch of 17 Charr, ten of which were juveniles 
                              (<12 cm fork length), and 23 adult Brown Trout 
                              up to 2.0 kg in weight, ten of which were Ferox 
                              Trout. It 
                              seemed likely that the Trout were attracted to the 
                              captive Charr, since some Trout were trapped within 
                              inches of the Charr, some of which had been chewed. Fewer fish, and only Charr of a similar size 
                              range, were caught during a 24-hour exposure of 
                              the same nets at the same site in May 1992. Most of these Charr were adults, and some 
                              were mature - one female with large eggs in the 
                              ovary and one male with large testes.
                            
                            Both fish catches 
                              by the D.A.F.S. vessel using a mid-water trawl in 
                              the South Basin are intriguing on two counts. Both the August 1988 and the November 1992 
                              trawling collected both Three-spined Sticklebacks 
                              Gasterosteus aculeatus and very small Charr from 
                              the full pelagial in waters down to 30 m depth. The collection of as many as 50 Sticklebacks 
                              in November 1992 is a surprising record, which should 
                              be investigated further because of its implications 
                              for acoustic studies (Kubecka, Duncan and Butterworth, 
                              1993). Both D.A.F.S. trawl hauls captured large numbers 
                              of Charr, some of which were the smallest Charr 
                              (lengths of 3.0, 4.5 and 5.0 cm) so far recorded 
                              from Loch Ness, or indeed, as far as we are aware, 
                              from the pelagic of any Scottish loch.
                            
                            Mr. R.B. Greer 
                              of D.A.F.S. identified these Charr as young but 
                              not larval fish, as might be expected if Loch Ness 
                              Charr spawn in the loch in late October/early November. Moreover, larval Charr were not collected 
                              in the plankton during
                            Vol 105, The 
                              Scottish Naturalist: Fish Habitats in Loch 
                              Ness p245 
                             autumn 1992 by 
                              tow-netting within the scattering layer, nor during 
                              May 1992, as might be expected if the Charr were 
                              spring spawners. Further study is urgently needed on the spawning 
                              sites and the life cycle of the Charr, which is 
                              probably the numerically dominant species in Loch 
                              Ness. A 
                              tow-net larger than 1.0 m diameter may be an advantage. The question as to whether Charr spawn on beaches 
                              in the loch or in local rivers also needs to be 
                              resolved. There are no known records, however, of Charr 
                            entering any feeder stream of Loch Ness.
                            
                            The capture of 
                              the larval Lamprey in May 1992 is a new pelagial 
                              record for Loch Ness, although there is also a recent 
                              profundal record (Martin, Shine and Duncan, 1993). It was possibly an individual drawn into the 
                            pelagial by wind-induced currents.
                            Sublittoral 
                              Habitat
                            The collection 
                              of mature Charr, whose gonads were 'running' ripe, 
                              during December 1992 by the L.N.M.P. gill-net series, 
                              set at sublittoral depths of 15-25 m off Brachla, 
                              adds some supportive evidence for autumn spawning 
                              in the loch, and may indicate the location of one 
                              possible spawning site. This was an unusual catch, since regular 
                              sampling of the sub-littoral depths between 10 and 
                              30 m with the same nets during May-June 1989 and 
                              March-September 1991 normally caught more Brown 
                            Trout than Charr.
                            Profundal 
                              Habitat
                            Sixteen more Charr 
                              were caught from the profundal regions on twelve 
                              sampling occasions during the 1988-92 period, thus 
                              giving an overall total of 29 profundal fish caught 
                              since 1982, with standard lengths ranging from 4.5 
                              to 27.9 cm. These were fish with profundal invertebrates 
                              in their guts. About a third of these fish were less than 
                              10 cm in length, and about 40% were greater than 
                              19 cm. The 
                              fish were caught either by benthic trawling or with 
                              an adequate area of gill-netting (about 400 m2) 
                            in deep water.
                            
