A comparative assessment of temporal variation in diet of powan, Coregonus lavaretus (L.), from Loch Lomond and Loch Eck, Scotland, U.K.

Stomach contents of powan caught throughout the year from Loch Lomond and Loch Eck were examined. Adult powan in Loch Lomond fed mainly on planktonic Cladocera, switching to benthic food during December-April. Bosmina coregoni was the principal planktonic food with Daphnia hyalina an important late summer and autumn prey. Loch Eck powan fed throughout the year on chironomid larvae and Pisidium spp. from the benthos and ingested large amounts of non-food material. Mean dry weights of food in stomachs were correlated with water temperature in Loch Lomond but not in Loch Eck and were higher in powan from Loch Eck over most of the year.     

Shift in pike, Esox lucius L., predation pressure following the introduction of ruffe, Gymnocephalus cernuus (L.) to Loch Lomond

Rapid and dramatic change in the fish community of Loch Lomond has resulted from a series of fish introductions in recent years. A comparison of the diet of pike, Esox lucius L., in 1989–1990 with data from 1955–1967, prior to recent introductions, demonstrates a shift in prey choice. In 1955–1967 powan, Coregonus lavaretus (L.), dominated in the diet (57% of prey by number) by 1989–1990 the introduced ruffe, Gymnocephalus cernuus (L.), was the commonest prey species (44% by number). This shift in pike predation to an abundant population of introduced ruffe has consequences for native species. Assuming that the pike population has not increased in response to increased food availability due to the introduction of ruffe, pike predation pressure on native species will be relieved. This is likely to have the greatest effect on powan. Comparison of the predation rate in 1955–1967 with 1989–1990 supports the hypothesis that the rate of predation on powan has declined, although the effect that this may have on the powan population is unclear, as the role of predation in the regulation of population size is unknown for this species.     

The fish community of Loch Lomond,Scotland: its history and rapidly changing status

The fish community of Loch Lomond is of national importance. Its diversity of species and rare populations of powan (Coregonus lavaretus) and freshwater feeding river lampreys (Lampetra fluviatilis) warrant high conservation status. It is also of value for its sport fisheries for sea-trout (Salmo trutta), salmon (Salmo salar) and pike (Esox lucius). Historical records demonstrate that the species composition of the fish community has remained stable over a very long period until recently when a series of introductions of fish species new to the catchment has resulted in successful colonisation by a number of species. These have resulted in fundamental changes in the ecosystem. Here using historical records the long-term stability of the fish community is examined, recent rapid changes in the fish community are documented and some of the resultant effects of changes in the fish community are demonstrated.     

The genetic relationships of the coregonid fishes of Britain and Ireland indicated by electrophoretic analysis of tissue proteins

The electrophoretic patterns of the general proteins, esterases, creatine kinase and phosphoglucomutase of white muscle from Lough Neagh pollan, Lough Erne pollan, Lake Bala gwyniad, Loch Lomond powan, Haweswater schelly and Bassenthwaite vendace are consistently of three types. It is therefore suggested that the coregonids examined are three distinct species: the pollans; gwyniad, powan and schelly; vendace. Vendace is genetically closer to the gwyniad group than to the pollans.     

Zooplankton in Loch Lomond: perspectives,predation and powan

Published and unpublished accounts of the zooplankton community of Loch Lomond are reviewed. Loch Lomond's zooplankton community is unique amongst those of Scotland's other large lochs. Holopedium gibberum appeared in Loch Lomond in the years between the studies of Slack & Hamilton in the 1950s and zooplankton surveys in the late 1970s.The annual cycles of Eudiaptomus gracilis, Cyclops strenuus abyssorum and Mesocyclops leuckarti are described for the mid basin of Loch Lomond. E. gracilis overwinter mainly as adult with variable egg production during their lifespan of 5–8 months. The cyclopoid copepods have variable life history characteristics. C. str. abyssorum produce a single generation per year, overwinter by means of resting eggs and a small population of planktonic adults, while M. leuckarti is present in the plankton throughout the year, but in low numbers during winter.Zooplankton populations in Loch Lomond are predated by the diverse fish community which includes a facultative planktivore in the powan, Coregonus lavaretus. Powan feed heavily on zooplankton from late spring until late autumn. Powan display selectivity in the prey species taken, in the size distribution and morph types taken within prey species. Zooplankton species, morphs and forms in Loch Lomond tend to reflect this predation.Zoology Department, University of St. AndrewsZoology Department, University of St. Andrews     

