The populations of fish in the River Tees system on the Moor House National Nature Reserve, Westmorland

The age, growth, population densities and annual production of an isolated population of brown trout ( Salmo trutta L.) and bullhead ( Coitus gobio L.) were investigated over a 5 year period in Trout Beck in the northern Pennines. Additional data were obtained from two other sites; Great Dodgen Pot Sike and the River Tees.
Trout growth rates were low, particularly in Dodgen Pot Sike, but ages up to VIII were recorded. Bullhead growth rates were similar to those in the Tees at Cow Green (Crisp et al. , 1974), though the Moor House bullheads had a higher survival rate. Population densities of trout ranged from 0.10–0.22/m², with a fry contribution of 0.80%. The bullhead population in Trout Beck was sparse, generally 0.1/m², though an 0 group value of 0.46/m^a was estimated in 1967. At Tees Bridge values ranged from 2.5–7/8/m². Most trout matured by age III, though in Great Dodgen Pot Sike only half of the females were mature at age IV.
Annual production was estimated from Allen graphs. Trout production ranged from 1.02–3.50 g/m²/year, the fry contributing up to 51.5% of the total. A value of 0.48 g/m²/year was obtained for bullheads in the Trout Beck system in a year of good recruitment, whilst bullhead production at Tees Bridge was 7.43 g/m²/year, with fry and I group contributing 48.7 and 37.5% respectively. The high survival rate, irregular recruitment, and poor growth of these high altitude populations is discussed. Growth and reproductive peculiarities of the Great Dodgen Pot Sike trout are also discussed.     

Growth and size of Triturus vulgaris and T. cristatus (Amphibia) in different parts of Norway

The growth and size of the newts, Triturus vulgaris (L.) and T. cristatus (Laurenti) in different parts of Norway (+ Jämtland, Sweden) were studied. Age was estimated from skeletal growth marks, size-frequency histograms and, for the males, from the number of testis lobes.
On average, the T. vulgaris Larvae in central Norway and Jämtland are smaller than those in southeastern Norway, although in good localities, at the same altitude, they are of approximately the same size and complete their development already within 2½–3 months. Larvae living in eutrophic habitats are larger than those in oligotrophic-dystrophic habitats. In bog habitats at similar times of year the T. cristatus larvae from central Norway are about the same size as those from similar habitats in southwestern and southeastern Norway, but somewhat smaller than those from eutrophic ponds in the Oslofjord area.
No significant differences in the mean size of adult T. vulgaris from southeastern Norway and from central Norway were found. The maximum lengths attained were recorded from central Norway, however. In eutrophic habitats T. vulgaris adults BK usually larger than those in oligotrophic/dystrophic habitats.
In southeastern Norway, where growth is rapid, T. vulgaris may become sexually mature when 2+ yr-old, in central Norway usually a year later, in Jämtland. on average, still later, and in their northernmost locality (Vefsn) they probably do not breed until 5+ or 6+ yr-old.
On average, T. cristatus adults from southeastern Norway are slightly larger than those from central Norway, and may become mature at 2+ yr-old. compared with 4 + yr-old, at the earliest, in the latter area. The differences in the growth rates of both larvae and metamorphosed stages, of both species, most probably represent a response to climatic differences, although biotope quality is also an important factor.     

Spatial organization and recruitment of lynx (Lynx lynx) in a re-introduced population in the Swiss Jura Mountains

We monitored seven resident (three males and four females) and six dispersing subadult Eurasian lynx from to in a population that was re-introduced to the Swiss Jura Mountains in the early 1970s. Home-range areas of the neighbouring adults were 71–281 km², and significant core areas 34–252 km². Males occupied significantly larger areas than females. Home-range overlap was 9% for neighbouring males and 3%) for females. Core areas of males did touch, but those of females were clearly separated. Each male's home range covered those of one or two females. Population density was 0.94 lynx/100 km² for resident animals. Pre-dispersal mortality was estimated to be 50%. Juveniles dispersed from their mothers' home area at the age of 10 months. Of six monitored subadults. only one survived the first year of independence. Human-caused mortality (traffic accidents. illegal killing) was high. This was also the case among resident adults. This might be a threat to the long-term survival of the reintroduced population.     