                            Littoral 
                              Habitat
                            Night-seining is 
                              one of the most cost-effective sampling techniques, 
                              which exploits the behavioural tendency of many 
                              fish species to migrate shorewards during twilight 
                              (Kubecka, 1993). The technique thus samples both the littoral 
                              fish, which spend their days in shallow water, as 
                              well as the deep or open-water fish,
                            Vol 105, The 
                              Scottish Naturalist: Fish Habitats in Loch 
                              Ness p248
                            
 
                            
                             
                            Vol 105, The 
                              Scottish Naturalist: Fish Habitats in Loch 
                              Ness p249 
                            which move shorewards 
                              at dusk. It proved valuable in Loch Ness, where it was 
                              used for the first time in May-June 1992.
                            
                            Table 2 lists the 
                              species composition of fish caught. During the day, no fish were caught in four 
                              out of five hauls, probably because the well-lit 
                              shallow water is a dangerous place for small fish. Nevertheless, small fish, such as Minnows, 
                              Sticklebacks and even small salmonids, form quite 
                              dense shoals in very shallow sites, thus avoiding 
                              predation by big fish (see the selective day catch 
                              in Table 2).
                            
                            At night, Sticklebacks 
                              were caught in large numbers in every haul, together 
                              with small salmonids and Minnows in some hauls. The numbers of salmonids caught increased 
                              with 'true darkness', but declined abruply with 
                              the advent of 'dawn', which arrives early in the 
                              short northern night (Figure 
                              3, 4K). It would be advantageous to repeat this exercise 
                            when nights are longer.
                            
                            Although only 29 
                              Brown Trout were caught, Figure 
                              4 (6K) shows that three year classes of 
                              fish were present, namely 1+, 2+ and 3+. With a bigger net, it seems likely that night 
                              shore-seining could be used to establish the ratios 
                              of young Brown Trout to young Salmon Salmo salar 
                              to littoral Charr, which is useful information for 
                              understanding the ecology of Loch Ness salmonid 
                            populations.
Before 
                            embarking on a longer term study, some care should 
                            be taken with the size of net used. The number of fish caught is a function of 
                            the area sampled quantitatively, and Figure 
                            5 (5K) illustrates the relationship between 
                            net length and area sampled.                            
                            
                            Sticklebacks and 
                              Minnows are an important food source for piscivorous 
                              salmonids, so any information about their population 
                              size structure is useful. Figure 
                              6 (6K) shows that three size groups were present in the Stickleback population 
                              sampled by shore-seining; these were 31-33 mm, 41-45 
                              mm and 51 mm or larger. Although Sticklebacks cannot be aged by their 
                              scales, further study on their growth rates would 
                              provide this information.
                              
                            
                            No Pike were caught 
                              by shore-seining in Loch Ness, perhaps because Pike 
                              were absent in the sampled areas, whereas the catch 
                              per unit effort of small Eels Anguilla anguilla 
                              (30-35 cm) was high compared with other localities 
                              where night-seining has been carried out (e.g. River 
                            Thames, Czech and London reservoirs). Both species need further study in Loch Ness.
                            Vol 105, The 
                              Scottish Naturalist: Fish Habitats in Loch 
                            Ness p253
Summary 
                              of Fish Habitats in Loch Ness
      
                            Sampling Loch Ness 
                              fish habitats is a difficult task. Table 1 gives a summary of the species composition 
                              and sizes of the fish catches in the different habitats 
                              of the loch. Charr 
                              is the most ubiquitous species in the loch and the 
                              most abundant species in the pelagial and the profundal. Brown Trout dominate sublittoral catches 
                              with Charr coming second (see also Shine and Martin, 
                              1988), whilst the Three-spined Stickleback is an 
                              important species in the littoral and the pelagial 
                              (Figure 
                            7, 46K colour chart).
                            