Histopathology, ultrastructure and immunohistochemistry of Coregonus lavaretus hearts naturally infected with Ichthyocotylurus erraticus (Trematoda)

Histopathological, ultrastructural and immunohistochemical investigations were conducted on 26 specimens of powan Coregonus lavaretus (L.) from Loch Lomond (Scotland). The hearts of all 26 powan (15 females and 11 males) investigated harboured metacercariae of the digenean trematode Ichthyocotylurus erraticus (Rudolphi, 1809). The vast majority of metacercariae were located either singly or as an aggregation of white cysts on the surface of the bulbus arteriosus. The intensity of infection ranged from 2 to 200 larvae heart(-1), although the number of metacercariae found on male powan did not exceed 13. Histochemically, the parasite cyst wall gave a strong positive reaction with periodic acid schiff (PAS) and a faint positive signal with Azan-Mallory stain. All the metacercariae cysts were embedded in a granulomatous proliferation of heart epicardium tissue, forming a reactive fibroconnective capsule around the parasite. The capsule enclosing the parasite (produced by the host's reaction to the parasite) measured 13.57 to 90.20 microm (37.43 +/- 3.56) in thickness. Within the capsule wall, eosinophilic granular cells (EGCs), granulocytes, melanocytes and, in some instances, partially degenerated or vacuolated epithelioid cells were observed in close proximity to the cyst wall. Pigment-bearing macrophages were scattered throughout the granulomatous host-tissue reaction and as macrophage aggregates (MAs) within the capsules surrounding parasites. Immunohistochemical tests were applied to infected heart sections using 12 different antisera. Nerve fibres immunoreactive to bombesin, substance P (SP), and atrial natriuretic peptide (ANP) antisera were observed in close proximity to the parasite larvae. The presence of a serotonin-like substance was also observed within host immune-cells surrounding trematode cysts. Large cells of the epicardium were found to be immunoreactive to met-enkephalin and vasoactive intestinal peptide (VIP) antisera but not immunoreactive to anti-protein gene-product 9.5 (PGP9.5) sera.     

Life histories of the powan,Coregonus lavaretus (L.) (Salmonidae,Coregoninae) of Loch Lomond and Loch Eck

Four aspects of the life histories of the two populations of powan Coregonus lavaretus (L.) in Scotland are described: growth (Eck powan are shorter and with greater year to year variance than Lomond); sexual maturation (Eck powan mature younger, but at similar weight to Lomond); spawning (timing in Eck varies, but is consistent in Lomond); and recruitment/mortality (fecundity, sex ratios, and mortality also vary in the short term). Short term differences between the physiological ecology of the populations can be ascribed to the size and topography of the lochs. Long term differences are more difficult to account for, and are more important in that they may signal changes in sustainability. Conservation of powan must be considered in terms of their synecological relationships, not in isolation.     

Mycological aspects of a disease of young perch in Windermere

A disease of young perch in Windermere was investigated. In contrast to the case of U.D.N. infections of salmon and trout, a variety of fungi were obtained from the diseased fish. These reflected the spora present in the water at the same time to a large extent. An exception was Saprolegnia ferax , which did not grow on the fish and indicated that some degree of host specificity was involved. The Achyla isolates, both from the fish and from the water, were found to be heterothallic and male in reaction.     

