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
JOHN MINSHULL
Loch Ness
and Morar Project
and MARALYN SHINE
Loch Ness
and Morar Project
Pioneer
Work
It is now
some ninety years since Sir John Murray
and Laurence Pullar published the first
part of their pioneer hydrographic survey
of Loch Ness (Murray and Pullar, 1903-08),
followed by their comprehensive account
of all the important Scottish fresh-water
lochs (Murray and Pullar, 1910). Then E.R.
Watson and James Wedderburn's discovery
of internal seiches in Loch Ness (Watson,
1904; Wedderburn, 1907a and 1907b; Wedderburn
and Watson, 1909) began a long tradition
of physical limnology (e.g. Wedderburn,
1907b, 1911 and 1912) which was ultimately
to attract other leading scientists, such
as C.H. Mortimer (Mortimer, 1952 and 1955)
and S.A. Thorpe (Thorpe, Hall and Crofts,
1972; Thorpe, 1974 and 1988), to this fascinating
body of water.
Nevertheless, apart from the pioneer work of the Bathymetrical Survey, the biology of the loch remained
rather neglected until Dr. Peter Maitland's
multi-disciplinary survey of 1977-80 (Maitland,
1981), which has remained the definitive
background for further studies.
1993
This year (1993)
also marks two other anniversaries. It
is now 125 years since the
Inverness Courier (8th October
1868) first referred to the loch's unusual
tradition, and sixty years since the same
newspaper (Anon., 1933) coined the term
'Monster'. This tradition has been blamed,
possibly somewhat unjustly, for scientific
reticence regarding the loch. In fact,
as the roll of distinguished scientists
shows, this is hardly the case, but equally
it remains to be explained why Britain's
premier body of fresh-water has not received
more attention. Perhaps one reason has
been the inadequacy of much freshwater sampling
equipment when confronted with windy surface
conditions and water depths of over 200
metres.
The
Monster Tradition
On
the other hand, although the Monster tradition
may have attracted enthusiastic amateur
naturalists but few professional zoologists,
it did attract trained engineers, and with
them some quite large-scale expeditions
to provide the logistics for the novel methods
of protracted acoustic and underwater camera
monitoring (e.g. Campbell and Solomon, 1972;
Mackal, 1976; Rines, Wyckoff, Edgerton and
Klein, 1976; Rines, 1988). It was from
here that the Loch Ness and Morar Project's
tradition of improvisation developed, and,
as the general scientific possibilities
became clear, so some marine equipment found
its first freshwater use in Loch Ness (Note
1).
Finally,
the opportunity afforded by the Caledonian
Canal was exploited, as a collaboration
with the Simrad Company and the Department
of Agriculture and Fisheries for Scotland
(Shine and Martin, 1988); this saw larger
vessels entering the loch, thus partly off-setting
the hostile surface conditions. None of
this would have been possible without a
considerable change in general scientific
outlook and perspective.
The
Parting of the Ways
In 1960 one of the earliest investigators, Dr. Peter
Baker, was already attempting to assess
the fish population by acoustics, and was
speculating about the effects of the loch
physics upon the horizontal transport of
biomass (Baker and Westwood, 1960; Baker,
1962). However, events in the 1970s, leading
to the naming of the Monster as Nessiteras
rhombopteryx (Scott and Rines, 1975),
severed any real connection between aquatic
science and popular expectations (Witchell,
1975: 147-156; Witchell, 1989: 139-146).
The
Project in the 1980s
The activities of the Loch Ness and Morar Project in
the 1980s sought to revise these expectations,
through the reintroduction of general scientific
objectives, and in 1987 analysis of unusual
sonar contacts culminated in 'Operation
Deepscan' (Shine and Martin, 1988) and the
slaying of the popular media Monster.
Since
then, the Project has been given a headquarters
at the loch-side by the Official Loch Ness
Exhibition Centre, whose proprietor Mr.
R.A. Bremner has also generously provided
a laboratory, harbour facilities, and land
for a field station. This permanent presence
has permitted the Project to invite and
encourage trained scientific workers from
all limnological disciplines to collaborate
and take advantage of our field-work and
equipment specially developed for deep-water
research.
