The reproductive success of pike, Esox lucius: aspects of fecundity, egg density and survival

Power-law relationships have been estimated between fecundity and fish length, weight and age for pike from two gravel pit lakes (Main Lake and St Peter's Lake) in Buckinghamshire, England. Relative fecundity, estimated at 17.6 eggs g⁻¹ for Main Lake pike and 19.4 g⁻¹ for St Peter's Lake pike, did not differ significantly between the lakes. Population fecundity for the Main Lake was estimated at 10.6 million eggs and 6.4 million eggs in 1986 and 1987, respectively.
Experimental work involving samples of eggs from 18 artificially fertilized Main Lake female pike showed a significant relationship between egg diameter and female length, and a significant difference between mean egg diameters for 41-cm and 101-cm females. No significant relationships were found between mean egg size and mean fry size, nor between fry length and adult length. Mean size differences between newly hatched, 18-day-old and 41-day-old fry from the 41-cm and 101-cm females were not significant.
Samples of eggs taken at 2-day intervals from 6 April to 5 May in a pike spawning area of the Main Lake revealed clumped distributions of eggs, with average egg densities ranging from 1 to 47 eggs per 0.071 m² (14–671 m ²) on sand and silt substrata. The maximum egg density of 51 per 0.071 m² (729 m⁻²) was found on flooded grass. The stages of development of the eggs were identified over the sampling period. Survival from stage 1 to fry was estimated at 3–6%. Egg losses were estimated at 9–10% day⁻¹.
Egg survival experiments gave overall hatching success rates of 11 % on sand/silt compared with 2.5% on aquatic plants (1986), and 18.9% on undisturbed sand/silt compared with 7.9% on disturbed sand/silt (1987). The effect of siltation on egg hatching is discussed.     

The production of juvenile Atlantic salmon, Salmo salar in the upper Wye, Wales

The production of juvenile Atlantic salmon ( Sulmo salur L.) was investigated in 16 study sites in the upper Wye catchment during the period February 1975 to November 1976. The population structure was characterised by large numbers of 0+ fish whose abundance decreased with respect to time so that the numbers of each of the older year-classes was a function of the year-class strength of the original fry. The range of parr densities was similar to that recorded for other rivers: the estimates of 2+ smolts did not exceed about 0.04 m⁻². Production ranged from 0.3 to 11.0 gm⁻² a^−l; Of and 1+ fish contributed over 72% and 3+ fish less than 1 % of the total annug production. The relationship between production P (gm⁻² a⁻¹) and mean biomass B (gm⁻²) for all yeatclasses considered separately and in combination can be expressed by the relationship P=a^b . The values for a vary with age-composition and season but the values for b are not significantly different. Differences in growth rate of salmon between sites are small and differences in production result principally from differences in standing crop (biomass density). In most cases low values for salmon production can be attributed to poor spawning.     

Cannibalism in pike fry, Esox lucius L.: some experiments with fry densities

Pike fry were kept in 0.054-m³ tanks at densities of 50, 100 and 150 (277, 555, 833 fry m⁻²) for seven weeks during which the development of social and feeding behaviour was observed. Zooplankton, macro-invertebrates and perch fry were provided sequentially as food; the pike fry were allowed to feed ad libitum.
Zooplanktivorous fry stopped growing at 22 mm whilst, in the presence of abundant suitable alternative prey, 1–4% turned cannibalistic at 5 weeks of age; cannibals subsequently grew rapidly (mean 1.88 mm day⁻¹). Cannibalism ensued in all tanks when the ratio of predator size: prey size was c .2:1.
Fry tended to space-out evenly in the tanks with no overt aggression or territoriality. Behaviour was typified by remaining still for long periods, particularly subsequent to the onset of cannibalism. Cannibals were attracted by fry movements which often initiated attacks.
Daily per capita mortality rates showed no crowding effects before cannibalism but significant density-dependent mortality due to cannibals. Consumption rates of cannibals varied between 0.63 and 6.0 fry per cannibal per day. Cannibals accounted for 54–96% of daily mortality in the experimental tanks. These results are discussed in relation to proposed mechanisms of pike population density regulation.     