                            The various fish 
                              habitats were sampled by appropriate methods, which 
                              are illustrated in Figure 
                              2a and Figure2b. At present, depths greater than 50 m in the 
                              open water have not been sampled by live capture 
                              techniques, and depths between 50-70 m and 200 m 
                              were not reached by 420 kHz dual-beam sonar (Kubecka, 
                              Duncan and Butterworth, 1993). Since then, however, a 38 kHz split-beam 
                              sonar has shown a small number of probable fish 
                              targets at these depths, and a greater number on 
                              the loch bed (Figure 
                              8, (19K colour chart). Having established a sampling technology 
                              for some of these fish habitats, and with the aid 
                              of quantitative echo-sounding (Kubecka, Duncan and 
                              Butterworth, 1993; Shine, Martin and Marjoram, 1993), 
                              the situation is ripe for studying how the more 
                              numerous species, Charr and Sticklebacks, exploit 
                              this deep oligotrophic loch, by finding out where 
                              they spawn, where their subsequent life-cycle stages 
                              live, and quantifying their impact upon the ecosystem.
                            
                            Acknowledgements
                            We should like 
                              to express our thanks to the Royal Holloway University 
                              of London for permission to use the fish sampling 
                              equipment, and to the Loch Ness Fisheries Board 
                              for permission to try out whether night shore-seining 
                              was a useful fish sampling method in one of Britain's 
                              largest lochs. We also acknowledge the presence and help 
                            of the bailiff, Mr. W. Hastings.
                             
                            References
                            Kubecka, J. (1993). Night 
                              inshore migration and capture of adult fish by shore 
                              seining. Aquaculture and Fisheries Management, 
                              24: in press.
                            
                            Kubecka, J. and 
                              Bohm, M. (1991). The fish fauna of the Jordan Reservoir, one 
                            of the oldest man-made lakes in central Europe. Journal of Fish Biology, 38: 935-950.
                            
                            
                            Vol 105, The 
                              Scottish Naturalist: Fish Habitats in Loch 
                              Ness p255
                            Kubecka, J., Duncan, 
                              A. and Butterworth, A.J. (1993). Large and small organisms detected in the 
                            open waters of Loch Ness by dual-beam acoustics. Scottish Naturalist, 105: 175-193.
                            
                            Maitland, P.S. (Ed.) (1981). The Ecology of Scotland's 
                              Largest Lochs: Lomond, Awe, Ness, Morar and Shiel. Monographiae Biologicae, Vol. 44. The Hague: Junk.
                            
                            Martin, D.S., Shine, 
                              A.J. and Duncan, A. (1993). The profundal fauna of Loch Ness and Loch Morar. Scottish Naturalist, 105: 113-136.
                            
                            Shine, A.J. and 
                              Martin, D.S. (1988). Loch Ness habitats observed by sonar and underwater 
                              television. Scottish 
                              Naturalist, 100: 111-199.
                            
                            Shine, A.J., Martin, 
                              D.S. and Marjoram, R.S. (1993). Spatial distribution and diurnal migration 
                              of the pelagic fish and zooplankton in Loch Ness. Scottish Naturalist, 105: 195-235.
                            
                              
                            Received July 
                              1993
                            Mr. Adrian J. 
                              Shine, Loch Ness and Morar Project,
                            Loch Ness Centre, 
                              DRUMNADROCHIT, Inverness-shire IV3 6TU.
                              
                              
                              Dr. 
                              Jan Kubecka, Hydrobiological Institute,
                            Czech Academy 
                              of Sciences, 370 05 Ceske Budejovice, Na 
                              Sadkach 7, CZECH Republic.
                            Present address: 
                              Hydroacoustic Unit, Department of Biology, 
                              Royal 
                              Holloway
                              University of London, EGHAM, Surrey TW20 0EX.
                            
                              Mr. David S. Martin, Loch Ness and Morar Project,
                            Loch Ness Centre, 
                              DRUMNADROCHIT, Inverness-shire IV3 6TU.
                            
                              Dr. 
                              Annie Duncan, Hydroacoustic Unit, Department of 
                              Biology,
                            Royal Holloway 
                              University of London, EGHAM, Surrey TW20 0EX.