Ecos
A
34-ft clinker-built motor cruiser was purchased
in the autumn of 1989. It was originally
built in the 1930s as a mail boat for the
Orkneys. Renamed Ecos, it was renovated and specially
adapted for freshwater research (Figure
1, 11K). The vessel was selected
as particularly suitable because of the
open cockpit close to the waterline for
easy recovery of equipment and samples. It also has a flush deck giving the necessary
space for the hand-hauling of over 100 km
of ropes during sampling, trawling, dredging
and coring over the last three years (Figure
2, 8K). Internally, bench space
was provided to mount the bulky instrumentation
necessary, for example, for acoustic fish
stock surveys. Externally, an out-rigger
system permits the rapid over-side deployment
of heavy transducers, to port and starboard.
Fixed Station
A
fixed station facility was constructed in
mid-loch, consisting of a submerged two-point
mooring laid in 200 m depth of water. This
involves 1.0 km of warp, and permits Ecos to
maintain station without drift while accurate
series of vertical samples are taken.
Over
14 tonnes of water have been raised from
the station during continuous studies over
the past three summers and two winters. The mooring has also enabled instrumentation,
such as thermistors and sediment traps,
to be maintained and serviced in deep water.
Coring Equipment
The
Project has designed and built its own gravity-coring
apparatus for deep-water and for use from
smaller vessels. The wide bore (10.3 cm)
was originally developed to collect the
large volumes of sediment necessary to detect
chemicals present at minute concentrations
(Sanders, Jones and Shine, 1993). However,
the system has proved very effective for
quantitative benthos studies (Martin, Shine
and Duncan, 1993; Griffiths and Martin,
1993) as well as sediment mapping (Bennett
and Shine, 1993). Very satisfactory 3.0
m cores can be taken with this system.
Personnel
The
authors provide a permanent loch-side presence
to conduct field-work. In London, David
Martin acts as a scientific clearing house
for materials destined for specialist workers
elsewhere. Each year, volunteers are recruited
and logistics organised to support the various
programmes. These volunteers are mainly
students, some in the course of B.Sc. dissertations,
but other volunteers come from all walks
of life.
Recent
Increase in Research
During
the past three years it is gratifying to
report that forty collaborators have responded
with research on many aspects of Loch Ness,
including a continuous three-year multi-disciplinary
plankton study by the University of Lancaster,
to which the Project has contributed the
field-work (see Appendix). At least seventeen
B.Sc., M.Sc., M.Phil. and Ph.D. dissertations
and theses, wholly or partly devoted to
the loch, have been completed or are in
preparation (see Appendix), and the Project's
Sediment Group is also being co-ordinated
by the Environmental Change Research Centre
at University College London.
During
1992 and 1993 a certain amount of material
was presented at meetings of the Institute
of Fisheries Management (Note 2), the Societas
Internationalis Limnologae Theoreticae et
Applicatatae (S.I.L.S.) in Barcelona (Note
3), and the British Ecological Society (Note
4); further papers and posters featured
in the 50th meetings of both the Scottish
(Note 5) and London (Note 6) Freshwater
Groups. Additional specialist publications
will follow in due course, but clearly it
is now time to place some collected observations
on record in order to emphasise some aspects
of recent work which are regarded as particularly
interesting, and to pose a few questions.
Project
Contributions
The
place of Loch Ness in relation to the other
major Scottish lochs has already been set
out in detail by Maitland (1981). In terms
of richness and diversity, Loch Ness was
found to be generally intermediate between
the 'richest', Loch Awe and Loch Lomond,
and the 'poorest', Loch Morar and Loch Shiel. In the main, the Loch Ness and Morar Project
has applied its energies to carrying out
work complementary to several aspects of
Dr. Maitland's studies.
Sediments
Following
a hydrographic and seismic survey (Young
and Shine, 1993), sediment studies have
been conducted in detail and have produced
answers to the questions of acidification,
eutrophication and pollution (Jones, Rose
and Appleby - Notes 4 and 5; Bennett and
Shine, 1993; Jenkins, 1993a; Sanders, Jones
and Shine, 1993). Much additional work
is being undertaken as longer cores are
obtained.