Juvenile salmonid production in a Lake Erie nursery stream

Normandale Creek (2531m²) provides spawning and nursery grounds for lake-run rainbow trout, Salmo gairdneri Richardson, brown trout, Salmo frutta L. , and coho salmon, Oncorhynchus kisutch (Walbaum). Upstream movements are significantly correlated with peak stream discharge (P < 0.05). In 1973–1974, 59 adult salmonids constructed 86 nests of which 60% were disturbed by re-use or sand deposition. From a calculated deposition of 90 400 ova, 483 juveniles, 6 cm in fork length ( f.l. ), were resident in the stream when estimates by electrofishing first became valid. Estimated within-stream (i.e. all sections combined) monthly densities of wild juveniles >6 cm f.l. ranged from six to 22 fish (100 m²)⁻¹, and biomass from 1.3 to 3.1 gm⁻². The highest within-section biomass was 8.3 gm⁻² in November. Additions of hatchery-reared rainbow trout temporarily increased monthly within-stream juvenile density up to 23 fish (100 m²)⁻¹, and biomass to 8.2 g m⁻². Density and biomass were positively correlated (P < 0.05) with both instream and bank cover, and biomass negatively correlated (P < 0.05) with gradient. The importance of substratum and flow characteristics on juvenile density and biomass was strongly indicated although not statistically significant. Annual within-section production of juveniles > 6 cm f.l. ranged from 4.79 to 5.93 g m⁻² year⁻¹ in Sections III and IV, respectively. Inclusion of calculated probable numbers of fish < 6 cm f.l. increased within-section production from 5.93 to 18.65 g m⁻² year^−l in upstream Section IV.     

Vertical distribution and abundance of invertebrates within the sandy substrate of a low-gradient headwater stream

SUMMARY. 1 The vertical distribution of invertebrates (>0.053 mm) was studied in a sandy-bottomed, first-order stream on the Coastal Plain of Virginia, U.S.A. Invertebrate species composition, abundance and bio-mass were determined monthly over one year at sediment depth intervals of 0–1, 1–5, 5–15,15–30 and 30–40 cm.
2. The subsurface community was numerically dominated by species of Chironomidae, Nematoda and Crustacea, while much of the biomass was due to early instars of several species of Trichoptera.
3. Invertebrate density and biomass decreased significantly with depth in the substrate (ANOVA; P <0.05). Annual mean density decreased from 1,346,844 individuals m⁻³ at the surface to 13,578 individuals m⁻³ at 15–30 cm. Annual mean biomass decreased from 66.30 g m⁻³ at the surface to 0.44 g m⁻³ at 15–30 cm.
4. Dissolved oxygen decreased markedly from the surface to the 5 cm depth in the substrate, anaerobic conditions often occurring below 10 cm. Density and biomass both showed a significant positive relationship with dissolved oxygen concentration (Linear regression; P <0.05).
5. Physical forces were important in structuring the subsurface invertebrate community. Besides low dissolved oxygen concentration, sediment scouring resulting from storm discharge dramatically reduced density and biomass     

The production of brown trout, Salmo trutta in tributaries of the Upper Wye, Wales

The population density, age structure, biomass, growth and production of brown trout were investigated in four tributaries of the upper River Wye. The populations at each site were largely maintained by immigration from nursery areas. Abundance of separate year classes at sites on the three largest tributaries reached a peak at age 2+. On the smallest stream numbers reached a peak at 1+. Recruitment occurred throughout the year but decreased with age of year classes. Maximum O+ densities ranged from 0.04 to 0.89 m⁻², and >0+ densities from 0.13 to 0.59 m⁻². Average total biomass in 1975 ranged from 2.6 to 14.2 g m⁻². Within the study sites annual trout production in 1975 ranged from 2.9–19.7 g m⁻². Production values were dependent on age structure and population mobility at the study sites. In the three largest streams 2+ and 3+ fish contributed 66.3–88.3% of total production whilst 1+ and 2+ fish contributed 74.5–84.5 % of the total in the smallest stream. The mobile (non-resident) component of the population accounted for up to 60–70% of production at certain times of the year, but over the year (1976) accounted for =30 % of total production. The resident component of the highest annual production value (19.7 g m⁻²) was estimated to be between 15.0 and 18.2 g m⁻².     

