Short reviews of Nordic publications

By Jonas E. Lawesson Moen, A. 1998. Nasjonalatlas for Norge: Vegetasjon (National Atlas of Norway: Vegetation). Fremstad, E. 1997. Vegetasjonstyper i Norge. (Vegetation types in Norway). Påhlsson, L. (ed.) 1998. Vegetationstyper i Norden (Vegetation types in the Nordic countries). TemaNord 1998.     

Book Reviews

Forty-fifth Annual Report of the Freshwater Biological Association . 1977.
Impact of Acid Precipitation on Forest and Freshwater Ecosystems in Norway . Braekke, F.H. (Ed.) (1976) .
Elliott, J. M. (1977) . Some Methods for the Statistical Analysis of Samples of Benthic Invertebrates .     

A flexible sand coating (Conniflex) for the protection of conifer seedlings against damage by the pine weevil Hylobius abietis

1 A new method for the physical protection of conifer seedlings against feeding damage by Hylobius abietis (L.), is described and evaluated in field trials in Swedish forest plantations.
2 The lower 60% of the stem of the seedling is protected by the Conniflex coating, consisting of fine sand (grain size = 0.2 mm) embedded in an acrylate dispersion that remains flexible after drying.
3 Seedlings are treated in the nursery by a large-scale application procedure involving four steps: (i) spraying the seedlings with water; (ii) application of fixative to the lower sections of the stems, (iii) application of fine sand to the fixative; and (iv) drying of the fixative.
4 A field experiment over three seasons demonstrated a significant increase in survival for coated seedlings compared with untreated seedlings. The survival rate increased from 29% to 97% for Scots pine and from 26% to 86% for Norway spruce. Coating the lower 30% of the stem (instead of 60%) provided inferior protection, resulting in only 64% survival in spruce.
5 Field trials in 11 commercial plantation areas indicated that the Conniflex sand coating was as effective in protecting seedlings as treatment with the insecticide imidacloprid.
6 The new method of coating conifer seedlings with fine sand provides an effective and environmentally sound alternative to insecticide treatment.     

Embryogenesis of Dinocras cephalotes, Perla grandis and P. marginata (Plecoptera: Perlidae) in different temperature regimes

1. Temperature dependence of embryogenesis of the three perlid stoneflies Dinocras cephalots , Perla grandis and P. marginata was investigated by means of incubation experiments. Special emphasis was laid on the effect of fluctuating temperatures and on intraspecific differences between populations from two different field sites in the Swiss prealps (i.e. River Necker and River Thur).
2. Dinocras cephalotes embryos develop between 6.3 and 26.6 °C. The lower threshold temperature is lower than has been reported for more northern populations (i.e. from England and Norway), indicating the existence of a latitudinal gradient. Perla grandis eggs only developed between 9.9 and 18.4 °C. In P. marginata , successful embryogenesis was observed between 9.9 and 18.4 °C, but not enough eggs were available to explore the threshold temperatures.
3. Embryogenesis of D. cephalotes and P. grandis was significantly faster at a 12/16 °C daily fluctuating temperature regime than at a constant 14 °C. However, no significant difference was found between the development under simulated field temperature regimes (with less distinct daily amplitudes) and constant temperatures.
4. D. cephalotes , hatching of eggs from the Necker population was much less synchronous than that in the Thur population. Since the Necker discharge regime is harsher than the Thur regime, it is possible that this asynchrony spreads the risk of destruction during bed-moving floods.     

Embryogenesis of Dinocras cephalotes, Perla grandis and P. marginata (Plecoptera: Perlidae) in different temperature regimes

1. Temperature dependence of embryogenesis of the three perlid stoneflies Dinocras cephalots , Perla grandis and P. marginata was investigated by means of incubation experiments. Special emphasis was laid on the effect of fluctuating temperatures and on intraspecific differences between populations from two different field sites in the Swiss prealps (i.e. River Necker and River Thur).
2. Dinocras cephalotes embryos develop between 6.3 and 26.6 °C. The lower threshold temperature is lower than has been reported for more northern populations (i.e. from England and Norway), indicating the existence of a latitudinal gradient. Perla grandis eggs only developed between 9.9 and 18.4 °C. In P. marginata , successful embryogenesis was observed between 9.9 and 18.4 °C, but not enough eggs were available to explore the threshold temperatures.
3. Embryogenesis of D. cephalotes and P. grandis was significantly faster at a 12/16 °C daily fluctuating temperature regime than at a constant 14 °C. However, no significant difference was found between the development under simulated field temperature regimes (with less distinct daily amplitudes) and constant temperatures.
4. D. cephalotes , hatching of eggs from the Necker population was much less synchronous than that in the Thur population. Since the Necker discharge regime is harsher than the Thur regime, it is possible that this asynchrony spreads the risk of destruction during bed-moving floods.     

