Silviculture of the Spruces in West Scotland

An account of the silviculture of Norway and Sitka spruce (Piceaabies and P. sitchensis) practiced in the West (Scotland) Conservancyof the Forestry Commission in the past 50 years. Present methodsare summarized and future developments forecast. A more intensiveform of silviculture is advocated for Sitka spruce.     

A comparison of litterfall and element fluxes in even aged Norway spruce,sitka spruce and beech stands in Denmark

Litterfall was investigated in three even-aged Norway spruce (Picea abies), sitka spruce (Picea sitchensis) and beech (Fagus sylvatica) stands on a nutrient poor-soil in Southern Denmark. Dry weights and N, P, K, S, Mg, Ca, Na, Al, and Fe concentrations and fluxes were examined in litterfall fractions. Foliage litter amounted to 90% of total litterfall. The tree stands showed a similar mean annual litterfall. In the spruce stands, annual litterfall was correlated negatively with the current year increment and positively with the previous year increment. Annual litterfall in beech was constant during the 6 study years whereas Norway spruce and sitka spruce showed large fluctuations between years caused by drought, spruce aphid infestations and probably sea salt stress. Norway spruce responded with a long lasting elevated needle loss. Sitka spruce responded to infestations with premature needle loss during short periods. The presence of a large syrphid (Coccinellidae) population was important in regulating aphid (Elatobium abietinum) population density. The between-year variation in element concentrations of litterfall was small whereas variations during the year were large. Interspecific levels were recognized: Norway spruce>beech>sitka spruce. High concentrations in Norway spruce were ascribed to a combination of drought, sea salt stress and elevated transpiration. In sitka spruce, aphid infestations reduced the litterfall N content. Sitka spruce showed the smallest amount of base cation fluxes with litterfall. In contrast, spruce and beech exhibited even litterfall element fluxes. Litterfall studies revealed reduced vitality in the non-native spruce stands and underlined the perception of a healthy stand of native beech.     

Norway or Sitka Spruce?

The deep and wide continental range and climate occupied bythe European species Picea abies is contrasted with the long,narrow, and humid coastal strip which the American species Piceasitchensis inhabits. Norway is the more plastic spruce and cansuffer thinner soils and much more drought, but neither spruceis a shallow rooter where soils are good and deep. Of the two,the litter of Sitka is less harmful to the soil. To date thereis no evidence that height for height Norway is more stableunder high wind pressures, but the belief that it is less sensitiveto frost than Sitka is well established. Sitka is far superiorto Norway for planting at high altitudes and in exposed placesand on Molinia peatlands in Britain, which accounts for thevast preponderance of the American species used in the westand north of the British Isles. The present popularity of Norwayspruce with foresters and timber merchants is admitted, butthe point is made that in planting it where Sitka could growwell, only some 75 per cent, of the volume will be secured.Finally, the types of site which each spruce requires and alsowhich each cannot take are considered in detail. A plea is madefor mixtures in certain circumstances, and the belief expoundedthat only deeper and better cultivation of the hardpacked materialunder-lying thin soils will make it possible to extend considerablyour acreage of really stable crops of these high-volume producers.     

THE CHECKING OF FOREST TREES BY HEATHER

Experiments on podzolized heather moor at Allerston Forest haveshown that a heather mulch applied around checked Sitka spruce(Picea sitcbensis Carr.), Norway spruce (P. abies Karst.), andLawson cypress (Cbamaecyparis lawsoniana Parl.), or more simplythe elimination of living heather by surface hoeing, producesimproved colour and vigour of the trees. It has also been demonstratedthat the addition of a nitrogenous fertilizer to checked Sitkaspruce can, at least temporarily, alleviate the condition ofcheck.     

Variation of Density and Growth Ring Width in Stems of Sitka and Norway Spruce

The variation with height and radial distance from the pithof basic density and ring width has been determined in stemsof 48-year-old Sitka and Norway spruce planted at two spacingsat Durris, Kincardineshire. The pattern of radial variationof density was broadly similar at all heights: high near thepith, falling to a minimum and then a gradual increase. Themean whole-stem densities were significantly different at thetwo spacings in Sitka but not in Norway spruce. Density at breastheight was inversely correlated with ring width, strongly inSitka but weakly in Norway spruce. The density/ ring width regressionequation for outer wood in Sitka spruce at age 48 differed fromthat at age 31. In Norway spruce a high correlation was foundbetween whole-stem density and the density of rings 16–25at breast height.     