The fauna of the muddy sediments of Lough Neagh, with particular reference to eutrophication

SUMMARY. The main groups of animals found in the muddy sediments of Lough Neagh were Chironomidae (Diptera) and Oligochaeta (mainly Tubificidae). At 8 m depth, the average biomass of the main species in these groups was: Chironomus anthracinus , 2.64gm⁻²; Procladius , 342.8 mgm⁻²; tubificids ( Tubifex, Potamothrix, Aulodrilus, Limnodrilus ), 418.7mgm⁻². At 25m depth C. anthracinus and Procladius were less abundant, C. plumosus was present and among the tubificids Limnodrilus had a higher density (up to 50000m⁻²) often an order of magnitude greater than at 8m. Kinnego Bay, a semi-enclosed, polluted area of the lough, had fewer chironomids than the open lough, with chiefly C. plumosus and Procladius. Limnodrilus was more abundant than at 8m. Dero obtusa (Naididae) was also present, sometimes in large numbers (31000m⁻²).
The response of this fauna to eutrophication in L. Neagh appears to be a diminution in the number of species present, with C. plumosus gradually replacing C. anthracinus as the dominant Chironomus , and tubificids, particularly Limnodrilus , becoming more abundant, A further stage, shown in Kinnego Bay, has Procladius as the most numerous' chironomid. Average total biomass at the three sites was 3.9gm⁻² at 8m, 13.2gm⁻² at 25 m in the main lake, 2.8gm⁻² in Kinnego Bay. At 8m depth, C. anthracinus contributed 76% of the total biomass, at 25m Limnodrilus contributed 74% and C. anthracinus 15%, and in Kinnego Bay, Limnodrilus contributed 40%, C. plumosus 28% and Procladius 20%.     

Stocking success of newly hatched pike evaluated by radioactive strontium (85Sr) marking

The stocking of newly hatched pike Esox lucius labelled with radioactive strontium (⁸⁵Sr) and later electrofishing of the same area was found to be a workable tool in assessing the success of pike stockings. ⁸⁵Sr‐labelled individuals constituted 96% of the 0+year pike catch from late summer electrofishing in the stocked littoral areas of the heavily regulated Lokka Reservoir. The density of 0+year pike reached 2·1 individuals 100 m⁻² in the stocked areas in comparison to 0·3 individuals 100 m⁻² in the reference areas. In the three other stocked lakes the proportion of labelled pike in the catch ranged from 0 to 42% in late summer. The results suggest that pike stockings are profitable in filling empty or nearly empty 0+year pike habitats of regulated lakes or other waterbodies, where the natural reproduction of pike has declined due to human activities.     

Radioactive strontium (85Sr) in marking newly hatched pike and success of stocking

Immersion of newly hatched pike fry in an aqueous solution of 1·85 MBq ml⁻¹⁸⁵Sr did not affect their growth or survival over 3 weeks. Cannibal pike fry retained only l.2–3·3% of the ⁸⁵Sr from their labelled prey. Newly hatched ⁸⁵Sr-marked fry stocked into two lakes in 3 years were distinguishable from wild fish 4–6 weeks later. Stocked and wild fry grew equally well, and stocked pike survived best where density of wild pike was highest. There was no evidence of increase in density as a result of Stocking.     

Long-term study of the ecology of wild Atlantic salmon smolts in a small Norwegian river

Backcalculated lengths at the end of the first growth season in wild Atlantic salmon Salmo salar differed significantly between parr smolting at age 1, 2 and 3 years over a period of 11 years (i.e. 1983–1993). Mean body lengths of the respective age groups at the end of the first growth period were 11·1, 6·2 and 4·7 cm, respectively. The mean percentage distribution of fish smolting at age 1, 2 and 3 was 14, 78 and 7%, and the mean smolt age was 1·95 years. Mean lengths at smolting of age groups 1, 2 and 3 were 13·6, 15·8 and 17·5 cm, respectively. Females outnumbered males among the downstream migrating smolts with a mean sex ratio (females/ males) estimated at 1·61, with a significant female surplus in 7 of the 11 years sampled. Of the smolts sampled, 14% exhibited enlarged gonads indicative of parr maturation, and all were males (37% of the parr males sampled). Mean annual smolt density from 1975 to 1996 was 13·4 individuals 100 m⁻² ranging between 0·3–31 smolts 100 m⁻². Mean densities (100 m⁻²) of the smolts aged 1, 2 and 3 years were 1·5, 9·3 and 0·9 fish, respectively. Mean annual biomass for the 22-year period (1975–1996) was estimated at 437 g 100 m⁻², with a range of variation from 136 to 683 g 100 m⁻². Smolt age 2 made up 81% of the mean annual biomass (355 g 100 m⁻²) and smolt age 1 and 3, 8% (35 g 100 m⁻²) and 11% (47 g 100 m⁻²), respectively.     