The
Littoral
The
littoral of Loch Ness is a steeply sloping
stony ribbon exposed to considerable wave
action. Examination of the sub-littoral
macrophytes, by underwater television, confirms
the low light penetration, the Isoetes
swards of more transparent lochs, such as
Morar, being replaced by strands of filamentous
algae. The chemistry of feeder streams
has been examined by Jenkins (1993b).
The
invertebrates of the littoral (<1.0 m)
were extensively surveyed by Smith, Maitland,
Young and Carr (1981). The Project's attentions
have therefore been restricted to sub-littoral
invertebrate sampling, for comparison with
the profundal.
The fish of the littoral are described
by Shine, Kubecka, Martin and Duncan (1993).
The
Profundal
In
terms of basic species lists, the only significant
gap in the Institute of Terrestrial Ecology's
comprehensive survey (Maitland, 1981) was
the profundal region, the size of which
is very considerable in Loch Ness because
of the loch's great mean depth (132 m). The Project's contribution (Martin, Shine
and Duncan, 1993) now fills this gap.
Ongoing
work with a quantitative wide-bore coring
technique is now defining community structures
and densities in a way similar to that achieved
for the ostracods (Griffiths and Martin,
1993). In the future, the life history
of some of the chironomids seems worth examination,
since they contribute to the diet of profundal
Charr as larvae and to that of pelagic Charr
as pupae. It is particularly important
to follow the pattern of their emergence
and behaviour in the pelagic at this time,
when they are sometimes associated with
acoustic scattering layers.
An
interesting discovery during the profundal
programme was a population of Charr ranging
in size between 4.0 cm and 30 cm. These
are described by Shine, Kubecka, Martin
and Duncan (1993).
The
Pelagic
The
pelagic zone has been the main area for
the Loch Ness and Morar Project's contributions
to Loch Ness research, because of its amenability
to investigation by acoustic methods (Kubecka,
Duncan and Butterworth, 1993; Shine, Martin
and Marjoram, 1993).
Physical Characteristics of Loch Ness
The
regular morphometry and orientation of Loch
Ness, in line with the prevailing south-west
wind, has produced a dynamic physical environment
(Shine and Martin, 1988). Powerful wind
mixing often extends deep into the water
column. This, together with low light penetration
and scarce nutrients, reduces the scope
of primary productivity, and the importance
of microbial productivity is therefore enhanced
(Shine, Martin, Bennett and Marjoram, 1993).
Investigations
The
balance between algae and bacteria, together
with nutrient and carbon flow, should soon
become clear as the Project enters the last
of three years of sampling on behalf of
the University of Lancaster's plankton survey,
funded by the Natural Environment Research
Council (N.E.R.C.) (see Appendix).
In
addition to the above acoustic studies,
the pelagic fish are described by Shine,
Kubecka, Martin and Duncan (1993) and Martin
and Shine (1993).
Further
efforts will be made to explore links between
vertical migration and predator-prey relationships
in the fish and zooplankton. The components
of the scattering layer also need to be
further resolved, and the behaviour of chironomid
pupae examined.
Acknowledgements
The
work described in this paper was made possible
by the enthusiasm of the volunteer members
of the Loch Ness and Morar Project over
the last three years. Particular thanks
are due to the following, who have given
considerable time in staff roles during
the field-work: Miss S. Bennett, Mrs E.
Gallagher, Mr. K. Hawley, Mr. B. Herring,
Miss R.S. Marjoram, Mr. James Reid, Mr.
P. Rimmer, Mr. J. Vallette, and Mr. K. Wilson.
Notes
Note 1
The
200 m depth of the loch has been exploited
in trials of a variety of marine equipment,
including Remote Operated Vehicles (R.O.V.s). Conversely, water of this depth demands
equipment with marine capabilities, such
as underwater television and cameras; marine
sonars have also been brought in for hydrography
and acoustic fish estimates. It has also
been possible to use large hull-mounted
units, as vessels are able to enter the
loch via the Caledonian Canal.