Comparison of Campylobacter fla-SVR genotypes isolated from humans and poultry in three European regions

Aims:  The genetic diversity of Campylobacter isolated from human infection and from poultry was assessed in strains originating in three different European regions in order to compare these two hosts and to investigate European regional differences.
Methods and Results:  Randomly chosen isolates originated from Norway, Iceland and Basque Country in Spain were genotyped by sequencing of the short variable region (SVR) of flaA . A total of 293 strains were investigated, c . 100 per country with half originated from either host. The results indicate extensive diversity in both hosts and identified differences in the nature and distribution of genotypes between the countries. These differences could in part be related to geographical location, in that Campylobacter genotypes from Iceland and Norway were more similar to each other than either was to Basque Country.
Conclusions:  Differences between the countries exceeded the observed differences between human and poultry isolates within a country.
Significance and Impact of the Study:  Regional differences are extensive and should not be ignored when comparing genotyping data originating from different international studies.     

Winter herbivory by voles during a population peak: the relative importance of local factors and landscape pattern

1. We studied the relative role of local habitat variables and landscape pattern on vole–plant interactions in a system with grey-sided voles ( Clethrionomys rufocanus (Sund.)) and their favourite winter food plant, bilberry ( Vaccinium myrtillus L.). The study was conducted during a vole peak year (1992–93) in a tundra area in northern Norway.
2. Using Mantel statistics we were able to separate the direct effects of the spatial patterning of habitats and the indirect effects due to spatial aggregations of similar habitats.
3. Results indicate that knowledge about the explicit spatial patterning of patches does not improve our understanding of the system. Instead, two local factors, vegetation height and bilberry biomass, explained more than 50% of the variation in cutting intensity in winter (defined as the proportion of above-ground shoots cut). Increasing vegetation height increased, and increasing bilberry biomass decreased, the cutting intensity.
4. The conclusion that grey-sided voles are able to distribute themselves relative to habitat quality was also partially supported by our estimated over-winter persistence by voles in the various habitats. Vole persistence was uncorrelated with vegetation height, the important predictor of autumn vole density, but tended to correlate with the deviation from the relation between vegetation height and autumn vole density. This conforms to the expectations from the theory of ideal-free habitat distribution.
5. The cue for vole habitat choice, i.e. vegetation height, indicates that either predation or freezing risk is important for voles when selecting over-wintering habitat.     

Laboratory studies of egg development and diapause in Isoperla obscura (Plecoptera) from a mountain stream in Norway

SUMMARY 1. A laboratory study of egg development of the stonefly Isoperla obscura (Zetterstedt) collected from the stream Flybekken (southern Norway, 61°25'N, 8°48'E, 1373 m a.s.l.) showed a short diapause followed by a prolonged period of postdiapause quiescence.
2. Diapause occurred over a wide range of temperatures (−20°C to +8°C), but 0–1°C was the most favourable for fast diapause development and successful hatching. Diapause development required temperatures below 12°C, but sub-zero diapause temperatures increased mortality during the postdiapause phase, as well as reducing hatching success.
3. The threshold for initiation of postdiapausal development was about 1–2°C. Development rate increased and variation in development time decreased at higher water temperatures. These may be important characteristics to ensure seasonal and geographical synchrony of development in individual populations. Low postdiapausal temperatures were unfavourable, despite a subsequent increase in water temperature. No response to postdiapausal photoperiod was detected.
4. The results agreed well with conditions noted in the natural environment, and with the abundance of this species at high altitudes and latitudes in Fennoscandia. Nevertheless, Isoperla obscura also occurs in streams in the maritime parts of western Norway. The results of the present study suggest that egg development in these populations will prove to be different.     