Population dynamics and feeding of mayfly larvae in some acid and alkaline New Zealand streams

SUMMARY. 1. Population dynamics (density, biomass, annual production), gut contents and feeding rates of mayflies ( Deleatidium spp.; Leptophlebiidae) were compared in two naturally acid (mean pH≃4.8). brownwater streams and two alkaline (mean pH 7.5), clearwater streams in South Westland, New Zealand.
2. Mean densities of larvae (range 234–2318 m⁻²) were higher in alkaline streams on most of the six bimonthly sampling dates. Mean biomass (range 0.020–0.376 g larval dry weight (LDW) m⁻²) was always highest at the stable, spring-fed, alkaline site and was lower at the acid sites and another alkaline site where the population was always dominated by small larvae.
3. Annual production was high at the more stable, alkaline site (10.35 gLDW m⁻²) but much lower at the other sites (2.49–3.77 g m⁻²).
4. Gut contents of larvae were dominated by fine (45–75 μm widest diameter) paniculate matter (69–99%), diatoms (up to 21%) and. at one site, filamentous algae (8–13%).
5. Grazing rates of mayflies on epilithon were significantly higher on stones taken from acid than alkaline streams and material grazed from the former contained a higher proportion of inorganic material (87–93% and 61–83% inorganics, respectively).
6. Higher grazing rates may reflect lower quality of epilithic food in acid, brownwater streams, a factor that could contribute to the lower productivity of Deleatidium populations at these sites.     

Life-history and production of the amphipod Gammarus pulex in a Dorset chalk stream

SUMMARY. The seasonal variation in population density of Gammarus pulex was studied in a Dorset chalk stream. The numbers increased markedly in June and July and reached a maximum of c. 10000m⁻² in September whilst the most rapid decline in density occurred in October-November and reached a minimum of 820 m⁻² in February. The animals occurred in greater densities in habitats containing Ranunculus or Callitriche than in those devoid of vegetation. The population structure was determined monthly and was split into juveniles (length <4mm), immature males, immature females, mature males and ovigerous females. The percentage of juveniles (39–76) was always the highest of any of the categories. Ovigerous females were found at all times of the year. The sex ratio varied with the time of year both for immatures and matures, although there was approximately a 1:1 ratio for the mature individuals. Seasonal variation in biomass showed a maximum of 7.l g dry wt m⁻² in September and a minimum of 1.4 g dry wt m⁻² in March. Production was calculated by two methods giving values of 12.9 g dry wt m⁻² year⁻¹ and 12.8 g dry wt m⁻² year⁻¹.     

Influence of stocking density of European eel (Anguilla anguilla, L.) elvers on sex differentiation and zootechnical performances

Survival rate, sex ratio and zootechnical performances were evaluated on 168 kg of Anguilla anguilla elvers (0.45 g) weaned into three groups (A1, A2, A3) at initial densities of 800, 1600 and 3200 g m⁻³, respectively. In order not to modify the sex ratio, animals weaned at the different densities were maintained separately during the trials, and no size grading was performed at the fattening phase.
Final mean weight achieved by females belonging to the different groups showed no statistically significant differences; weight of males was higher (P < 0.05) in A1 than in A3 where a higher percentage of males was observed. Sex ratio of eels was different among the groups, with a higher percentage of males in A3 (96%) than in A2 (78%) or in A1 (69%). This finding testifies to a sex differentiation strongly affected by pre-fattening stocking density of elvers. Final load showed an increase in males corresponding to a significantly reduced biomass between A1 and A2 (10.97 and 10.24 kg m⁻³) and A3 (8.44 kg m⁻³). Final survival rate ranged from 87% (A1) to 90% (A3). As to food conversion, a better rate was found in A1 (1.9:1) compared to A2 (2.1:1) and A3 (2.3:1) eels.     