Note 2
Institute
of Fisheries Management, Greater London
and South-East Branch. Branch Meeting,
King's College, London, 3rd March 1992.
Paper
read. The Fish and Other Fauna of Loch
Ness. By Annie Duncan and J. Kubecka (Department
of Biology, Royal Holloway University of
London) and D.S. Martin and A.J. Shine (Loch
Ness and Morar Project).
Note 3
Societas
Internationalis Limnologae Theoreticae et
Applicatatae. XXV International Congress,
University of Barcelona, Spain, 21st-27th
August 1992.
Poster
paper. Longitudinal and Vertical Patterns
of Pelagic Fish Distribution in Loch Ness:
Acoustic Sizes and Numbers. By Annie Duncan
and J. Kubecka (Department of Biology, Royal
Holloway University of London) and A.J.
Butterworth (National Rivers Authority,
Thames Region).
Poster
papers. Studies of the Plankton of Loch
Ness. By R.I. Jones, Johanna Laybourn-Parry,
M. Walton and Judith M. Young (Institute
of Environmental and Biological Sciences,
University of Lancaster), and A.E. Bailey-Watts
(Institute of Freshwater Ecology, Penicuick,
Midlothian).
1)
Phytoplankton. By Judith M. Young, R.I.
Jones, and A.E. Bailey-Watts.
2)
The Microbial Loop. By Johanna Laybourn-Parry
and M. Walton.
3)
Rotifers. By Alison Fulcher.
Later published as abstracts: 1 - Young, Jones and Bailey-Watts
(1993); 2 - Laybourn-Parry and Walton (1993);
3 - Fulcher (1993).
Note 4
British
Ecological Society. Winter meeting and
A.G.M., University of Lancaster, 15th-17th
December 1992.
Paper
read. Functional Aspects of the Microbial
Plankton in Loch Ness. By M. Walton and
Johanna Laybourn-Parry (Institute of Environmental
and Biological Sciences, University of Lancester).
Paper
read. Bioassay Studies of Loch Ness Phytoplankton. By R.I. Jones and Anne Hartley (Institute
of Environmental and Biological Sciences,
University of Lancaster).
Paper
read. Dual-beam Echo-sounding for Fish
in Loch Ness. By Annie Duncan and J. Kubecka
(Department of Biology, Royal Holloway University
of London).
Paper
read. Picophytoplankton of Loch Ness, a
Deep Oligotrophic Lake. By Judith M. Young
and R.I. Jones (Institute of Environmental
and Biological Sciences, University of Lancaster.
Institute of Environmental and Biological Sciences, University
of Lancaster.
Poster papers: Studies of the Plankton of Loch Ness.
1)
Phytoplankton. By Judith M. Young and R.I.
Jones.
2)
The Microbial Loop. By Johanna Laybourn-Parry
and M. Walton.
3)
Rotifers. By Alison Fulcher.
4)
Crustacean Zooplankton. By U. Jayakodi.
Later published as abstracts: 1 - Young, Jones and Bailey-Watts
(1993); 2 - Laybourn-Parry and Walton (1993);
3 - Fulcher (1993).
Poster
paper. The Recent History of Loch Ness. By Vivienne Jones and N. Rose (Environmental
Change Research Centre, University College
London) and P.G. Appleby (University of
Liverpool).
Poster
paper. Longitudinal and Vertical Patterns
of Pelagic Fish Distribution in Loch Ness:
Acoustic sizes and Numbers. By Annie Duncan
and J. Kubecka (Department of Biology, Royal
Holloway University of London) and A.J.
Butterworth (National Rivers Authority,
Thames Region).
Poster
paper. Profundal Benthos in Loch Ness. By D.S. Martin and A.J. Shine (Loch Ness
and Morar Project) and Annie Duncan (Department
of Biology, Royal Holloway University of
London).
Poster
paper. Loch Ness Project, Reviews and Previews. By A.J. Shine (Loch Ness and Morar Project).
