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The Profundal Fauna of Loch Ness and Loch Morar

Vol 105, The Scottish Naturalist: Profundal Fauna of Loch Ness and Loch Morar p119

Reproduced with the permission of the Scottish Naturalist
Copyright: May be used for private research. All other rights reserved
 

By DAVID S. MARTIN
Loch Ness and Morar Project
ADRIAN J. SHINE
Loch Ness and Morar Project
ANNIE DUNCAN
Hydroacoustic Unit, Department of Biology,
Royal Holloway University of London

 Introduction 

The importance and biological interest of the profundal faunas of Loch Ness (230 m) and Loch Morar (310 m) is that they inhabit depths greater than 200 m or 300 m, which are deep on a world scale.  The profundal sediments are also very extensive, occupying over 50% of the area of Loch Ness greater than 150 m deep (Smith, Cuttle and Maitland, 1981) and 27% greater than 200 m deep; the corresponding percentages for Loch Morar are 28% and 19% (Mr. D.S. Martin, pers. comm.).  After the early studies instigated by Sir John Murray (Murray, 1904 and 1908; Murray and Pullar, 1910), few authors published on the fauna inhabiting the deep sediments of Loch Ness or Loch Morar until Maitland (1981), Shine and Martin (1988), and Griffiths, Martin, Shine and Evans (1993).  Little work was done on the profundal fauna of five Scottish lochs during the 1977-80 survey (Maitland, 1981) but Smith et al. (1981) reviewed what had been recorded earlier.  Dredgings from 90-230 m depths in Loch Ness by Murray and Pullar collected the bivalve Pisidium, subsequently identified as P. conventus and P. personatum, the lumbriculid Stylodrilus heringianus, and Chironomus sp., together with the 'casual' occurrence of Hydra and Lymnaea peregra from depths of 120-180 m.  Samples taken from c. 200 m depth in Loch Morar contained two lumbriculid worms, Lumbriculus variegatus and Stylodrilus heringianus (Smith et. al., 1981).

These authors also recorded that the profundal sediments of Loch Ness, taken by core, consisted of fine size fractions with clay (<39 m, 45.8%), fine silt (39-63 m, 18.1%) and sand (63-125 m, 35.4%), low organic content (loss on ignition, 8.9%) and a pH of 5.4.  The paucity of records is largely due to the difficulties of sampling sediments at the great depths of these lochs.  As part of the sampling

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programme undertaken by the Loch Ness and Morar Project, Shine and Martin (1988) designed a variety of collecting techniques, and compiled a list, not previously published, of thirty profundal zoobenthic species, at an average density of 295 individuals/m2, from Loch Ness.  Ostracods formed an important component, comprising over 60% of the fauna by numbers.  The four species recorded from the deep sediments (>200 m) of Loch Ness were Candona angulata G.W. Müller, C. candida (O.F. Müller), Cryptocandona reducta (Alm) and Cypria ophthalmica (Jurine) (Griffiths et al., 1993).

This present paper publishes the full listing of macrobenthic and meiofaunal species from the profundal sediments of both lochs, collected over the past ten years during the Loch Ness and Morar Project's benthic sampling programme.

Methods

Sampling Sites 

The samples from Loch Ness were mostly collected between June and October of 1982-85, from mid-loch and in depths between 189 and 230 m, from the areas Horseshoe Scree to Alltsigh and between Inverfarigaig to Urquhart Bay (Figure 1a, 7K ).  The profundal sites (>200 m) sampled during 1991-93 are plotted in Figure 2 (9K).  Samples from Loch Morar, collected during July or August in 1978-80 and 1985, were all taken from depths of 295-310 m in the deep basin between Meoble Bay and Swordlands (Figure 1b, 7K).  Echo-sounders were used to measure the depths being sampled, as well as to detect and control the descent of the sampling gear in situ, especially the Ekman sampler (Figure 3, 16K chart). 