Establishment of the Second Rotation in the Jutland Heath Plantations

The paper describes briefly the silvicultural methods used toregenerate stands of pure Norway spruce (Picea abies Karst.)or Norway spruce mixed with a small proportion of Silver fir(Abies alba Mill.) on the poor heath soils of Jutland. Regenerationafter clear felling, uniform shelterwood, and shelterwood stripfelling is described. The last method has advantages over theother two.     

Fertilization Experiments on Forest Mineral Soils: A Review of the Norwegian Results

The paper gives a brief overview of the background, history and main results of forest fertilization experiments on mineral soils in Norway. Positive results of initial phosphorus (P) fertilization on survival and growth of Norway and Sitka spruce have only been achieved in the coastal districts of western Norway. Other elements have seldom given any significant effect. In young and old stands of Norway spruce and Scots pine nitrogen (N) fertilization with 150 kg N ha -1 usually gave increment increases in the range of 1-2 m 3 ha -1 yr -1 , for a period of 6-8 yrs after application. Given individually, no other element has proved stimulating to stem growth in a similar way. In young Norway spruce stands P has often given additional growth response when given together with N. Liming has been shown to have no or negative effects on tree growth. Fertilization experiments have changed from being management orientated towards addressing the problems of possible nutrient imbalances.     

The chemical composition of needle and leaf litter from Scots pine, Norway spruce and white birch in Scandinavian forests

Needle litter from 14 stands of Scots pine (Pinus silvestris,L.), 13 stands of Norway spruce (Picea abies (L.) Karst.) andleaf litter from three stands of white birch (Betula pubescensEhrh.) were analysed for chemical composition. The concentrationsof the elements N, P, K, Ca, Mg and Mn as well as solid organiccomponents (lignin, cellulose and hemicelluloses) and solubleswere determined. When the average chemical compositions werecompared the Scots pine needle litter was clearly the most nutrient-poorlitter type. Of the solid organic-chemical components the ligninfraction dominated in the spruce and birch litter whereas thecellulose dominated in the pine needle litter. When Norway spruce and Scots pine were growing in adjacent standson soils with the same bedrock origin the spruce litter hadsignificantly higher concentrations of nutrients (N, P, K, Ca,Mg, Mn) than the pine needle litter. At sites where Norway spruceand white birch were growing in adjacent stands, the birch leaflitter had generally higher concentrations of nutrients. However, significant or nearly significant differences were onlyobtained for Mg (P = 0.002), K (P = 0.056) and N (P = 0.087),probably due to the few replicates of stands compared. Concerningorganic chemical components, the spruce needle litter had significantlyhigher concentrations of lignin and mannan than all the otherlitters and lower levels of ethanol-soluble substances, celluloseand galactan than the pine needle litter. Further, it had lowerconcentrations of water solubles, rhamnan and xylan than thebirch litter. No relationships were established between the nutrient statusof the conifer litters and the site index H100 (the dominantheight of the trees at a reference age of 100 years) of thestands. Concentrations of solid carbohydrates in the litterswere, however, positively correlated with site index (P <0.001). Further, the concentration of nitrogen in the pine needlelitter was negatively correlated with the latitude of the sites(P < 0.01). The influence of litter chemistry on the decompositionof litter and nutrient cycling of forests is discussed.     

Mechanical Properties of Picea sitchensis

This study establishes the basis for the assignment of Danish - grown Sitka spruce (Picea sitchensis) to the European strength class system (EN 338). In total, 712 boards were chosen to be representative of the normal Danish commodity. These were visually graded for strength according to Nordic rules (INSTA 142), and subsequently tested to failure in either bending or tension. The results show that Danish - grown Sitka spruce graded to levels T1 and T2 satisfy the requirements of EN 338 at the C18 and C24 levels, respectively. Results for Danish - grown Sitka spruce concur with those for Danish - grown Norway spruce (Picea abies). This study shows that Sitka spruce and Norway spruce of similar origins exhibit highly comparable mechanical properties.     