Poster
paper. Loch Ness Undercurrents 1 and 2. By A.J. Shine (Loch Ness and Morar Project).
Note 5
Scottish
Freshwater Group. 50th Meeting, University
of Stirling, 2nd-3rd February 1993.
Institute of Environmental and Biological Sciences, University
of Lancaster.
Poster
papers: Studies of the Plankton of Loch
Ness.
1)
Phytoplankton. By Judith M. Young and R.I.
Jones.
2)
The Microbial Loop. By Johanna Laybourn-Parry
and M. Walton.
3)
Rotifers. By Alison Fulcher.
4)
Crustacean Zooplankton. By U. Jayakodi.
Later published as abstracts: 1 - Young, Jones and Bailey-Watts
(1993); 2 - Laybourn-Parry and Walton (1993);
3 - Fulcher (1993).
Poster
paper. The Recent History of Loch Ness. By Vivienne Jones and N. Rose (Environmental
Change Research Centre, University College
London) and P.G. Appleby (University of
Liverpool).
Poster
paper. Application of Hydroacoustics to
Scottish Freshwaters: Case Studies in Loch
Ness and Orkney Lochs. By Annie Duncan
and J. Kubecka (Department of Biology, Royal
Holloway University of London) and A.J.
Butterworth and W. Duncan (National Rivers
Authority, Thames Region).
Poster
paper. Profundal Benthos in Loch Ness. By D.S. Martin and A.J. Shine (Loch Ness
and Morar Project) and Annie Duncan (Department
of Biology, Royal Holloway University of
London).
Poster
paper. Loch Ness Project, Reviews and Previews. By A.J. Shine (Loch Ness and Morar Project).
Poster
paper. Loch Ness Undercurrents 1 and 2. By A.J. Shine (Loch Ness and Morar Project).
Note 6
London
Freshwater Group. 50th Meeting, Linnean
Society, London, 19th March 1993.
Paper
read. The Biology of Loch Ness. By D.S.
Martin (Loch Ness and Morar Project).
Appendix
Research
work carried out at Loch Ness towards the
degrees of B.Sc., M.Sc., M.Phil. or Ph.D.
include the following:
Baker,
D.L. (1993). A
Palaeolimnological Reconstruction of the
Recent History of the Catchment of a Scottish
Loch. B.Sc. Dissertation, University
of Wolverhampton.
Bennett,
S. (1993). Patterns
and Processes of Sedimentation in Loch Ness. B.Sc. Dissertation, University of Staffordshire.
Bracewell,
C.E. (1993). A
Geochemical Study of Natural and Pollutant
Compounds in Loch Ness, Scotland. M.Sc.
Dissertation, University of Newcastle-upon-Tyne.
Fulcher,
A.S. (Ongoing).
Rotifers of Loch Ness and the Cumbrian
Lakes. Ph.D. Thesis, University of
Lancaster.
Griffiths,
H.I. (Ongoing). Applications of Freshwater Ostracods
in the Study of Late Quaternary Palaeoenvironments
of North Western Europe. Ph.D. Thesis,
University of Cardiff.
Hartley, A. (1993). Plankton Bioassay of Loch Ness Water. B.Sc. Dissertation, University of Lancaster.
*
Jayakodi, U. (Ongoing). The Ecology of Zooplankton in Loch Ness. Ph.D. Thesis, University of Lancaster. *
Jenkins, P.H. (Ongoing). Comparative Effects of Environmental
Change, Human Impact and Climatic Change
on a Large and Small Lake. Ph.D. Thesis,
University of Wolverhampton.
Mansfield, C.A. (1992). A Study of Biogenic and Anthropogenic
Compounds in Sediment Cores from Loch Ness,
Scotland. M.Sc. Dissertation, University
of Newcastle-upon-Tyne.
Marjoram, R.S. (1993). An Investigation of the Identification
and Behaviour of an Acoustic Scattering
Layer in Loch Ness, Scotland. B.Sc.
Dissertation, Worcester College of Higher
Education.