Collecting Gear 

In the absence of the expensive Jenkins mud-sampler, several techniques were used to collect samples from the deep sediments during the earlier sampling period (1978-85) (Figures 4a, 4b, 4c,and 4d).  Unquantitative samples were taken in both lochs, using a weighted dredge with a fine mesh cod-end (Figure 4a, 8K) which collected the larger organisms; additional unquantitative methods, used only in Loch Ness, were fine-meshed eel-traps on the loch bottom and a set of gill nets of 50 mm mesh (1.4 m high and 125 m long) kept suspended with the lead-line resting on the sediments.  Left overnight, these caught profundal fish and some invertebrates.  Quantitative samples were mostly taken by a weighted Ekman grab (15 cm x 15 cm) or, in Loch Morar only, by a 5.0-litre brass Patalas volume sampler, which sampled an area 10 cm x 10 cm in the deep basin of 295-310 m depth.  Two special bottom samplers were designed by Adrian Shine to collect the surface few centimetres of sediment: a 0.25 m2 quadrat bottom sampler (Figure 4b, 20K) and a 0.25 m2 scoop bottom sampler (Figure 4c, 17K).

More recent sampling in Loch Ness (1991-93) was conducted using a new quantitative 10.3 cm diameter corer (Figure 4d, 7K photo), which permitted sub-samples to be cut at 1.0 cm intervals, thus giving samples of 83.3 cm3 for each 1.0 cm depth interval.  A second technique used was to dredge the sediment surface.  In Loch Morar during the 1990s, only the deep basin (310 m) was sampled, by leaving a buoyed sterilised hessian sack on the bottom and retrieving it after two weeks.

Sample Treatment

Profundal samples were kept alive in a refrigerator, seived, and sorted live as soon as possible; samples for fine-sorting in London were transported overnight in a cold container.  Some chironomids were kept alive until they emerged as adults.  Only Loch Morar sediment samples were washed through a series of seives down to 75 micro/m mesh.  The most effective method of quantitative sorting of the Loch Ness quantitative samples was by visually panning a series of diluted 10 ml sub-samples until the whole sample was completed.  This was continued until no more live animals were detected.  Sorted animals were preserved and sent to taxonomic specialists for identification (Tables 1 and 2).  The 1990s Loch Ness samples from the corer were mostly cut into slices, 0.0 - 1.0 and 1.0 - 2.0 cm from the surface mud, and totally sorted.  The whole mud sub-sample was examined microscopically, one spatula load at a time, and diluted to the thinnest layer possible in a petri dish.  The surface dredge samples were seived through 100 m mesh after settlement of larger particles.

The Loch Ness records for the period 1982-85 came from a total of 37 samples (eight by dredging, six by the scoop or quadrat samplers, and 23 by Ekman sampler) and for the 1991-93 period from a total of 11 samples (seven by dredging and four by coring); the Loch Morar records for 1978-85 came from a total of 12 samples (eight by Patalas sampler, three by Ekman sampler, and one by dredging). 

Results 

Twenty-nine species or genera of profundal invertebrates, and two species of fish, were recorded in Loch Ness during the earlier period of sampling, compared with only seventeen species or genera of invertebrates in Loch Morar (Table 1).  The lesser number of species recorded in Loch Morar reflects the smaller sampling

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effort, and the fewer collecting techniques employed, rather than implying any real difference.  The species came from 200 m depth.  Benthic collections were made at other depths, but are excluded from this paper.  In Table 1, most of the species collected were the larger macrobenthic forms present, due to the methods of collection.  That these methods did collect the small ostracods and copepods probably reflects their numerical abundance.  Some collecting methods turned out to be selective; for example, the scoop sampler (Figure 4c) was better at catching cyclopoid copepods, which swim just above the sediments, than the Ekman sampler.

Table 2 lists the species collected during the more recent sampling series (1991-93), using better methods of collection (e.g. a corer), with smaller volumes of mud, more easily sorted accurately and quantitatively.  This list includes additional species/genera recorded, and also contains more of the smaller meiobenthic groups of animals which are more difficult to extract from the mud.  This brings the total for Loch Ness to sixty-four invertebrate species/genera, plus two fish species, and for Loch Morar to thirty six species/genera.  These lists are based on extracting whole live specimens.

Further species were later added to the list, by accepting the chironomid head capsules sorted from the top few cms of Loch Ness core samples.  These further species were identified by Mr. D.P. Gallagher as follows: Heterotrissocladius, Orthocladius, Synorthocladius, Paratrichocladius, Psectrocladius, Pseudo-chironomus and Tanytarsus.

Comparing the two lochs, twenty-one species or genera were recorded in both lochs.  With more frequent and more extensive sampling, it seems likely that the number of species recorded from Loch Morar will increase.  In Loch Lomond, the only other major Scottish loch in which the profundal benthos has been sampled, only nine species (four oligochaete, two of Pisidium, and three chironomid species) were recorded (Weerekoon, 1956; Slack, 1965; Maitland, 1981).