Invertebrate food sources for waterbirds provided by the reconstructed wetland of Nyirkai-Hany,northwestern Hungary

Detailed studies of the macroinvertebrate benthos and zooplankton communities in Loch Leven, the largest shallow lowland lake in Scotland, UK, were carried out from 1966 to 1973 as part of the International Biological Programme (IBP). The results revealed a reduction in species diversity that was attributed to increasing eutrophication. This work provides a baseline against which the response of the invertebrate communities to subsequent changes in management can be assessed. This article compares macroinvertebrate benthos and zooplankton data from the IBP study with the post-IBP era during which changes at Loch Leven included a 60% reduction in the phosphorus input from external sources and variations in fish stocking rates. Only in recent years has there been evidence of ecological recovery by the invertebrate communities: the number of macroinvertebrate and zooplankton taxa has increased (including taxa considered to be sensitive to nutrient enrichment) and invertebrate abundances have declined. These changes appear to reflect the improvements in water quality and habitat conditions at Loch Leven that have occurred as a result of the recent reduction in nutrient loads, albeit with a substantial delay before any ecological response could be detected. This time lag in recovery has important implications for assessing improvements in the ecological status of other lake systems, as is required by the EU Water Framework Directive.     

Alternative migration and host parasitism strategies and their long-term stability in river lampreys from the River Endrick, Scotland

The stability of a discrete body size dimorphism of sexually mature river lamprey Lampetra fluviatilis from the River Endrick, Scotland, was examined over a 21 year period. Stable isotope analysis was used to test the hypothesis that the two size forms comprise individuals with differing migration and parasitic foraging strategies. Maturing river lamprey and the brook lamprey Lampetra planeri were trapped over 3 months each year in the periods 1983–1984 and 2004–2005. Brook lamprey catches and catches of both species combined showed no significant trend in catch rate with time. The catch rate of small body size river lamprey declined between 1983–1984 and 2004–2005 (although the difference did not reach statistical significance; P = 0·055). In contrast, there was a significant increase in the catch rate of the large body size river lamprey and as a consequence, a significant change in the relative proportion of each of the two river lamprey morphs over the study period. Analysis of the stable isotopes of C and N in muscle tissue showed that brook lamprey tissue derived its carbon from a freshwater source and had a δ¹³C more consistent with that of the River Endrick than with Loch Lomond. δ¹⁵N values for this species showed it to be feeding at the base of the food chain, consistent with filter feeding as an ammocoete. The large body size and the small body size river lamprey adults differed substantially in their δ¹³C values, with the small body size δ¹³C signature indicative of a freshwater carbon source and the large body size morph of a marine source. The small body size morph had a δ¹³C signature that was consistent with that of Loch Lomond powan Coregonus lavaretus suggesting that they share a common carbon source. The large body size morph was clearly feeding at a higher trophic level than the small body size morph. A single small body size river lamprey individual with typical morphology for that group, however, had C and N signatures that clustered with those of the large body size morphs. This individual had either migrated to sea to forage, as is typical for the species, or had been feeding on an anadromous fish with a strong marine C signature in fresh water. It is concluded that the body size dimorphism is indicative of a differential migration and foraging strategy in the parasitic phase of the life cycle of river lamprey at this site.     

Ferox Trout (Salmo trutta) as `Russian dolls': complementary gut content and stable isotope analyses of the Loch Ness foodweb

1. Conventional collection methods for pelagic fish species (netting, trawling) are impractical or prohibited in Loch Ness, U.K. To investigate trophic relationships at the top of the Loch Ness food web, an alternative strategy, angling, provided samples of the top predator, the purely piscivorous ferox trout ( Salmo trutta ).
2. The gut contents of these fish provided further samples of prey-fish, and subsequent examination of prey-fish guts revealed their dietary intake, analogous to the famous nested `Russian dolls'. Each trophic level separated by gut content analysis provided further complementary samples for stable isotope analysis and thus information on the longer term, assimilated diet.
3. Ferox trout exhibited considerable cannibalism to supplement a diet of Arctic charr ( Salvelinus alpinus ). However, conspecifics stemmed from a lower isotopic baseline in relation to charr, so ferox trout exhibited a lower trophic level than predicted (4.3) by using the δ¹⁵N values. Charr displayed dietary specialisation with increasing length, and isotopic values supported by the gut data placed the charr at a trophic level of 3.5. The isotope data also indicated that charr carbon was primarily autochthonous in origin.     

