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.     

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.     

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Published and unpublished accounts of the zooplankton community of Loch Lomond are reviewed.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     

A multivariate morphometric and meristic description of a population of freshwater-feeding river lampreys, Lampetra fluviatilis (L.), from Loch Lomond, Scotland

Adults of a freshwater-feeding population of Lampetra fluviatilis (L.) in Loch Lomond, Scotland, were trapped on their upstream (spawning) migration in the main inflow to the loch, the Endrick Water. These animals could be distinguished from the more typical anadromous upstream migrant L. fluviatilis , from the Endrick Water and the nearby catchment of the River Forth, principally by their smaller total length, greater disc, snout and eye size and much darker pigmentation. These, and other morphometric and meristic characteristics have been compared for populations of L. fluviatilis and Lampetra planeri (Bloch) from the Lomond and Forth catchments using univariate and multivariate statistics. The Endrick Water 'dwarf' L. fluviatilis was found to differ significantly from the other Lampetra studied, in a number of body proportions in addition to those traits mentioned above, and also in mean numbers of trunk myomeres and teeth. These differences are likely to be due to a combination of the effects of genetic isolation, and the short (freshwater) feeding phase of the 'dwarf' L. fluviatilis , which may, therefore, represent the intermediate stage between parasitic and non-parasitic forms in the L. fluviatilis/L. planeri species pair.     

Confirmation of the presence of schelly, Coregonus lavaretus, in Brotherswater, U.K.

The presence of the nationally-rare schelly. Coregonus lavaretus , in Brotherswater, a small lake in the English Lake District, was confirmed in the summer of 1992. However, pronounced deepwater oxygen depletion may pose a threat to the continued survival of this seventh U.K. population.     

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.     

A hermaphrodite specimen of Coregonus lavaretus (L.) (Salmoniformes, Salmonidae) from Loch Lomond, Scotland

A single hermaphrodite specimen of Coregonus lavaretus (L.) was caught mid-January in Loch Lomond, Scotland.     

Ultrastructure of the retina of two subspecies of Coregonus lavaretus (Teleostei) from Lake Constance (Germany)

In the two studied subspecies of Coregonus lavaretus , the pollan ( C. l. wartmanni ) (which lives deep in the pelagial) and the gangfish ( C. l. macrophthalmus ) (which lives near the slope, closer to the bottom), duplex retinae containing rod and cone photoreceptors are found. Four morphologically different cone types were observed: unequal double cones, short single cones, long single cones and triple cones. The cones are arranged in a square pattern (four double cones around a central short single cone) in the ventral and ventrotemporal and in a row pattern in the nasal and dorsal areas of the retina. Moreover, intermediate patterns can be observed in several regions indicating that double cone twisting occurs, i.e. double cones twist about their longitudinal axis. The highest cone densities are found in the ventrotemporal area. Conversely, the rod photoreceptor density is the highest in the dorsal retina. While the basic morphology of the retina is the same in both subspecies, the distribution of long single and triple cones differs between the studied animals. While these cone types are very rare in the pollan, they are common in the gangfish, though not exhibiting a regular pattern. The findings are discussed with regard to the photic habitat conditions, the systematic position of coregonids and variation of retinal morphology in the two subspecies.     

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.     

Radioactive strontium (85Sr) in marking whitefish [Coregonus lavaretus (L.)] larvae and the dispersal of larvae from river to sea

Anadromous whitefish [ Coregonus lavaretus (L.)] yolk-sac larvae were marked with a radioactive strontium isotope, ⁸⁵Sr. Before marking the method was tested in accumulation-elimination tests using various strontium concentrations. The first-phase effective half-life of ⁸⁵Sr in yolk-sac larvae was 4 days. The mean ⁸⁵Sr radioactivity was 81 Bq per larva at the day of release and the half-life of the slow component was 51 days, so the strontium label was easily detected up to 60 days after marking. The larvae were released into the River Simojoki about 12 km upstream from the river mouth and recaptured from early May to the end of July. Marked larvae were found at the river mouth 12–13 days after release, while the last marked fish in the river were found at 16 days. It is likely that all leave the river within 2 to 3 weeks post-hatching. Two months after marking some larvae had attained a distance of 10 km from the river mouth.     

Environmental effects of introducing whitefish,Coregonus lavaretus (L.), in Lake Ring