Adult survival and numbers in a coastal breeding population of Redshank Tringa totanus in northwest England

The results of a long-term capture-mark-recapture ringing programme carried out on a coastal population of breeding Redshanks Tringa totanus between 1974 and 1988 are presented.
Both sexes were equally likely to be recaptured in subsequent years, as were birds captured for the first time compared with those that had been captured previously. Older birds were more frequently recaptured than were young birds. There was no significant difference in male and female adult survival rates, with a mean of 72% of females and 75% of males surviving each year.
The breeding population fluctuated annually with estimated breeding densities of 122–285 pairs per km². Variation in breeding numbers (males), but not survival, was partially attributable to winter air temperature.     

The population dynamics of the crested newt Triturus cristatus at a pond in southern England

The population dynamics of a population of crested newts Triturus cristatus were studied in 1983 at a pond in southern England, Three length classes of individuals were distinguished. Adult newts between 67 and 84 mm snout-vent length migrated to the pond in spring, on days when the temperature approached 5°C. Males tended to arrive earlier than females and to remain in the pond for longer. A second period of immigration of smaller adults, between 57 and 70 mm in length, took place in the autumn; these newts overwintered in the water and may have been about to breed for the first time. Juvenile newts measuring between 40 and 51 mm in length also visited the pond during the spring and summer. Immigrant male and juvenile crested newts were significantly higher in weight than emigrants: there was no significant difference between the weights of immigrating and emigrating females.     

ASYMMETRIC VIABILITY OF RECIPROCAL-CROSS HYBRIDS BETWEEN CRESTED AND MARBLED NEWTS (TRITURUS CRISTATUS AND T. MARMORATUS)

Hybridization between divergent lineages often results in reduced hybrid viability. Here we report findings from a series of independent molecular analyses over several seasons on four life stages of F₁ hybrids between the newts Triturus cristatus and T. marmoratus . These two species form a bimodal hybrid zone of broad overlap in France, with F₁ hybrids making up about 4% of the adult population. We demonstrate strong asymmetry in the direction of the cross, with one class ( cristatus -mothered) making up about 90% of F₁ hybrids. By analyzing embryos and hatchlings, we show that this asymmetry is not due to prezygotic effects, as both classes of hybrid embryos are present at similar frequencies, implicating differential selection on the two hybrid classes after hatching. Adult F₁ hybrids show a weak Haldane effect overall, with a 72% excess of females. The rarer marmoratus -mothered class, however, consists entirely of males. The absence of females from this class of adult F₁ hybrids is best explained by an incompatibility between the cristatus X chromosome and marmoratus cytoplasm. It is thus important to distinguish the two classes of reciprocal-cross hybrids before making general statements about whether Haldane's rule is observed.     

Predation and competition within an assemblage of larval newts (Triturus)

The impact of crested newts (Triturus cristatus) on the smaller-bodied palmate and smooth newts (T helveticus and T vulgaris) was studied during the larval stages using a combination of field and laboratory experiments In pond enclosures T cristatus larvae had no effect on the two smaller species over the first four weeks of development By eight weeks, however. T cristatus had achieved a size advantage which enabled it to eliminate T helveticus and severely reduce T vulgaris by predation In laboratory trials under food-limited conditions, T helveticus and T vulgaris were slightly smaller when raised with T cristatus, suggesting that this predatory effect was complemented by interspecific competition during early development Predation of the smaller species started when T cristatus reached a threshold size of c 27 mm No reciprocal effects on T cristatus growth or survival were observed Although T cristatus may be a significant predator of congeneric species in natural ponds, other factors, such as differences in microhabitat selection, higher-order predator-prey interactions and the occasional desiccation of pond habitats may facilitate coexistence between the species     