The reproductive cycle of Coregonus lavaretus (L) in Loch Lomond, Scotland, inrelation to seasonal changes in plasma cortisol concentration

The reproductive cycle of Coregonus lavaretus (L.) in Loch Lomond, Scotland, was investigated by monthly sampling. Spawing takes place during the first 3 weeks of January, on offshore gravel banks. Males congregate on the spawning-grounds throughout the spawning period, while ripe females cruise in unisexual shoals in deeper water. Individual females migrate to the spawning-grounds as ovulation takes place, spawn, and return to deep water. After spawning, the gonadosomatic ratio and somatic condition factor of both sexes decrease until June or July. Gonad recrudescence occurs in males between July and October, and in females between July and December. The somatic condition factor of both sexes rises from its minimum in June or July to its maximum in September. A competitive protein binding assay was used to determine cortisol levels in 0–1 ml plasma samples of individual fish. The lowest cortisol levels occurred in fish caught by seine-netting and killed immediately by anaesthesia in MS 222 (Sandoz). Seine-netting and killing by concussion induced higher cortisol levels, in the range of 3–5 ng/100 ml. Cortisol levels in fish caught by gill-netting for 18 h were 3–6 times higher than in seine-netted fish, and the method of killing was immaterial in this case. The length of time spent in the gill-net had no significant effect on cortisol level, but maintenance of the fish in aquaria for 24–80 h elicited high cortisol levels of over 50 μg/100 ml. Post-mortem delay before blood-sampling resulted in lowered cortisol levels. Seasonal variations in cortisol level were determined in gill-netted fish throughout the year, and in seine-netted fish when available. Cortisol levels were high in both sexes in September; and very high in ovulating females caught on the spawning-grounds, though not in ovulating females caught off the spawning-grounds.     

Parasite community structure in Pimephales promelas (Pisces: Cyprinidae) from two converging streams

Parasites of the fathead minnow, Pimephales promelas, were examined in fish collected from Elk Creek (40.88534 degrees N, 96.83366 degrees W) and West Oak Creek (40.90821 degrees N, 96.81432 degrees W), Lancaster County, Nebraska. These 2 streams are part of the Salt Valley watershed and flow together approximately 2 km downstream from the collection sites to form Oak Creek. This study examined the extent to which the 2 tributaries constitute a continuous habitat with respect to fish hosts. The parasite community included Trichodina sp., Myxobolus sp., Dactylogyrus simplex, D. bychowskyi, and D. pectenatus (all on gills); Gyrodactylus hoffmani (gill and body surface); Posthodiplostomum sp. (neascus, body cavity); and Uvulifer ambloplitis (encysted in skin). Among 46 fish from Elk Creek and 56 fish from West Oak Creek taken on 5 dates during April-July 1998, U. ambloplitis was found in Elk Creek fish at prevalences of 44-100% but in only 2 West Oak fish on 1 date. Prevalence and mean abundance of D. simplex also differed between the 2 sites. On the basis of these observations, fish populations in the 2 streams were considered to be distinct, with little or no fish movement between the tributaries.     

Factors limiting fish populations in the Loch Fleet system, an acidic drainage system in south-west Scotland