Stump infection by Armillaria in first-rotation conifers

Information about the entry of Armillaria into first-rotation pine and spruce stands was obtained by searching for infected stumps, rhizomorph systems or trees that had been killed. In pines Armillaria foci were very rare. In pure Norway spruce Armillaria lutea and A. mellea were detected in stumps but rhizomorphs did not extend into the soil; in Norway spruce mixed with oak, by contrast, A. lutea sometimes produced extensive rhizomorph systems. In Sitka spruce small groups of trees had been killed by A. ostoyae. All foci investigated in conifers contained different genotypes of Armillaria and probably originated from spore infection of stumps created by thinning. Some implications of these findings are discussed.     

AN EXPERIMENT ON THE SAMPLING OF EARLY THINNINGS

Several subjective and objective methods of sampling the averagevolume per tree in first and second thinnings were tested instands of Sitka spruce (Picea sitcbensis Carr.), Norway spruce(Picea abies Karst.), Scots pine (Pinus sylvestris L.), Corsicanpine (Pinus nigra var. calabrica Schneid.), Douglas fir (Pseudotsugataxifolia Brit.), and European larch (Larix decidua Mill.).Altogether 7,000 thinnings were measured at Fernworthy in Devon,Thetford Chase in Norfolk, Forest of Ae in Dumfriesshire, andKinellar in Aberdeenshire. The subjective methods, consistingof visual estimates and the measurement of groups of trees selectedas appearing representative, tended to give biased results nomore precise than those obtained by the objective methods whichare unbiased. The objective methods tested were random groups,systematic sampling by trees, and systematic sampling by rows.The systematic sampling by rows was the most satisfactory incombining a reasonable degree of precision with simplicity inexecution.     

Interception of precipitation by pole-stage Sitka spruce and lodgepole pine and mature Sitka spruce at Kielder Forest, Northumberland

Interception loss was measured indirectly for 3 years in adjacentunthinned 25-year-old stands of Sitka spruce and lodgepole pineand for 2 years in a 63-year-old selectively thinned stand ofSitka spruce. For each stand measurements were made of grossprecipitation, throughfall and stemflow. Interception loss wasderived by subtracting the sum of throughfall and stemflow (netprecipitation) from gross precipitation. For the years 1977–78, 1978–79 and 1979–80,interception losses in the pole-stage Sitka spruce were 32,28 and 27 per cent of gross precipitation respectively whilethose in the lodgepole pine were 33, 29 and 26 per cent. For1979–80 and 1980–81 interception losses in the matureSitka spruce were 44 and 53 per cent of gross precipitation.The average interception loss was 29 per cent for both pole-stagecrops and 49 per cent for mature Sitka spruce. The proportions of net precipitation reaching the ground asstemflow and throughfall were 0.18 and 0.82 respectively forthe pole-stage Sitka spruce, 0.14 and 0.86 for the pole-stagelodepole pine and 0.02 and 0.98 for the mature Sitka spruce.     

Growth and Wood Properties of Sitka Spruce (Picea sitchensis (Bong.) Carr.) in Nursing Mixtures Established on Nitrogen-deficient Mineral Soils

Rate of growth and wood properties of Sitka spruce (Picea sitchensis (Bong.) Carr.) were investigated in triplet mixtures with lodgepole pine (Pinus contorta Loud.) (Alaskan provenance) and Japanese larch (Larix kaempferi (Lamb.) Carr.), established on nitrogen-deficient, iron pan soils. These ?nursing? mixtures were compared with pure stands of Sitka spruce that had been either regularly or periodically fertilized with nitrogen. Japanese larch promoted a growth rate in Sitka spruce greater than that achieved in the lodgepole pine-nursed spruce and equivalent to the two pure spruce treatments over the duration of the experiment (current age 28 yrs). Growth of regularly fertilized pure Sitka spruce was not significantly greater than that of periodically fertilized pure spruce. Alaskan lodgepole pine controlled branch size on the lower part of the spruce stems more effectively than the other treatments, although this may have been a function of tree size. Branch characteristics of Japanese larch-nursed spruce, however, were similar to those of the pure spruce treatments. Japanese larch caused an imbalance in crown development in the spruce, although it is unclear from the present study whether this will have an influence on stem and wood quality by the end of the rotation. Overall, the evidence from this study suggests that Japanese larch is an effective nurse of Sitka spruce on nitrogen-deficient iron pans, maintaining a rate of growth similar to that of pure Sitka spruce periodically fertilized with nitrogen and higher than that observed in spruce nursed by Alaskan lodgepole pine.     