High survival rate and site fidelity in the Siberian Tit Parus cinctus, a focal species of the taiga

The Siberian Tit Parus cinctus population of Finland, and probably of the whole of Fennoscandia, has declined dramatically during this century. Understanding its population dynamics is essential for its conservation. We studied annual survival rate and dispersal distance of both breeding and fledgling Siberian Tits during 1989-97 in a moderately managed forest habitat near the southern border of its range in Kuusamo, northeastern Finland. Breeding density was low, averaging 0.51 pairs/km in a nestbox area of about 42 km². This was probably an underestimate, because we did not search for nests in natural holes. However, following its population decline, overall densities at the southern border of the range are low. The study area was, however, suitable breeding habitat as reflected by their high nesting success. Brood size at fledging averaged 7.45 young, and reproductive output was 6.16 young per breeding pair, including failed nests. Clutch size adjustment was successful among pairs producing fledglings because, on average, 92.6% of the eggs laid produced young in successful broods. Median distance between consecutive breeding attempts was 100 m for both males and females (range 0–2500 m and 0–6000 m, respectively). Natal dispersal distance was significantly longer in females than in males. We applied Cormack-Jolly-Seber modelling to estimate survival and recapture probabilities separately. Survival of breeding birds was not sex-related, averaging 0.69 annually. Some of the ringed fledglings (3.0%) were later captured as breeders in the area. This is an underestimate of local survival rate due to incomplete recapturing of breeding birds. The implications of these results with respect to the conservation status of the species are discussed.     

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.     

Turnover and recruitment in a tundra population of Peregrine Falcons Falco peregrinus

The population dynamics of tundra-nesting Peregrine Falcons Falco peregrinus tundrius was studied over 7 years on a 450 km² study area in the Keewatin District of Canada's Northwest Territories. Peregrines showed strong fidelity to nest sites; none of 25 males ringed changed territories, while five of 38 females ringed were recorded changing territories. Such changes usually occurred after nesting failure. Annual turnover of territorial adults was estimated to be 22% (15% for males and 26% for females). Annual mortality of adults was estimated to be 17% (15% for males and 19% for females). If we assumed that territory vacancies, in addition to mate replacements, were indicative of mortality, then maximum annual mortality was estimated at 24% for each sex. Territories were held only by adult Peregrines. The oldest male on territory was at least 7 years old, the youngest was 2. The oldest female on territory was at least 7 years old, the youngest was 3. Territories were held by individuals of each sex for at least 6 years. One pair remained together for at least 4 years. Less than 4% of all young Peregrines produced on the study area in the first 5 years of the study were recruited into the breeding population. More male than female young were recruited despite an even sex ratio among nestlings. Peregrines did poorly in their first breeding attempts. The single young female recruited into the study population dispersed more than three times the median dispersal distance of six recruited males, suggesting that other females probably dispersed beyond the boundaries of the study area.     

The effect of pH on feeding behaviour in newt larvae (Triturus: Amphibia)

The feeding responses of three species of newt larvae were compared under circumneutral and sublethal acid conditions. Under acid conditions (pH 4.5) feeding behaviour was suppressed in palmate newts, Triturus helveticus, and smooth newts. T. vulgaris , but not in crested newts, T. cristatus. At low pH, approach and orientation towards food occurred in T. helveticus and T. vulgaris , but snapping was inhibited; T. cristatus snapped and consumed food immediately it was offered under the same conditions. These differences are not consistent with the apparent greater tolerance of T helveticus for acidified ponds. The observations suggest that the chemosensory system of T. helveticus and T. vulgaris may be impaired at low pH.     