 

Discussion

Under the stable environmental conditions of the profundal (200 m) of Loch Ness, i.e. constant cold (5.6C), darkness, high levels of dissolved oxygen (>80% saturated), low conductivity (<100 uS.cm-1) and great hydrostatic pressure (Shine and Martin, 1988), a fauna of surprising diversity exists, which can be sub-divided as follows:

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(A)The true profundal macrobenthic groups of chironomids (with 13 species/genera), oligochaetes (14 species) and the sphaeriid bivalve Pisidium (four species), each containing a dominant species within their group.

(B) The true profundal meiobenthic groups of the ostracods (six species), the nematodes (five species), harpacticoid copepods (three species) and cyclopoid copepods (two species), each contributing dominant species within their group.

(C) A series of macrobenthic groups, comprising turbellarians (two species), isopods (two species), amphipods (one species), and Trichoptera (three species); and meiobenthic groups, comprising protozoans (five species), microturbellarians (one species), rotifers (two species), tardigrades (one species), and oribateids (one species), with fewer representatives and whose distribution is not characteristically profundal.  The chydorid cladocerans (seven species) have also been placed in this group, for reasons given below.

The two fish species, i.e. Lamprey Lampetra sp. and Charr.

Apart from the fish, these organisms are all living on or in the sediments, or in the water just above the sediments of very fine fraction (Smith et al., 1981), and with an organic content which ranges between 18.29% at Inverfarigaig at 210 m to 33.9% at Invermoriston at 210 m (Mr. J.D. Hamilton, pers. comm.).  The new values are higher than the 8.9% organic content recorded by Smith et al. (1981), which came from a single core taken in the less deep shoulder of Loch Ness between the North and South Basins where the loch is under the influence of the River Foyers (Figure 2).  In the profundal, most of the listed species will be detritivorous, apart from the carnivorous groups like the turbellarians, cyclopoid copepods, tanypodine chironomids and fish.

Sub-division (A)

The chironomids, oligochaetes and species of Pisidium belonging to sub-division (A) contribute species to the profundal macrobenthos in some other deep, oligotrophic lakes of Europe, e.g. Lake Malaren (Ahren and Grimas, 1965), Thingvallavatn (Lindegaard, 1992), Lago Maggiore (Lenz, 1954; Della Croce, 1955; Stella, 1964; Bonomi, 1967; Kuiper, 1981), Lago di Mergozzo (Bonomi and Ruggin, 1966), and Lake Geneva (Zschokke, 1911; Juget, 1958; Dussart, 1967); and of North America, e.g. Great Slave Lake (Rawson, 1953), and the St. Lawrence Great Lakes (Cook and Johnson, 1974).

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Sub-division (B)

Similarly, the sub-division (B) groups contribute to the profundal meiobenthos in Finnish lakes, e.g. Paajarvi (Holopainen and Paasivirta, 1977) and Paijanne (Sarkka, 1992); in Polish lakes (Prejs and Papinska, 1983); and in pre-alpine Austrian lakes, e.g. Vorderer Finstertaler See (Bretschko, 1973, 1975, 1984), Piburger See (Pehofer, 1977) and Mondsee (Danielopol et al., 1988).

 

Sub-division (C)

Sub-division (C) is very heterogeneous, and consists of species able to survive in the profundal zone but not especially characteristic of it.  For example, nine of Fryer's (1985) list of crustacean species, collected more from large rather than small water bodies in his survey of 207 lowland Yorkshire waters, are recorded in Loch Ness or Loch Morar:  of these, four are chydorid cladocerans (Alona affinis, Chydorus sphaericus, Eurycercus lamellatus and Acroperus harpae), plus the ostracods Cyclocypris ovum and Cypria ophthalmica, the cyclopoids Paracyclops fimbriatus and Megacyclops viridis, and the isopod Asellus aquaticus.  These might be termed vagile (widely dispersive) organisms.  Some of sub-division (C) are introduced species:  Phagocata woodworthi (turbellarian) (Reynoldson, Smith and Maitland, 1981) and Crangonyx pseudogracilis from North America.  The presence of Asellus spp. in the profundal is less surprising when one considers how the amphipod Pontoporeia affinis dominates the deep profundal benthos of the St. Lawrence Great Lakes (Cook and Johnson, 1974) and the Great Slave Lake (Rawson, 1953) in North America, although to a greater extent than seems to occur in European lakes.  The Trichoptera larvae (and Plecoptera nymphs in Loch Morar) are probably littoral in origin, but are able to survive in the well-oxygenated profundal sediment surface.  However, in the few biological studies of the deep sediments of oligotrophic lakes which include the profundal meiobenthos, species are recorded in the profundal which belong to the same animal groups listed in sub-division (C), plus some others (Holopainen and Paasivirta, 1977).  Many of the sub-division (C) species are rare or exist as sparse populations able to survive under the stable environmental conditions of Loch Ness deep sediments (Giller, 1984).