The functional relationship between peak spring floods and survival and growth of juvenile Atlantic Salmon (Salmo salar) and Brown Trout (Salmo trutta)

1. The effects of high spring floods on survival and growth of juvenile Atlantic Salmon, Salmo salar, and Brown Trout, Salmo trutta , are explored, using data from a long-term study in the River Saltdalselv, northern Norway. The flow regime in this river is typical for northern rivers.
2. There was considerable variation in year class strength of both species.
3. Mortality of Atlantic Salmon increased significantly in years with high discharge during the alevin stage as well as the first week after emergence. High discharge during the egg stage and more than 1 week after emergence seemed to be of minor importance. Water temperature at emergence was rather high (average 10·5 °C) and did not significantly affect year class strength.
4. Brown Trout emerged earlier than Atlantic Salmon at an average water temperature of 8·2 °C. Highest mortality was observed in years with low water temperatures at emergence as well as high discharge during the alevin stage.
5. For 1-year-old fish or older, the size of the spring peak flood did not influence mortality significantly.
6. Growth of Atlantic Salmon parr was diminished in years with a high peak spring flood. A similar effect on Brown Trout was not detected.     

Density-dependent melanism in sub-arctic populations of winter moth larvae (Operophtera brumata)

Abstract.  1. The aim of this 4-year observational study was to test for the presence of direct and delayed density-dependent larval melanism in the geometrid moth species Operophtera brumata (winter moth) in northern Norway.
2. Data from many populations with a wide range of population densities in time and space facilitated statistical analyses that could separate the effects of current and past density. The data also included different phases of the 10-year population cycle of this species so that eventual non-linear density effects due to population phase could be detected.
3. The results showed that the prevalence of melanism had a strong positive, linear relation to population density within years, whereas there was no evidence for a delayed effect from the year before or dependency on the phase of the population cycle.
4. In combination, these results limit the range of possible explanations of larval melanism in this outbreaking species. The possible reasons why winter moth larvae might benefit from crowding-induced melanism are discussed.     

Biological activity of aryl hydrocarbon receptor ligands in sediments from remote European lakes

1. Sediment extracts from 63 European remote lakes were tested for the presence of ligands of the aryl hydrocarbon receptor (AhR), using a yeast-based bioassay. The lakes belong to different European mountain regions in Central Norway, Scotland, Piedmont (Italy), Tyrol (Austria and Italy) and the Julian Alps (Slovenia).
2. AhR ligand activity varied about 40-fold among the examined lakes, with Scotland having the highest proportion of lakes with high AhR ligand activity, whereas samples with low AhR ligand activity predominate in the Julian Alps.
3. AhR ligand activity in sediment samples correlated with the concentration of several environmentally relevant compounds, including polycyclic aromatic hydrocarbons, hexachlorobenzene and several polychlorobiphenyl ether congeners. These correlations suggest an anthropogenic origin for the detected AhR ligands, rather than being generated from natural processes.
4. We conclude that pollutants deposited in remote lakes by atmospheric processes are active AhR ligands at the concentrations found in these lakes. Given the high toxicity associated with ectopic activation of AhR in different animal systems, this activity may represent a significant risk for the exposed fauna.     

What limits the altitudinal distribution of Craspedolepta species (Sternorrhyncha: Psylloidea) on fireweed?

Abstract.  1.  Craspedolepta nebulosa and C. subpunctata were studied on their shared host plant Epilobium angustifolium along an altitudinal transect in the Hardangervidda area of southern Norway.
2. The following was investigated: whether (a) the altitudinal distribution and abundance of each psyllid species was determined by summer heat budgets, acting directly on rates of insect development and/or through their host plant and (b) whether parasitism, predation, and interspecific competition played any role in restricting the distribution of the two species .
3.  Craspedolepta nebulosa extended to higher elevations than C. subpunctata but within their ranges neither species showed a clear altitudinal trend in abundance per plant. However, the former species grew to a smaller size with increasing elevation.
4. There was little evidence for interspecific competitive effects and observed levels of parasitism and predation were negligible across the transect.
5. Development and distribution of both species mirrored significant differences in the phenology, size, leaf number, and reproductive output of E. angustifolium with increasing altitude.
6. Distributions were related to available heat budgets acting on psyllid/plant development rates, with C. nebulosa developing more efficiently at lower heat availability.
7. Population transplant experiments supported the overall conclusions, with neither psyllid species able to establish viable populations above their current upper altitudinal range limit, despite the presence of their host plant.     