Meacham, N.J. (1993). The Fecundity and Associated Ecological
Factors of the Arctic Charr, Salvelinus
alpinus, and Brown Trout, Salmo
trutta,
in Loch Ness, Scotland. B.Sc. Dissertation, University of
Hull.
Miller, K.C. (1993). A Study of Sedimentary Markers within
the Lacustrine Environment. B.Sc. Dissertation,
University of Edinburgh.
Millward,
D. (1992). A Palynological Sedimentation
Study of a Core from the South Basin of
Loch Ness. B.Sc. Dissertation, University
of Hull.
Picots, A. (Ongoing).Studies
of Bacterioplankton in Loch Ness, plus Limiting
Factors. M.Sc. Dissertation, University
of Lancaster.
Walton, M. (Ongoing). The Population Dynamics of Bacteria and
Flagellated Protozoa in Loch Ness. Ph.D.
Thesis, University of Lancaster. *
Wheeler, A. (Ongoing). Structure, Origin and Periodicity of
Laminations in Holocene Sediment Cores from
Loch Ness, Scottish Highlands. Ph.D.
Thesis, University of Wolverhampton.
Young, J.M. (Ongoing). Picoplankton in Loch Ness. M.Phil.
Dissertation, University of Lancaster. *
*Research
carried out under the auspices of the University
of Lancaster's multi-disciplinary plankton
study - Plankton Community Dynamics of a
Large Oligotrophic Freshwater System (Loch
Ness) - funded by the Natural Environment
Research Council (N.E.R.C.).
References
Anon. (1933). Strange
spectacle on Loch Ness. What was it? (From
a correspondent). Inverness Courier,
2nd May 1933.
Baker, P.F. (1962). Cambridge University Loch Ness Expedition
Report. Cambridge.
Baker, P.F. and Westwood, M. (1960). Underwater detective work. Scotsman,
14th September 1960.
Bennett S. and Shine, A.J. (1993). Review of current work on Loch Ness sediment
cores. Scottish Naturalist, 105: 55-63.
Campbell, E.M. and Solomon, D. (1972). The Search for Morag.
London: Tom Stacey.
Fulcher, A.S. (1993). Studies of the plankton of Loch Ness, Scotland. 3. Rotifers. Verhandlungen der Internationalen
Vereinigung fur Theoretische und Angewandte
Limnologie, 25: 460.
Griffiths,
H.I. and Martin, D.S. (1993). The spatial distribution of benthic ostracods
in the profundal zone of Loch Ness. Scottish
Naturalist, 105:
137-147.
Jenkins,
P.H. (1993a). Loch Ness sediments: a preliminary report. Scottish Naturalist, 105:
65-86.
Jenkins,
P.H. (1993b). Results of a water chemistry
study of Loch Ness. Scottish Naturalist,
105: 45-54.
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.
Laybourn-Parry, J.E.M. and Walton, M.C. (1993). Studies of the plankton
of Loch Ness, Scotland. 2 The microbial
loop. Verhandlungen der Internationalen
Vereinigung fur Theoretische und Angewandte
Limnologie, 25: 459.
Mackal, R.P. (1976). The
Monsters of Loch Ness. London: Macdonald
and Jane's.
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.
Mortimer, C.H.(1952). Water movements in lakes during summer stratification. Philosophical Transactions of the Royal
Society of London, Series B, 236: 355-404.
Mortimer, C.H. (1955). Some effects of the Earth's rotation on
water movement in stratified lakes. Proceedngs
of the International Association of Applied
Limnology, 12:
66-77
Murray, J. and Pullar, L. (Eds.) (1903-08). Bathymetrical
survey of the fresh-water lochs of Scotland. Geographical Journal, Vols. 22-31. A series of papers by various
contributors, preliminary to the six-volume
publication of 1910.
Murray, J. and Pullar, L. (Eds.) (1910). Bathymetrical
Survey of the Scottish Fresh-Water Lochs. Vols.
1-6. Edinburgh: Challenger
Office.
Rines, R.H. (1988). A review of research contributions at Loch
Ness by the Academy of Applied Science. Scottish Naturalist, 100:
201-208.