Sub-division (D)

 The Lamprey was a larva and was caught by the hessian sack technique.  More information on the numbers and sizes of profundal Charr caught can be found in Shine and Martin (1988) and Shine, Kubecka, Martin and Duncan (1993).

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More detailed accounts of the seven groups in sub-division (A) and sub-division (B), i.e. Chironomidae, Oligochaeta, Sphaeriidae, Ostracoda, Nematoda, Harpacticoid Copepoda, and Cyclopoid Copepoda, are as undernoted:

Chironomidae
The taxonomic composition of the profundal Chironomidae was more diverse in Loch Morar than in Loch Ness, despite the fewer species collected.  The Loch Ness forms belonged to only two sub-families, the Tanypodinae (six taxa), with predacious head capsules and capable of feeding on small invertebrates, and the Chironominae (six taxa), with a non-carnivorous head capsule, a variety of feeding habits and diets, and capable of producing haemoglobin and silken threads for tube-building (Bryce and Hobart, 1972).

Among the Loch Morar taxa, one was tanypodine, three were chironomine, and two belonged to the Orthocladiinae, another non-carnivorous sub-family capable of producing silk but not haemogloblin (that is, without preadaptation to low oxygen conditions).  In Loch Ness the two most numerous species were the chironomine Sergentia coracina and the tanypodine Procladius spp., and in Loch Morar Sergentia spp. were dominant.  The presence of the orthocladine Heterotrissocladius grimshawi and tanytarsine Stempellinella spp. in Loch Morar suggests a greater degree of oligotrophy than in Loch Ness (Cook and Johnson, 1974; Lindegaard, 1992).

Oligochaeta
In both lochs, three families of oligochaetes were represented: Naididae (Specaria josinae - Lake Paijann; Sarkka, 1992), Tubificidae (Spirosperma ferox and Tubifex tubifex - Thingvallavatn; Lindgaard, 1992), and Lumbriculidae (Stylodrilus heringianus and Lumbriculus variegatus - Thingvallavatn; Lindegaard, 1992), in each case with characteristic deep-water profundal species (indicated in brackets) recorded in similar circumstances elsewhere.  In the oligotrophic Finnish Lake Paijanne (Sarkka, 1992), a higher proportion of naidid over tubificid species indicated a greater degree of oligotrophy compared with eutrophicated areas of the lake.  The most abundant oligochaete species was the tubificid Spirosperma ferox in Loch Ness, and S. velutinus in Loch Morar.

Sphaeriidae

According to Odhner (1923) and Kuiper (1974), Pisidium conventus is an arctic species which survives in the cold regions of alpine and deep temperate

 

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lakes.  In Loch Morar it was the only species collected, and of the four species in Loch Ness, it was the most abundant species, together with P. personatum.

Ostracoda

This was the most abundant group collected in both lochs. Of the four species recorded in Loch Morar, Cyclocypris ovum and Cypria ophthalmica are listed by Fryer (1985) as crustacean species showing a preference for large water bodies.  Candona candida and Cryptocandona reducta, present in both lochs, were less common in Loch Ness than Cypria ophthalmica and Candona angulata (Griffiths et al., 1993), who comment upon the low ostracod species diversity they found.  Danielopol et al., (1985, 1988) recorded as many as nine species of ostracods in the profundal of Mondsee, Austria, two species of which occur in Loch Ness, Candona candida and C. neglecta (Table 1).