The population density, growth rate and production of roach Rutilus mtilus (L.) in Tjeukemeer, The Netherlands

A method of estimating the population density of roach in Tjeukemeer (21.3 km²) using 20 529 introduced fin-clipped fish is described. Fyke nets proved to be an effective method of sampling the population for marked fish during the spawning season. A total of 20 277 roach were processed during the recapturing period. The population density of roach (⋝ 14 cm) was estimated to be 1 246 458. The growth rate of roach in the lake although relatively poor (von Bertalanffy's L ∞ for males and females, 22 and 26 cm respectively) for the species was similar to that recorded in three other Friesian lakes. The mortality rate of males was higher than that of females. Logarithmic length-weight regression analyses showed that the value of the coefficient varied both within and between the sexes, that of females being higher (range 3.03–3.375) than that of males (range 2.76–3.254). Seasonal changes in the size of the coefficient were due to a disproportionate loss of gonad weight in larger fish. The fecundity of the population was comparatively high for the species.
The total production of the population was estimated to be 95 hg ha⁻¹ of which 39 kg ha⁻¹ was contributed by fry. In older (⋝ IV) fish the production of females (12 kg ha⁻¹) exceeded that (2 kg ha⁻¹) of males, due to differences in their growth and mortality rates. The relatively poor performance of roach in Tjeukemeer, in terms of biomass and production was due to a scarcity of zoobenthos and competition from other species offish. There is no evidence either from this study or the literature that the productive potential of roach in lakes is high, even though macrophytes and detritus can be consumed in significant amounts.     

Philopatry and colony fidelity of Shags Phalacrocorax aristotelis on the east coast of Britain

Shag Phalacrocorax aristotelis colonies from the Firth of Forth south to the Fame Islands (SE Scotland and NE England) were surveyed in 1982 and 1983 for nesting Shags ringed as chicks or adults in the previous 20 years. On average, 5% of 863 Shags ringed as chicks were breeding away from their natal colony, and only 1% of 401 marked adults had left their breeding colony. The proportion p of chicks that settled at or beyond distance D from the natal colony followed the relationship p = 0.263D-^0.771, implying that more Shags moved 10 km or beyond to breed than expected from diffusion models or a constant-rate dispersal model; adjusted for birds that settled outside the study area, the relationship was p = 0.314D^-0771. There was no effect of colony size on emigration rate, but females showed a greater tendency to settle away from the natal colony than did males. At the natal colony, more males (69%) than females (47%) nested within 300 m of their birthplace. Two mechanisms appear to determine where a Shag settles to breed: a navigational one governing the return to the natal colony, plus the competitive process of nest-site establishment.     

Studies on feeding, growth and production of a recruited inshore population of Pleuronectes platessa (L.) at East Anglesey, North Wales

Trawl surveys in an area of 237 km² to the east of Anglesey (Irish Sea) indicated a population of recruited plaice ( c . 480000 fish) between 2 and 15 years old. Seasonal growth began in early April and annual growth in the dominant cohorts was analysed for males and females separately. Net annual production was estimated to be 0.17 g m^-2 from a standing crop of 0.3 g m^-2; c . 50% of the production is removed by fishing. The annual consumption (2.5 g m^-2) of Abra and Pectinaria was converted with 6.8% efficiency by weight (8.0% in energy units).     

The forest of Vorsø, Denmark: Succession towards a natural, decidous boreal forest influenced by breeding cormorants

The forest on the strictly protected island Vorsø in Horsens Fjord has since 1952 been examined every ten years to discover the number, basal area, wood volume and aboveground biomass of the species. This paper describes the results from 1982 and the development up to that. The dominating species are Fraxinus excelsior, Ulmus glabra and Fagus sylvatica. The two forests, "Vesterskov" (6.09 ha) and "Østerskov" (2.48 ha) respectively, were, in 1982, still immature. Their number of stems per ha were: 756 and 509; basal area: 38 m² and 45 m². volume: 544 m³ and 615 m³; biomass 207 t and 268 t. The annual increase 1972–1982 was respectively 8.8 m³/ha and 0.3 m³/ha. The impact of the cormorant ( Phalacrocorax carbo sinensis ) is especially in "Østerskov" increasing, since the colony here was established in 1976. The succession of the forest is discussed in relation to the disturbance by the cormorants.     

