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.