An examination of the fish population in the River Trent, Nottinghamshire using angler catches

During this study, anglers caught over 580 kg of fish, comprising over 29 500 individuals of 14 species. Nine times as many gudgeon, Gobio gobio , as roach, Rutilus rutilus , were caught. Gudgeon represented 35.94% of the total catch weight compared with 44.64% for roach. The mean total catch per man-hour was 8.9 fish and the mean catch rate was 176 g per man-hour. Population estimates for fish longer than 12 cm for all species other than gudgeon and bleak, Alburnus alburnus , gave fish densities of 0.21 fish m⁻², the roach population was 0.15 m⁻², and that for dace Leuciscus leuciscus , 0.02 m⁻². The population density for all sizes of all species caught was estimated to be 2.25 fish m⁻².
The fish biomass available to the angler was 447 kg ha⁻². The rate of exploitation of the fishery over the period of sampling was high at 17%, representing an annual exploitation rate of 94%.     

Life history and production of Paranephrops zealandicus in a forest stream, with comments about the sustainable harvest of a freshwater crayfish

1. Life history and production were assessed for the crayfish Paranephrops zealandicus in three reaches of a headwater stream with a catchment of regenerating coniferous-broadleaf forest in the south-east of the South Island of New Zealand.
2. Crayfish density ranged from 3 to 4 m⁻² in riffles and 4–12 m⁻² in pools, depending on reach. Crayfish biomass (4–33 g AFDW m⁻²) and annual production (2–11 g AFDW m⁻²) were high compared with values reported elsewhere, while P:B ratio was low (0.33–0.43). This substantial production was dependent primarily upon high biomass rather than high growth rate.
3. The crayfish of this population rank amongst the longest lived and slowest growing ever recorded. Individuals estimated to be 16+ year of age were not uncommon. Females became reproductively active at 6+ year. Fewer than 4% of females carried eggs, and young remained attached to females for at least 15 months.
4. We propose that characteristics of this population are the consequence of a cool thermal regime (mean daily stream temperature = 7.0 °C, range 1.8–11.9 °C), and that low biomass turnover and poor reproductive rate precludes any sustainable commercial harvest of crayfish from streams in New Zealand with similar thermal regimes.     

Distribution, life cycle and population dynamics of the aquatic enchytraeid Propappus volki (Oligochaeta) in an English chalkstream

Propappus volki was recorded in Dorset chalk streams from gravel substrata where it represented up to 88% of the oligochaete fauna. It was absent from silted sediments associated with marginal and submergent macrophytes. P. volki showed an annual life cycle with a peak of mature individuals in the period April to June, whilst young worms were most abundant after July. The mean monthly biomass in Bere Stream was estimated to be 0.29–0.45 g wet weight m⁻² and the highest mean density was 5465 m⁻² in April 1979.     

Holocene carbon burial by lakes in SW Greenland

The role of the Arctic in future global change processes is predicted to be important because of the large carbon (C) stocks contained in frozen soils and peatlands. Lakes are an important component of arctic landscapes although their role in storing C is not well prescribed. The area around Kangerlussuaq, SW Greenland (66–68°N, 49–54°W) has extremely high lake density, with ∼20 000 lakes that cover about 14% of the land area. C accumulation rates and standing stock (kg C m⁻²), representing late- to mid-Holocene C burial, were calculated from AMS ¹⁴C-dated sediment cores from 11 lakes. Lake ages range from ∼10 000 cal yr  bp to ∼5400 cal yr  bp , and reflect the withdrawal of the ice sheet from west to east. Total standing stock of C accumulated in the studied lakes for the last ∼8000 years ranged from 28 to 71 kg C m⁻², (mean: ∼42 kg C m⁻²). These standing stock determinations yield organic C accumulation rates of 3.5–11.5 g C m⁻² yr⁻¹ (mean: ∼6 g C m⁻² yr⁻¹) for the last 4500 years. Mean C accumulation rates are not different for the periods 8–4.5 and 4.5–0 ka, despite cooling trends associated with the neoglacial period after 4.5 ka. We used the mean C standing stock to estimate the total C pool in small lakes (<100 ha) of the Kangerlussuaq region to be ∼4.9 × 10¹³ g C. This C stock is about half of that estimated for the soil pool in this region (but in 5% of the land area) and indicates the importance of incorporating lakes into models of regional C balance at high latitudes.     