The distribution of genetic variation in Norway spruce (Picea abies Karst.) populations in the western Alps

Aim  In order to look for a possible centre of survival for the Norway spruce ( Picea abies Karst.) in the south-western Alps, six natural populations of this area were investigated by means of genetic markers in order to assess the degree and the distribution of genetic diversity within the species.
Location  Western and South-western Alps.
Methods  Populations were genotyped using seven simple sequence repeat (SSR) markers. Basic population genetics parameters were estimated and the amount of genetic differentiation calculated.
Results  A large amount of variability was found (0.59 <  H _e < 0.67); genetic differentiation as measured by F _ST was 0.05, close to other similar studies; no isolation by distance was detected by a Mantel test. Analysis of molecular variance confirmed a high degree of variability within populations and a low degree of variability among populations. Finally, the number of populations from which those observed could have arisen was estimated by Bayesian analysis.
Main conclusions  The results presented here suggest that the present populations derive their genetic make-up from three inferred clusters. The possible existence in this area of a relict/refuge population during the last glaciation is discussed.     

Dynamics of a harvested moose population in a variable environment

1. Population size, calves per female, female mean age and adult sex ratio of a moose ( Alces alces ) population in Vefsn, northern Norway were reconstructed from 1967 to 1993 using cohort analysis and catch-at-age data from 96% (6752) of all individuals harvested.
2. The dynamics of the population were influenced mainly by density-dependent harvesting, stochastic variation in climate and intrinsic variation in the age-structure of the female segment of the population.
3. A time delay in the assignment of hunting permits in relation to population size increased fluctuations in population size.
4. Selective harvesting of calves and yearlings increased the mean age of adult females in the population, and, because fecundity in moose is strongly age-specific, the number of calves per female concordantly increased. However, after years with high recruitment, the adult mean age decreased as large cohorts entered the adult age-groups. This age-structure effect generated cycles in the rate of recruitment to the population and fluctuations introduced time-lags in the population dynamics.
5. An inverse relationship between recruitment rate and population density, mediated by a density-dependent decrease in female body condition, could potentially have constituted a regulatory mechanism in the dynamics of the population, but this effect was counteracted by a density-dependent increase in the mean age of adult females.
6. Stochastic variation in winter snow depth and summer temperature had delayed effects on recruitment rate and in turn population growth rate, apparently through effects on female body condition before conception.     

Late autumn eclosion in the winter moth Operophtera brumata: compromise of selective forces in life-cycle timing

Abstract. 1. In eclosion experiments at constant temperatures (6, 9, 12, 15, 18, 21, and 25 °C), Operophtera brumata (L.) pupae were found to respond nonlinearly to temperature, with 9 °C giving the highest developmental rate.
2. Pupal development rate decreased and mortality increased at the highest and lowest temperatures. No pupae eclosed at 6, 21, or 25 °C.
3. Exposing pupae to periods of cold did not enhance their rate of development consistently, indicating that no pupal diapause occurred. Light did not affect the length of the pupal period significantly.
4. Variation in mean developmental rates across temperatures was modelled both for data in the present study (northern Norway) and for a previously published German study (20° further south). The German population had a longer pupal period at all temperatures than the northern population (mean difference of 88 days). The difference is assumed to be genetically based, and hence an adaption to (and not a consequence of) differences in phenology between the two sites.
5. The diversity of the life cycle of O. brumata is discussed. Timing of eclosion can be explained by the selective forces of predation by birds on the one hand and mortality due to early snowfall on the other.     

Life history and habitat use of Norwegian brown trout (Salmo trutta)

SUMMARY. 1. Brown trout ( Salmo trutta) life history and habitat use were studied in two Norwegian rivers: the Vosso river system, western Norway, and the Søre Osa, eastern Norway.
2. Age-groups were partly segregated in feeding habitats, the youngest fish living mainly in running water and in the littoral zone of lakes, the older fish also exploiting pelagic waters and deeper epibenthic habitats. In a population with free access to and from the sea, some individuals smoltified and became sea-run migrants, performing yearly migrations to the coastal sea, whereas others stayed as freshwater residents throughout their entire life span.
3. Within local populations, females were larger and less variable in size than males. This was partly because females matured at an older age than males, partly because the sexes tended to exploit feeding habitats with different food and growth Conditions. Within age-groups, females were more pelagic and migrated more than males, whereas males were more confined to running water and epibenthic areas than females. In the pelagic zone, males were more abundant in near-surface water, and females more abundant in deeper areas. When exploiting the same feeding areas, the two sexes grew at the same rate. There therefore appears to be a connection between feeding habitat and the reproductive ecology of brown trout.     