Fomes annosus Butt-Rot of Sitka Spruce IV. Observations on the Distribution of Cryptosporiopsis abietina (Petrak) in the Stems of Rotted Trees

Cryptosporiopsis abietina was isolated from the heartwood of12 out of 18 Sitka spruce suffering from butt rot caused byFomes annosus, but from none of eight healthy trees from thesame plantations. C. abietina was present in a distinct zoneabove the tissue colonised by F. annosus. It was present inboth sound wood, and in wood with stain or incipient decay. C. abietina inhibited the growth of F. annosus on malt agar,and to a lesser extent in autoclaved Sitka spruce heartwood.The possible influence of this potentially antagonistic funguson infection by F. annosus is briefly discussed.     

CONIFEROUS LITTER AMENDMENTS AND THE GROWTH OF SITKA SPRUCE

The growth of semi-checked Sitka spruce (Picea sitcbensis Carr.)on heathland has been stimulated to different degrees by theannual application of litter of Scots pine (Pinus sylvestrisL.), Corsican pine (P. nigra var. calabrica Schneid), Lodgepolepine (P. contorta Dougl.), Japanese larch (Larix leptolepisGord.), and Sitka spruce, at rates corresponding to normal plantationconditions, following an initial heavier rate. Foliar analysissuggests that the response is at least partially attributableto the influence of the litters on the nitrogen nutrition ofthe spruce and differences between litters appear to be largelydetermined by their total nitrogen contents. In this respect,Japanese larch, with an annual needle fall generally greaterthan that of the pines and a higher nitrogen content, offersthe best promise as a nurse species for spruce.     

Movement of Injected Dyes in Gymnosperm Stems in Relation to Tracheid Alignment

The path of upward movement of acid fuchsin dye was traced inpole-size stems of Japanese larch (Larix leptolepis Murr.),European larch (Larix decidua Mill.), Lawson cypress (Chamaecyparislawsoniana (A. Murr. Parl.)), Sitka spruce (Picea sitchensis(Bong. Carr.)), Norway spruce (Picea abies (L.) Karst.), andScots pine (Pinus sylvestris L.). In all of these species amarked, preponderantly clockwise spiral pattern of dye ascentwas observed. In general the upward movement of the dye followedtracheid alignment, except at the sapwood-heartwood boundary.The characteristic patterns of dye movement indicated that spiralgrain was much commoner than was straight grain among the pole-sizegymnosperms investigated. Large differences in spiral grainwere found among species, among trees within species, in differentannual rings within trees, and at different stem heights. Patternsof dye uptake were influenced somewhat by the method of injection.Dye injections through tubes driven into the stem were foundmore suitable for determining tracheid arrangement than wereinjections through individual roots immersed in dye. Factorsinfluencing spiral grain development and applications of thedye injection technique to spiral grain determination are discussed.     

Capercaillie habitat, diet and management in a Sitka spruce plantation in central Scotland