Nematoda

The nematodes listed in Table 2 are the commoner species collected from the profundal sediments of Loch Ness 200 m where they attained densities of up to 10,000 individuals/m2 (Mr. D.S. Martin, pers. comm.).  It is not an exhaustive list, and more species (Eumonhystera filiformis group, Eumonhystera cf. longicaudatula, Ethmolaimus pratensis group, Aphanolaimus sp. and Dorylaimus cf. stagnalis) and greater densities (particularly of Ironus tenuicaudatus, with densities of up to 20,000 individuals/m2) are found in lesser depths such as 50-170 m (Dr. F. Schiemer, pers. comm.).  The species listed consist of widely distributed nematode species known from a wide range of habitats, such as ponds, rivers, and especially lakes, but together these species form a nematode association characteristic of the profundal zone of deep oligotrophic lakes with fine sediments, low food availability and good oxygen conditions.

Such nematode associations, with different species missing depending upon particular local conditions, have been found in the Austrian alpine and pre-alpine lakes, such as Vorderer Finstertaler See (Bretschko, 1984), Attersee (Dudinski, 1979), and Piburger See (Pehofer, 1977); in Polish alpine lakes (Prejs, 1977b); in the Finnish oligotrophic Lake Paajarvi (Holopainen and Paasivirta, 1977); and in the arctic Lake Char (Prejs, 1977a).  It seems that the species composition of profundal nematodes, and the carrying capacity of the sediments, are influenced less by interspecific competition and more by the combined influence of the nature of the sediments, the degree of oxygenation and the level of food available (Dr. F. Schiemer, pers. comm.).

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Harpacticoid Copepoda

The same four species of harpacticoid copepods have been found by Holopainen and Paasivirta (1977) in the deep oligotrophic Lake Paajarvi, along with profundal nematode and chironomid species associations.  In the Finnish Lake Paijanne, Sarkka (1992) found that Attheyella crassa showed a marked preference for the well-oxygenated end of this oligotrophic lake, which is undergoing eutrophication at the other end.

Cyclopoid Copepoda

Paracyclops fimbriatus, which is a crustacean preferring larger water bodies (Fryer, 1985), occurred in both Loch Ness and Loch Morar, and has been recorded by Dussart (1967) in Lake Geneva and by Sarkka (1992) in Lake Paijanne, where the profundal cyclopoids are species of Diacyclops.  Another species commonly recorded in the profundal literature is Megacyclops viridis, which was often taken in the scoop samples and is a large enough animal to be a significant predator.


Conclusion

Without further study, it is difficult to determine whether the profundal benthos of these two deep Scottish lochs is a random collection of species populations accumulated over time, or a structured assemblage of interacting species populations of all trophic levels inhabiting the deep, cold, fine-structured and well-oxygenated loch sediments (Giller, 1984).  The profundal benthos is surprisingly species diverse, and may well represent a saturated equilibrium community (that is, with filled ecological niches), as defined by Giller (1984).  The age of the lochs, their environmental stability, their well-oxygenated conditions, and the not too low levels of organic food available in the sediments, all support this possibility.

Further biological studies on both the macrobenthos and meiobenthos should concentrate on quantifying both the vertical distributions and any differences between the North and South Basins in Loch Ness.  In order to understand the role of the deep benthos on the loch ecosystem, there is also a need for life-cycle studies of the dominant species, by seasonal sampling and experimental rearing, and, later, unravelling the biotic interactions between species.

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Acknowledgements

The Loch Ness and Morar Project would like to record its deep gratitude to the undernoted experts who gladly undertook the identification of specimens sent to them, without which Table 1 and Table 2 would not exist, or who contributed valuable assistance in other ways:

Dr. P.C. Barnard (Natural History Museum), Dr. J.A. Bass (Institute of Freshwater Ecology, I.T.E. Monks Wood), Dr. G. Bird (National Rivers Authority, Yorkshire Region), Dr. G.A. Boxshall (Natural History Museum), Dr. S. Brooks (Natural History Museum), Dr. P.C. Cranston (Natural History Museum, and C.S.I.R.O., Canberra, Australia), Dr. C. Duigan (Countryside Commission for Wales), Dr. J.P. Ellis (Natural History Museum), Dr. G. Fryer (Freshwater Biological Association), Mr. D.P. Gallagher (University of Dublin), Mr. R.B. Greer (Department of Agriculture and Fisheries for Scotland), Mr. H.I. Griffiths (University of Leeds), Mr. J.D. Hamilton (University of Paisley), Dr. R. Hamond, Dr. P.A. Henderson (Marine Biology Unit, Fawley), Dr. M.P. Kerney (Natural History Museum), Dr. M. Ladle (Institute of Freshwater Ecology, Wareham), Dr. M. Learner (University of Wales), Professor H. Löffler (University of Vienna), Mr. D.S. Martin (Loch Ness and Morar Project), Dr. P. Mordan (Natural History Museum), Dr. L.C.V. Pinder (Institute of Freshwater Ecology, I.T.E. Monks Wood), Dr. R.M. Pontin (Royal Holloway University of London), Professor F. Schiemer (University of Vienna), Dr. R.W. Sims (Natural History Museum), Dr. I. Strachan (Scottish Natural Heritage), Mr. A.C. Wheeler (Natural History Museum), and Dr. J.F. Wright (Institute of Freshwater Ecology, Wareham).