Production of a stream shredder, Peltoperla maria (Plecoptera: Peltoperlidae) in disturbed and undisturbed hardwood catchments

SUMMARY. (1) The average benthic density of Peltoperla maria in an undisturbed southern Appalachian stream was more than twice that of a nearby stream draining a previously clear-cul catchment in its tenth year of natural secondary succession.
(2) Peltoperla production estimates, using three methods, do not show a significant difference in production between streams draining the two catchments. We attribute these results to quicker growth and slightly higher densities of larger nymphs in the disturbed stream. Production estimates for the disturbed stream ranged from 498 to 560 mg (ash free dry weight) m⁻²y⁻¹ while those for the undisturbed stream were 41–4–515 mg m⁻² y⁻¹.
(3) Our results reinforce the view that conclusions based solely upon numerical densities may lead to erroneus interpretations about the roles organisms play in ecosystems.
(4) Annual frass production by this shredder is about 20 times (10 g m⁻² y⁻¹) the secondary production of P. maria.     

Early life characteristics of pike, Esox lucius, in rearing ponds: temporal survival pattern and ontogenetic diet shifts

Survival, biomass and diet of pike, Esox lucius , larvae and juveniles were studied over 3 years by stocking free embryos at a density of six fish m⁻² in 12 drainable outdoor ponds. The ponds were sequentially drained at six larval/juvenile developmental stages, up to a total length (T.L.) of 139 mm. The mean rate of survival at harvest decreased irregularly over time and the highest mortality rates were recorded during the early larval period (13 to 27 mm t.l .) and two intervals of the juvenile period (46 to 99 mm T.L. and 121 to 139 mm t.l .). Mean biomass increased dramatically between 46 mm T.L. (19.8 kg ha⁻¹) and 121 mm T.L. (181.8 kg ha⁻¹) and stabilized between 121 and 139 mm t.l . Sharp increases in the mean weight coefficient of variation were recorded during the early larval period (13 to 27 mm t.l .) and between 74 and 121 mm t.l . (development of cannibalism). Diet breadths were relatively narrow in pike larvae and reached maximum levels in 99 mm t.l . juveniles. Average-sized pike exhibited a sequence of size-dependent shifts from a diet composed primarily (in terms of weight) of micro-crustaceans (at 13 mm t.l .), to chironomid larvae (at 74 to 99 mm t.l .), and finally macrocrustaceans (at 121 to 139 mm t.l .). Cannibalism was detected first among the largest fish at the 74 mm t.l . stage. Between-year diet similarity at various developmental stages was consistently high. In 70 mm t.l . fish harvested from different ponds, we found significant among-pond differences in diet composition; however, similar trends of diet changes in relation to fish size were observed from pond to pond. Our results are discussed in light of existing knowledge of young pike trophic ecology and current aquaculture practices.     

A light intensity threshold for schooling in the Atlantic mackerel, Scomber scombrus

The schooling behaviour of Atlantic mackerel was studied in a large tank at different light intensities in the range 12.6–1.8 × 10⁻¹⁰μEs⁻¹ m⁻². Variable light intensity was produced by accurately controlling the current to a green light-emitting diode (LED) 3 m above the experimental tank. Under high light levels (1.8 × 10⁻⁶μEs⁻¹ m⁻²) mackerel always formed a single school, whereas at lower levels (1.8 × 10⁻⁸μEs⁻¹ m⁻²) they swam as individuals. At light levels down to 1.0 × 10⁻⁶μEs⁻¹ m⁻² the mean nearest neighbour distance in a school remained relatively constant (0.3–0.9 body lengths), and individual mackerel swam along a path which deviated from the position of their nearest neighbours by less than 14°. As light dropped below 1.8 × 10⁻⁷μEs⁻¹ m⁻², both nearest neighbour distance and heading angle between nearest neighbours increased, with mean values of 1–1.8 body lengths and 23–92°, respectively, at 1.8 × 10⁻⁹μEs⁻¹ m⁻². The results are discussed in terms of ambient light conditions in the sea.