Loch Fleet is an oligotrophic upland lake in Galloway, south-west Scotland. It once supported a brown trout, Salmo trutta L., sport fishery with low annual catches (< 150 fish year ^?1) but catches declined markedly after 1950 and no fish were caught after 1975. Diatom records for the lake sediments indicate acute acidification since 1975, pH changing from c. 5–8 to 4–6. In 1984 a project was set up at Loch Fleet to investigate techniques of acidity mitigation, with a view to restoring fisheries in this and similarly affected waters. An underlying assumption of the project was that fish had been lost as a direct result of acidity and associated factors. Studies were therefore undertaken during a 2-year baseline period (1984–1986) to validate this assumption. Fish surveys using a variety of techniques (gill-netting, trapping, electrofishing) confirmed that trout were absent from the Loch and its afferent streams, and also from its main outlet stream, the Little Water of Fleet, for a distance of 7 km downstream. Trout were present below this point but are prevented from passing upstream by a 5-m waterfall. Eels, Anguilla anguilla L., were present throughout the Little Water of Fleet, though not in the Loch itself. Population densities of both species were low, with less than 7 eels and 5 trout per 100 m². Survival studies using brown trout ova and yolk-sac fry indicated that conditions in the Loch and its afferent streams were acutely toxic to these stages as a result of the low pH (pH 4.5), low calcium (I mg l^?1) and high aluminium concentrations (200 μg 1 ^?1 total Al, 60 μg 1 ^?1 inorganic monomeric Al). Trout fingerlings could survive these conditions in short-term tests (9 days) but, in chronic exposure tests lasting up to 180 days carried out in situ in streams adjoining the loch, no fish survived this period. This toxicity was eliminated in experiments where pH was raised to 5.4 by KOH addition. It is concluded that the loss of the brown trout fishery at Loch Fleet occurred as a direct result of acidity and related factors, probably acting in the first instance on the sensitive intra-gravel ova and yolk-sac fry stages, leading to recruitment failure.     

Morphometric analysis of two ecologically distinct forms of Arctic charr, Salvelinus alpinus (L.), in Loch Rannoch, Scotland

Gill–netted samples of Arctic charr from Loch Rannoch, Scotland were bimodal when tested by univariate and multivariate morphometric analyses. The separation into two morphs corresponded very closely (95–98%) with fish classified subjectively in the field as benthic or pelagic, based largely on colour differences and ecological observations. The benthic charr had relatively longer heads, larger eyes and more powerful jaws than the pelagic charr. Unlike sympatric morphotypes described from Scandinavia and Greenland, neither Rannoch morph is dwarfed.     

Immune response of U.D.N.-infected salmon to Saprslegnia

Sera from salmon Sulnio salar L. suspected of U.D.N.-infection were tested (by double gel diffusion) against antigens prepared from diseased salmon tissues and Suprolegniu . Precipitating antibodies to the fungus were found in 93 % of the salmon, but only 66% of these fish were colonized by fungus. The antibodies were detected in parr and adult salmon.     

The Relationship Between Spatial Distribution and Diet of Arctic Charr, Salvelinus Alpinus, In Loch Ness, U.K.

The pelagic fish community of Loch Ness, U.K., is dominated by Arctic charr, Salvelinus alpinus. Previous studies have shown that the distribution of zooplankton along the south-west to north-east axis of this elongate loch is very dynamic and determined largely by prevailing winds, but Arctic charr are consistently more abundant in the southern half of the loch. In July 1993, the diet compositions of 161 Arctic charr from 53 to 330mm in length were determined and related to their spatial distribution and those of microcrustacean zooplankton and pelagic chironomid larvae and pupae. Diets were dominated by chironomid larvae, with chironomid pupae, Bythotrephes longimanus, Bosmina coregoni and Daphnia hyalina also frequently taken. Over the whole study period, B. longimanus were more important in the diet of fish from the northern half of the loch, while chironomid larvae were more important for fish from the southern half. As a result, per capita prey weight of fish from the south was greater than that of fish from the north. However, wind-induced changes in the distribution of zooplankton along the loch were accompanied by a change in the diet composition of Arctic charr from the south.     