The ancestral habitat of capercaillie in most of its range isScots pine forest with an understorey of blaeberry. Keillourforest in Perthshire is dominated by Sitka spruce, has littleblaeberry, yet had a high density of capercaillie. Food andhabitat use by capercaillie were studied there from 1991 to1994. Seven cocks and two hens were caught, and marked withsmall radio transmitters. The habitats they occupied and foodremains in their droppings provided information on home range,habitat utilization and diet through the year. Sitka sprucewas the main food in autumn, winter and spring but was equalledby the fruiting heads of sedge in summer. Larch was the secondmost abundant item in spring and autumn diets. Sedge was plentifulon the wet restocks associated with the gleyed soils at Keillourand may provide a partial substitute for blaeberry in summer.Capercaillie showed a clear preference for the oldest blocksof trees, especially in winter. Sitka spruce, Norway spruceand Scots pine were used roughly in proportion to their availabilityin the birds' home range, but larch stands were used more thanexpected from availability. In summer, all birds used restocksto some extent. Management recommendations to encourage capercailliein commercial plantations largely comprising exotics are thatmature stands over 45 years and well-vegetated restocks shouldeach occupy at least 20 per cent of the forest, larch and blocksof mixed conifers each at least 15 per cent of the ground. Blaeberryshould be encouraged and, if possible, patches of windthrowinvolving living trees with the main root intact should be retained.     

Annual and spatial variation in coning shown by the Forest Condition Monitoring programme data for Norway spruce, Sitka spruce and Scots pine in Britain

Conifer seed is a valuable forest resource; as well as beingreproductive material, it is an important foodstuff for manywoodland species. Information on cone and therefore seed productionof the main forest species are useful in the fields of bothforest management and forest ecology. Coning has been recordedsince 1989 in plots of Norway spruce, Sitka spruce and Scotspine across Britain as part of the Forestry Commission's ForestCondition Monitoring programme. An 11-year period of these recordswas analysed and a summary of annual cone production by speciesand coning synchrony is presented. Both Norway spruce and Sitkaspruce coned sporadically, with a 4-year period between mastcrops during which cone density was very low or cone productionabsent, whereas Scots pine produced a good cone crop in mostyears throughout Britain. Coning was synchronous within andbetween Norway and Sitka spruce across the whole of Britain(at distances >600 km) but coning within Scots pine was insynchrony only over distances of up to 200 km. Synchrony ofcone production was not evident between the spruce species andScots pine. These findings have implications when planning andinterpreting monitoring used to predict cone crops and in thedesign of forests for wildlife conservation.     

Relationships between insect diversity and habitat characteristics in plantation forests

A methodological approach to the identification of biodiversity indicators in commercial forest stands is illustrated by analysis of the relationships between syrphid (hoverflies) and carabid (ground beetles) community composition and diversity, and stand structure and field layer vegetation. Data were collected from 12 commercial forest sites encompassing a range of climatic conditions and different crop types (Scots pine, Sitka spruce, Norway spruce and Corsican pine) across the UK. Comparisons were also made between unmanaged semi-natural Scots pine woods and Scots pine plantations. For both syrphids and carabids, no differences in species richness and diversity were recorded between semi-natural stands and plantations; one rare syrphid considered to be restricted to semi-natural pine stands was also found in spruce. Syrphid species diversity and richness was higher in southern spruce sites than in the northern sites. Northern sites had distinctive carabid communities, as did sites in the New Forest, a large ancient woodland in southern England. Of the measured habitat variables, vertical stand structure showed the best correlation with species richness and diversity of both carabids and syrphids. Richness and diversity were less in stands with high vertical cover values for canopy layers. Stands with higher field layer cover supported greater syrphid diversity, but lower carabid diversity. Measures of stand structure could be used as potential indicators of syrphid and carabid diversity, but additional habitat parameters also need to be tested.     

Sustainability in multi-aged stands: an analysis of long-term plenter systems

Long-term research plots in multi-aged stands managed with theplenter system were assessed to evaluate sustainability of theplenter system in Central Europe. Plots primarily consistedof Norway spruce (Picea abies (L.) Karst., silver fir (Abiesalba Mill.) or European beech (Fagus sylvatica L.) and weremeasured for seven to 16 measurement intervals over 60–91years. Sustainability was assessed with four types of criteria:stand density, tree species diversity, basal area increment,and stand structure. Comparable even-aged stands were also analysedto compare and evaluate the performance of the measures of sustainability.Measures of species diversity, increment and stand structuraldiversity generally experienced increasing trends over timein these even-aged stands. Basal area generally increased andtrees ha^-1 decreased in multi-aged stands following similarpatterns as in even-aged stands. These results suggest thatthe plenter system is still evolving and is not the model ofsustainability often assumed. Many of the measures used havepotential as indicators of sustainability in multi-aged stands.