The impact of whitefish (Coregonus lavaretus (L.)) on the trophic structure of eutrophic lakes was studied in Lake Ring, a small eutrophic Danish lake (22.5 ha, mean depth 2.9 m) in which the natural fish fauna is dominated by pike (Esox lucius L.), perch (Perca fluviatilis L.), and eel (Anguilla anguilla (L.)), roach (Rutilus rutilus (L.)) and burbot (Lota lota (L.)) being the only other fish species present. A total of 10993 0+ whitefish were stocked in the lake from October 1989 to July 1990 and the structure of the fish, zooplankton and benthic invertebrate communities studied during the period 1989–1991. Stomach contents analysis revealed that the whitefish mainly ate Daphnia and copepods in 1990–1991, the proportion of copepods decreasing with increasing size of the fish and Daphnia being the overall most important food source. The density of Daphnia in the lake decreased from 72 ind. 1^-1 in 1989 to 9 ind. 1^-1 in 1991; concomitantly the large species Daphnia magna and D. pulex almost disappeared and the density of cyclopoid copepods increased from 72 to 101 ind. 1^-1, presumably because of improved food conditions, while that of calanoid copepods remained virtually unchanged. As a result chl-a increased from 19 to 47 µg 1^-1 and Secchi depth decreased from 2.4 m to 1.7 m, despite there being no change in total P and total N (0.6 mg P 1^-1 and 1.3 mg N 1^-1, respectively). Changes were also observed in the benthic invertebrates; Chaoborus, oligochaetes, and chironomids all decreased, whereas Pisidium increased. It is concluded that the stocking of whitefish in eutrophic lakes for commercial purposes may delay their recovery, or even lead to enhanced eutrophication.     

Survival, growth and feeding of vendace, Coregonus albula (L.), larvae in net enclosures

Vendace, Coregonus albula (L.), larvae reared in net enclosures anchored in the littoral zone of Lake Ylä-Enonvesi, eastern Finland, were observed for a period of 44 days in order to determine the effects of density on mortality, growth and feeding of the larvae. The ratio of newly-hatched fish to their food was regulated by controlling the number of fish stocked into the enclosures. Prey animals were expected to move freely in and out of the enclosures. The effect of nylon mesh (500 μm) on the penetration of zooplankters into the enclosures was negligible. The influence of possible starvation was determined by analysing the growth of the larvae, their Fulton condition factor, the instantaneous amount of food ingested, and the food composition in their guts. All these indicators were inversely correlated with larval density. Vendace larvae were found to be resistant to mortality from direct starvation. At all densities the survival rate was over 60%. At the time of complete yolk absorption, there was no increase in mortality.     

Continuous variation in the pattern of marine v. freshwater foraging in brown trout Salmo trutta L. from Loch Lomond, Scotland

Carbon stable-isotope analysis showed that individual brown trout Salmo trutta in Loch Lomond adopted strategies intermediate to that of freshwater residency or anadromy, suggesting either repeated movement between freshwater and marine environments, or estuarine residency. Carbon stable-isotope (δ¹³C) values from Loch Lomond brown trout muscle tissue ranged from those indicative of assimilation of purely freshwater-derived carbon to those reflecting significant utilization of marine-derived carbon. A single isotope, two-source mixing model indicated that, on average, marine C made a 33% contribution to the muscle tissue C of Loch Lomond brown trout. Nitrogen stable isotope, δ¹⁵N, but not δ¹³C was correlated with fork length suggesting that larger fish were feeding at a higher trophic level but that marine feeding was not indicated by larger body size. These results are discussed with reference to migration patterns in other species.     

The relationship between Trichocerca pusilla (Jennings), Aulacoseira spp. and water temperature in Loch Leven,Scotland, U.K.

Loch Leven is a shallow, eutrophic lake in the Scottish lowlands that is famous for its brown trout (Salmo trutta L.) fishery. Studies of planktonic rotifer populations began here in January 1977. Since then, samples have been collected and analysed at more or less weekly intervals. Additional information on the composition and abundance of phytoplankton and crustacean zooplankton species, and on a variety of physical and chemical determinants, has been recorded on each sampling occasion.Long-term datasets, such as that described above, are invaluable for identifying interactions between components of the plankton that only appear for short periods each year, as these interactions would probably be overlooked in data spanning a shorter period of time. This study uses the long-term data from Loch Leven to examine the food and temperature requirements of the summer rotifer species Trichocerca pusilla (Lauterborn). The results suggest that T. pusilla prefers water temperatures above 12 °C and that it feeds, primarily, on the filamentous diatom Aulacoseira spp. During the summer months, its abundance was closely related to the availability of this diatom. When filaments of Aulacoseira spp. were abundant, rotifer densities reached 1000–3000 ind. l^–1 and when they were scarce (e.g. 1980, 1997 and 1998) T. pusilla densities also remained low (i.e. less than 100 ind. l^–1). The reason for the success or failure of Aulacoseira during the summer months each year is unclear but, in general, its abundance was related to the availability of dissolved silica in the water.     

The genetics of sympatric Arctic charr [Salvelinus alpinus (L.)] populations from Loch Rannoch, Scotland

Two morphologically and ecologically distinct forms of Arctic charr, Salvelinus alpinus , are found in Loch Rannoch, Scotland. The differences in morphology are adaptations to different modes of life, one being pelagic, the other benthic. Both forms have been the subjects of extensive genetic studies including cytogenetics, nuclear and mitochondrial DNA analysis, and protein electrophoresis. Significant differences between the two forms are revealed by some techniques but not others and provide evidence for the reproductive isolation of these two morphs. The findings are discussed in relation to the derivation of sympatry and the phylogenetics of Arctic charr.