Population biology of the helminth parasites of Phoxinus phoxinus (L.), the minnow, in a Cardiganshire lake

Specimens of Phoxinus phoxinus (L.) were collected over a period of a year from September 1967 to September 1968 from Frongoch Lake, Cardiganshire, Wales. Of a total of 650 fish examined for helminth parasites, 32 % were definitive hosts for Macrolecithus papilliger , and 16·5% for Neoechinorhynchus rutili (Müller), both parasites of the alimentary canal. All the fish examined were second intermediate host for Diplostomum phoxini (Faust) in the brain and 1·67% for Ligula intestinalis (L.) in the coelom. An increase in incidence of infestation with M. papilliger in spring and autumn and with N. rutili in summer were evident. The sex of the fish had no apparent effect on worm burden whereas size was more significant. There was a decrease in incidence of infestation with M. papilliger and an increase with N. rutili together with an increase in intensity of infestation with D. phoxini as the fish became larger. The latter species accumulated in the brain as the host increased in age.     

Climate-induced changes in high elevation stream nitrate dynamics

Mountain terrestrial and aquatic ecosystems are responsive to external drivers of change, especially climate change and atmospheric deposition of nitrogen (N). We explored the consequences of a temperature-warming trend on stream nitrate in an alpine and subalpine watershed in the Colorado Front Range that has long been the recipient of elevated atmospheric N deposition. Mean annual stream nitrate concentrations since 2000 are higher by 50% than an earlier monitoring period of 1991–1999. Mean annual N export increased by 28% from 2.03 kg N ha⁻¹ yr⁻¹ before 2000 to 2.84 kg N ha⁻¹ yr⁻¹ in Loch Vale watershed since 2000. The substantial increase in N export comes as a surprise, since mean wet atmospheric N deposition from 1991 to 2006 (3.06 kg N ha⁻¹ yr⁻¹) did not increase. There has been a period of below average precipitation from 2000 to 2006 and a steady increase in summer and fall temperatures of 0.12 °C yr⁻¹ in both seasons since 1991. Nitrate concentrations, as well as the weathering products calcium and sulfate, were higher for the period 2000–2006 in rock glacier meltwater at the top of the watershed above the influence of alpine and subalpine vegetation and soils. We conclude the observed recent N increases in Loch Vale are the result of warmer summer and fall mean temperatures that are melting ice in glaciers and rock glaciers. This, in turn, has exposed sediments from which N produced by nitrification can be flushed. We suggest a water quality threshold may have been crossed around 2000. The phenomenon observed in Loch Vale may be indicative of N release from ice features such as rock glaciers worldwide as mountain glaciers retreat.     

The parasite fauna of the fish of the Shropshire Union Canal, Cheshire

A survey was made of the parasites of 603 roach Rutilus rutilus (L.), 15 bream Abramis brama (L.), 201 perch Perca fluviatilis L., 30 pike Esox lucius L. and four eels Anguilla anguilla (L.) caught in the Shropshire Union Canal at Backford, Cheshire from December 1964 to August 1966. Twenty-seven species of parasites were found, 17 in roach, seven in bream, ten in perch, ten in pike and two in the eel. Two species of Protozoa, seven species of Monogenea, five species of Digenea, six species of Cestoda, two species of Nematoda, one species of Acanthocephala, two species of Hirudinea, one species of Crustacea and mollusc glochidia were recorded. The following parasites are believed to be new records for the British Isles: Henneguya oviperda (Cohn, 1895), Dactylogyrus suecicus Nybelin, 1936, D. wunderi Bykhovskii, 1931, Asymphylodora kubanicum (Isaichikov, 1923), and Philometra rischta Skryabin, 1917.
One table shows the species of parasites found, together with data on the site of occurrence in the host, and the percentage and intensity of infection of the fish. A second table compares the occurrence of the parasites in four other localities in the British Isles, Loch Lomond Scotland, Llyn Tegid (Bala Lake), Merionethshire, Rostherne Mere, Cheshire and the River Lugg, Herefordshire. A further column gives the normal hosts in the U.S.S.R.
A limited comparison is made of the numerical occurrence of six species of parasites in the canal, Llyn Tegid and Druzno Lake, Poland. It is concluded that unless relatively large samples of fish of all length groups are collected on a regular basis throughout the year such comparisons will have little meaning.
The concept of the characterization of parasite faunas is briefly noted. It is suggested that the high degree of host specificity shown by many of the species of parasites is evidence in support of the concept.