The Project also gratefully acknowledges help from Mr. John Minshull at Loch Ness, and the early enthusiastic contributions by Mr. Richard Grinvalds to the Loch Morar species list.

 

References 

Ahren, T. and Grimas, U.  (1965).  The composition of the bottom fauna in two basins of Lake Malaren.  Report, Institute of Freshwater Research, Drottningholm, 46: 49-57.

Bonomi, G. (1967).  L'evoluzione recente del Lago Maggiore rivelata dalle conspicue modoficazioni del macrobenton profundo.  Memorie della Istituto Italiano di Idrobiologia, 21: 197-212.

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Bonomi, G. and Ruggin, D.  (1966).  Il macrobenton profundo del Lago di Mergozzo, Italia.  Memorie della Istituto Italiano di Idrobiologia, 20: 153-200.

Bretschko, G.  (1973).  Benthos of a high-mountain lake: Nematoda.  Proceedings of the International Association of Theoretical and Applied Limnology, 18: 1421-1428.

Bretschko, G.  (1975).  Annual benthic biomass distribution in a high-mountain lake (Vorderer Finstertaler See, Tyrol, Austria).  Proceedings of the International Association of Theoretical and Applied Limnology, 19: 1279-1285.

Bretschko, G.  (1984).  Free-living nematodes of a high-mountain lake (Vorderer Finstertaler See, Tyrol, Austria).  I.  Monhystera cf. stagnalis and Ethmolaimus pratensis.  Archiv für Hydrobiologie, 101: 39-72.

Bryce, D. and Hobart, A. (1972).  The biology and identification of the larvae of Chironomidae (Diptera).  Entomologist's Gazette, 23: 175-217.

Cook, D.G. and Johnson, M.G.  (1974).  Benthic macroinvertebrates of the St. Lawrence Great Lakes.  Journal of the Fisheries Research Board of Canada, 31: 763-762.

Danielopol, D.L., Geiger, W., Tölderer-Farmer, M., Orellana, C.P. and Terrat, M-N.  (1985).  The Ostracoda of Mondsee: spatial and temporal changes during the last fifty years.  In: Contributions to the Paleolimnology of the Trumer Lakes (Salzburg) and the Lakes Mondsee, Attersee and Traunsee (Upper Austria). (Ed. D.L. Danielopol, R. Schmidt and E. Schultze).  Pages 98-121.   Mondsee: Eigenverlag Institut für Limnologie.

Danielopol, D.L., Geiger, W., Tölderer-Farmer, M., Orellana, C.P. and Terrat, M-N.  (1988).  In search of Cypris and Cythere - a report of the Evolutionary Ecological Project on limnic Ostracoda from the Mondsee (Austria).  In: Evolutionary Biology of Ostracoda.  (Ed. T. Hanai, N. Ikeya and K. Ishizaki).  Pages 485-500.  Oxford: Elsevier.

Della Croce, N.  (1955).  The conditions of the sedimentation and their relationship with Oligochaeta populations of Lake Maggiore. Memorie della Istituto Italiano di Idrobiologia, 8: 39-62.

Dudinski, S.  (1979).  Das meiobenthos das Altersee profundals unter besonderdet berücksichtigung der freilebenden nematoden.  Arbeiten Labor Weyregg, 3: 198-208.

Dussart, B.H.  (1967).  Les Copépodes des Eaux Continentales.  Volumes 1 and 2.  Paris:  N. Boubée et cie.

Fryer, G.  (1985).  Crustacean diversity in relation to the size of water bodies: some facts and problems.  Freshwater Biology, 15: 347-361.

Giller, P.S.  (1984).  Community Structure and the Niche.  London:  Chapman & Hall.

Griffiths, H.I., Martin, D.S., Shine, A.J. and Evans, J.G.  (1993).  The ostracod fauna (Crustacea, Ostracoda) of the profundal benthos of Loch Ness.  Hydrobiologia, 254: 111-117.