Population dynamics of two species of hyraxes in the Matobo National Park, Zimbabwe

Populations of the syntopic rock hyrax, Procavia capensis (Pallas), and yellow-spotted rock hyrax, Heterohyrax brucei (Gray), in the 42,400-ha Matobo National Park (MNP), Zimbabwe, were characterized from 1992 to 1996. The Procavia:Heterohyrax species ratio varied significantly ( P < 0.01) across years from 1.43:1.00 to 0.74:1.00. Estimates of abundance (31,114–39,869 Procavia and 21,833–46,619 Heterohyrax ) and densities (0.73–0.94 Procavia ha⁻¹ and 0.51–1.10 Heterohyrax ha⁻¹) indicated that populations had experienced a considerable decline since 1978, with drought suspected as the principal cause. The Heterohyrax population fluctuated more than the Procavia population. Although sex ratios of individuals trapped and collected in 1991–92 did not differ from 1:1 for either species, female Procavia nearly outnumbered males ( P < 0.10) among the prey of black eagles ( Aquila verreauxii Lesson), suggesting the possibility of divergence in the social organization and vigilance of the two species. In each year of the study, pups comprised < 30% of individuals in the two populations after the interspecifically synchronous annual birth event in March–April, and evidence exists for an effect by drought on fecundity. Females appear to breed, on average, no more than every other year. We estimated juvenile (0–1 year of age) mortality at 52.4–61.3% for Heterohyrax and 59.6–75.6% for Procavia . Mean mass (±1 SE) of Procavia adults was 3.4±0.2 kg and of Heterohyrax adults, 2.4±0.1 kg. Procavia had a significantly ( P < 0.01) greater condition index (body mass/total length) than Heterohyrax . Adults are especially important reservoirs of biomass for the diverse and numerous predators of the MNP, accounting for > 84% of hyrax biomass intraspecifically.     

An estimate of the sustainable rate of shell extraction from the Dutch Wadden Sea

1. Shell production by cockles Cerastoderma edule was studied to examine whether or not the present licensed rate of shell extraction in the Dutch Wadden Sea exceeds the current rate of shell addition to the exploitable stocks.
2. Long-term data on numbers of cockles and weights of their shells were used to estimate their annual production on Balgzand, a 50-km² tidal flat area in the western-most part of the Wadden Sea. During the 1969–97 period, it amounted to an average of 125 g m^–2, including 107 g m^–2 of shells large enough to be exploitable for shell-lime fishery.
3. The very irregular annual recruitment of cockles was the main cause of the wide 95% confidence limits (74 and 140 g m^–2 year^–1) of this 28-year estimate. Moreover, high mortality rates in severe winters substantially reduced production per recruit in some year classes.
4. About one-third of the estimated production does not reach exploitable stocks, because it is fragmented by birds (particularly eider ducks), permanently buried in the sediment, or removed by the fishery for live cockles.
5. During the last few decades, the estimated mean amount added annually to the exploitable stocks was 88 million kg or 132 000 m³ of large cockle shells. This amount compares favourably with the current annual level of removal of 134 000 m³ of shells, three-quarters of which are cockles.
6. Even at temporarily lower production rates, the exploitation of shell stocks at its present rate is not expected to lead to a rapid exhaustion of the existing stocks in the tidal inlets of the Dutch Wadden Sea, as these stocks will be in the order of a few million m³.     