Rines, R.H., Wyckoff, C.W., Edgerton, H.E.
and Klein, M. (1976). Search for the Loch Ness Monster. Technology
Review, March/April 1976: 25-40.
Sanders, G. , Jones, K.C. and Shine, A.J. (1993). The use
of a sediment core to reconstruct the historical
input of contaminants to Loch Ness: PCBs
and PAHs. Scottish Naturalist, 105: 87-111.
Scott, P. and Rines, R.H. (1975). Naming the Loch Ness Monster. Nature, 258: 466-468.
Shine,
A.J., Kubecka, J., Martin, D.S. and Duncan,
A. (1993). Fish habitats in Loch Ness. Scottish Naturalist, 105: 237-255.
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., Bennett, S. and Marjoram,
R.S. (1993). Allochthonous
organic inputs as an explanation of spatial
biomass gradients observed in the pelagic
and profundal zones of Loch Ness. Scottish
Naturalist, 105:
257-269.
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.
Smith, B.D., Maitland, P.S., Young, M.R.
and Carr, M.J. (1981). The littoral zoobenthos. In: The Ecology
of Scotland's Largest Lochs: Lomond, Awe,
Ness, Morar and Shiel. (Ed. P.S. Maitland). Monographie Biologicae, 44: 155-203. The
Hague: Junk.
Thorpe, S.A. (1974). Evidence of Kelvin-Helmholtz billows in
Loch Ness. Limnology and Oceanography, 19:
973-976.
Thorpe, S.A. (1988). "...
light on obscure oceanographical problems";
an historical review of studies of the physics
of Loch Ness. Scottish Naturalist,
100: 17-33.
Thorpe, S.A., Hall, A. and Crofts, I. (1972). The internal surge in Loch Ness. Nature,
237: 96-98.
Watson, E.R. (1904). Movements of the waters of Loch Ness, as
indicated by temperature observations. Geographical Journal, 24: 430-437.
Wedderburn, E.M. (1907a). The temperature of the fresh-water lochs
of Scotland, with special reference to Loch
Ness. With appendix containing observations
made in Loch Ness by members of the Scottish
Lake Survey. Transactions of the Royal
Society of Edinburgh, 45: 407-489.
Wedderburn, E.M. (1907b). An experimental investigation of the temperature
changes occurring in fresh-water lochs. Proceedings of the Royal Society of Edinburgh,
28:
2-20.
Wedderburn, E.M. (1911). Some analogies between lakes and oceans. Internationale Revue der Gesamten Hydrobiologie
und Hydrographie, 4: 55-64.
Wedderburn,
E.M. (1912). Temperature
observations in Loch Earn. With a further
contribution to the hydrodynamical theory
of the temperature seiche. Transactions
of the Royal Society of Edinburgh, 48: 629-695.
Wedderburn, E.M. and Watson, W. (1909). Observations with a current
meter in Loch Ness. Proceedings of the
Royal Society of Edinburgh, 29: 619-647.
Witchell, N. (1975). The Loch Ness Story. London: Penguin
Books. Revised edition (1989). London:
Corgi Books.
Young, I. and Shine, A.J. (1993). Loch Ness bathymetric and seismic survey,
December 1991. Scottish Naturalist, 105: 23-43.
Young, J.M., Jones, R.I. and Bailey-Watts,
A.E. (1993). Studies
of the plankton of Loch Ness, Scotland.
1. Phytoplankton. Verhandlungen der Internationalen
Vereinigung fur Theoretische und Angewandte
Limnologie, 25: 458.
Received
July 1993
Mr. Adrian J. Shine, Loch Ness and Morar
Project,
Loch Ness Centre, DRUMNADROCHIT, Inverness-shire
IV3 6TU.
Mr. John Minshull, Loch Ness and Morar Project,
Loch Ness Centre, DRUMNADROCHIT, Inverness-shire
IV3 6TU.
Mrs Maralyn Shine, Loch Ness and Morar Project,
Loch Ness Centre, DRUMNADROCHIT, Inverness-shire
IV3 6TU.