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Holopainen, I.J. and Paasivirta, L.  (1977).  Abundance and biomass of the meiozoobenthos in the oligotrophic and mesohumic Lake Paajarvi, southern Finland.  Annales Zoologici Fennici, 14: 124-134.

Juget, J.  (1958).  Recherche sur la faune de Fond du Léman et du Lac d'Annecy.  Annales de la Station Centrale de Hydrobiologie Appliquée, 7: 9-96.

Kuiper, J.G.J. (1974). Een pleistocene vondst van Pisidium conventus Clessin Nederland en de huidige geografische verspreiding van deze soort in Europa. Basteria, 38: 27-40.

Kuiper, J.G.J.  (1981).  Zur frage der geographischen unterarten bei Pisidien insbesondere bei Pisidium personatum Malm.  Archiv für Molluskenkunde, 112: 9-19.

Lenz, F.  (1954).  Die bodenfauna des Lago Maggiore und ihre lebensbedingungen.  Memorie della Instituto Italiano di Idrobiologia, 8: 273-322.

Lindegaard, C.  (1992).  Zoobenthos ecology of Thingvallavatn: vertical distribution, abundance, population dynamics and production.  Oikos, 64: 257-304.

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.

Murray, J.  (1904).  Loch Ness.  Part V:  Biology of Loch Ness.  Geographical Journal, 24: 442-443.

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Murray, J. and Pullar, L.  (1910).  Bathymetrical Survey of the Scottish Fresh-Water lochs.  Vols. 1-6.  Edinburgh: Challenger Office.

Odhner, N.H.  (1923).  Mollusca.  Pisidium conventus Clessin.  Report of Scientific Results of the Norwegian Expedition to Noyaya Zemlya 1921.  No. 6: 3-7.  Kristiania, Norway:  Videnskapsselskapet i Kristiania.

Pehofer, H.E.  (1977).  Bestand und produktion benthoscher nematoden im Piburger See (Otztal, Tirol).  Dissertations of Department of Limnology, University of Innsbruck, 7: 1-87.

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Prejs, K.  (1977b).  The species diversity, numbers and biomass of benthic nematodes in central part of lakes with different trophy.  Ekologia Polska, 25: 31-44.

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Vol 105, The Scottish Naturalist: Profundal Fauna of Loch Ness and Loch Morar p136

Rawson, D.S.  (1953).  The bottom fauna of Great Slave Lake.  Journal of the Fisheries Research Board of Canada, 10: 486-520.

Reynoldson, T.B., Smith, B.D. and Maitland, P.S.  (1981).  A species of North American triclad (Paludicola; Turbellaria) new to Britain found in Loch Ness, Scotland.  Journal of Zoology, 193: 531-539.

Sarkka, J.  (1992).  Lacustrine profundal meiobenthos as an environmental indicator.  Hydrobiologia, 243/244: 333-340.

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.

Slack, H.D.  (1965).  The profundal fauna of Loch Lomond, Scotland.  Proceedings of the Royal Society of Edinburgh, Section B, 69: 272-297.

Smith, B.D., Cuttle, S.P. and Maitland, P.S.  (1981).  The profundal zoobenthos.  In:  The Ecology of Scotland's Largest Lochs:  Lomond, Awe, Ness, Morar and Shiel.  (Ed. P.S. Maitland).  Monographiae Biologicae, 44: 205-222. The Hague: Junk.

Stella, E.  (1964).  Il Megacyclops viridis Jurine forma batiale de Lago Maggiore.  Memorie della Istituto Italiano di Idrobiologia, 17: 57-79.

Weerekoon, A.C.J.  (1956).  Studies on the biology of Loch Lomond.  1.  The benthos of Auchentullich Bay.  Ceylon Journal of Science, Section B, 7: 1-94.

Zschokke, F.  (1911).  Die tiefseefauna der seen Mitteleuropas.  Eine geographische faunistische studie.  Monographien und Abhandlungen zur Internationalen Revue der Gesamten Hydrobiologie und Hydrographie, 4: 1-246.

Received June 1993

Mr. David S. Martin, Loch Ness and Morar Project,

Loch Ness Centre, DRUMNADROCHIT, Inverness-shire IV3 6TU.

Mr. Adrian J. Shine, 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.

 

 

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Loch Ness Profundal Fauna