The annual number of breeding adults and the effective population size of syntopic newts (Triturus cristatus, T. marmoratus)

Pond-breeding amphibians are deme-structured organisms with a population genetic structure particularly susceptible to demographic threats. We estimated the effective number of breeding adults (Nb) and the effective population size (Ne) of the European urodele amphibians Triturus cristatus (the crested newt) and T. marmoratus (the marbled newt), using temporal shifts in microsatellite allele frequencies. Eight microsatellite loci isolated from a T. cristatus library were used, five of which proved polymorphic in T. marmoratus, albeit with high frequencies of null alleles at two loci. Three ponds in western France were sampled, situated 4-10 kilometers apart and inhabited by both species. Parent-offspring cohort comparisons were used to measure Nb; samples collected at time intervals of nine or 12 years, respectively, were used to measure Ne. The adult population census size (N) was determined by mark-recapture techniques. With one exception, genetic distances (FST) between temporal samples were lower than among populations. Nb ranged between 10.6 and 101.8 individuals, Ne ranged between 9.6 and 13.4 individuals. For the pond where both parameters were available, Nb/N (overall range: 0.10-0.19) was marginally larger than Ne/N (overall range: 0.09-0.16), which is reflected in the temporal stability of N. In line with the observed differences in reproductive life-histories between the species, Nb/N ratios for newts were about one order of magnitude higher than for the anuran amphibian Bufo bufo. Despite of the colonization of the study area by T. cristatus only some decades ago, no significant genetic bottleneck could be detected. Our findings give rise to concerns about the long-term demographic viability of amphibian populations in situations typical for European landscapes.     

Comparative primary productivity of algal epiphytes on three species of macrophyte in the littoral zone of Lake Ohrid, Yugoslavia

Primary productivity of algal epiphytes on the surfaces of Phragmites, Potamogeton , and Nuphar was measured seasonally from June 1978, through June 1979, in the littoral zone of Lake Ohrid, using ¹⁴C methodology. Surface areas of individual macrophytes were determined throughout the study period through the use of a non-miscible surfactant and a calibration curve of surfactant weight versus known, calculated surface areas.
Mean total surface area available for epiphytic colonization during the study period was 1.032 m² macrophyte surface per m² of littoral zone for Phragmites , 0.810 m² for Potamogeton , and 0.167 m² for Nuphar . Seasonal rates of mean primary productivity of algal epiphytes on Phragmites from the surface to the light-compensation depth ranged from 84–1406 mg C m⁻² littoral zone d⁻¹; ranges for epiphytes on Potamogeton and Nuphar were 77–586 and 69–268 mg C m⁻² littoral zone d⁻¹, respectively. Maximum rates were observed typically during June; minimum rates were observed typically during August to December. Mean daily productivity rates over the 12 month period were for epiphytes on Potamogeton 167.0, on Nuphar 100.4 and on Phragmites 671.2 mg C m⁻² littoral zone d⁻¹. Calculated annual production for epiphytes on Nuphar was 36.65, on Potamogeton 60.95 and on Phragmites 245.0 g C m⁻² littoral zone yr⁻¹. Epiphytic production data were typically considerably higher than production data obtained for littoral and pelagial planktonic algae and compare favorably with published data for epiphytic and periphytic production in Lawrence Lake, Marion Lake, and Borax Lake.     

Energy flow through small mammal populations

In order to compile an annual or seasonal energy budget for a small mammal population, it is necessary to derive estimates of standing crop, consumption, egestion, digestion, excretion, assimilation, respiration, growth and reproduction. This review gives an account of the methods available for compiling such budgets, and uses results obtained from a study of Microtus agrestis in south west Britain as a worked example. This population was live–trapped on twenty occasions during the course of 2 years. Total energy flow was 309 kJ/m² in the first year and 173 kJ/m² in the second. The average coefficient of digestibility of the natural diet was 54–3%, the average net production efficiency was 1 –0% and the average gross ecological efficiency was 1 –2%. Peak energy flow occurred in spring and late summer each year, and coincided with peaks in the abundance and growth rate of the grasses which formed the main source of food. Previously published estimates show that the annual flow of energy is higher amongst small mammal populations inhabiting open habitats, such as grasslands, than it is in successionally more mature habitats, such as woodlands.