Genetic differences among brown trout, Salmo trutta, stocks and their importance for the conservation and management of the species

SUMMARY. 1. Review of published studies on genetic variation, as shown by electrophoretic studies of protein variation, in natural brown trout ( Salmo trutta L.) populations from Britain and Ireland, Finland, France, Greece, Iceland, Norway, Sweden, U.S.A. and U.S.S.R., revealed abundant geographical variation in gene frequency with individual populations containing only a limited part of the gene diversity of the species.
2. Thirty-eight (54%) of the seventy gene loci examined have been found to be polymorphic in the species with an average population showing polymorphism at 16% of its loci (range 0-34.8%).
3. The brown trout is naturally subdivided into a large number of reproductively isolated and genetically distinct populations within, as well as among, drainages.
4. Two independent post-glacial colonizations, by genetically distinct races, followed by independent evolution in separate drainages over the past 13,000 years is seen as responsible for the genetic diversity of brown trout in north-western Europe.
5. Many genetically unique populations have been lost in the past 100 years and there is an urgent need to identify and conserve the remaining genetic diversity. Genetically unique populations are an irreplaceable resource for rational management in relation to angling and future aquaculture potential.     

Gallery construction and oviposition of the bark beetle Ips typographus (Coleoptera: Scolytidae) at different breeding densities

Abstract. 1. The bole of a Norway spruce, Picea abies (L.) Karst., was cut in 25 cm sections and infested in the laboratory with 2 or 8 females/dm² of Ips typographus (L.) and an equal number of males.
2. One log of each beetle density was dissected every second day, beginning 2 days after female introduction.
3. The oviposition rate was highest at the beginning of the breeding period for both densities and then declined, especially at the high density. Residence time is a useful approximation for the number of laid eggs in the beginning of the oviposition period or when the density is low.
4. Final gallery length and egg number per gallery were on average significantly smaller at density 8 than at density 2.
5. Linear relationships between total gallery length and number of egg niches were found to be similar for the two densities.
6. An estimate of the egg to adult mortality at different densities was found by using data from a field study that related gallery length to breeding density in attacked living trees, together with published relationships between density and egg niches or emergence of new adults.
7. It was concluded that both decreased oviposition rate and earlier re-emergence at higher densities contribute to a declining oviposition with increasing density. The final offspring production is futher reduced by lower survival due to larval competition as density increases.     

Ceratophyllum demersum hampers phytoplankton development in some small Norwegian lakes over a wide range of phosphorus concentrations and geographical latitude

1. Data on submerged and floating-leafed macrophytes, phytoplankton, nutrients (N, P) and calcium were collected from twenty-four small lakes ( 1 km²) over a wide range of latitudes in Norway. The majority of the investigated lakes were mesotrophic or eutrophic, and most of the lakes were markedly affected by diffuse and point-source runoff from agriculture. According to their macrophyte species composition, the majority of the lakes can be classified as Potamogeton lakes or Chara lakes, or a combination of these.
2. This study is consistent with the 'two alternative stable states' hypothesis. We observed clearwater lakes with dense macrophyte cover over a wider range of total P concentration than has been reported previously: from 30 to more than 700 mg P m^–3. The clearwater state was only observed in lakes with mean depths of less than 1.9 m.
3. Most clear lakes with high cover of submerged vegetation showed indications of N limitation.
4. In this study nearly all the macrophyte-dominated lakes with P concentrations above 30 mg m^–3 had dense stands of Ceratophyllum demersum (L.). This indicates that Ceratophyllum may also play an important role in stabilizing and maintaining a clearwater state at high P concentrations.     

Ceratophyllum demersum hampers phytoplankton development in some small Norwegian lakes over a wide range of phosphorus concentrations and geographical latitude

1. Data on submerged and floating-leafed macrophytes, phytoplankton, nutrients (N, P) and calcium were collected from twenty-four small lakes ( 1 km²) over a wide range of latitudes in Norway. The majority of the investigated lakes were mesotrophic or eutrophic, and most of the lakes were markedly affected by diffuse and point-source runoff from agriculture. According to their macrophyte species composition, the majority of the lakes can be classified as Potamogeton lakes or Chara lakes, or a combination of these.
2. This study is consistent with the 'two alternative stable states' hypothesis. We observed clearwater lakes with dense macrophyte cover over a wider range of total P concentration than has been reported previously: from 30 to more than 700 mg P m^–3. The clearwater state was only observed in lakes with mean depths of less than 1.9 m.
3. Most clear lakes with high cover of submerged vegetation showed indications of N limitation.
4. In this study nearly all the macrophyte-dominated lakes with P concentrations above 30 mg m^–3 had dense stands of Ceratophyllum demersum (L.). This indicates that Ceratophyllum may also play an important role in stabilizing and maintaining a clearwater state at high P concentrations.