The influence of compound fertilizer and cupric sulfate on the growth and the bioelement content of cembra pine seedlings (Pinus cembra)

Summary It was the purpose of this study to investigate the effect of compound fertilization and compound fertilization with cupric sulfate on the growth and the bioelement content of cembra pine seedlings. At the same time microbiological investigations of the soil were carried out under the same conditions.Five months after fertilization the seedlings of the plot treated with compound fertilizer showed lower total weights and lower bioelement contents in their organs than the seedlings in the untreated control plot and in the plot treated with copper.In addition to these results it was found that in the plot treated with compound fertilizer there was an increase in the activity of those microorganisms which are probably nutrient competitors to the cembra pine seedlings whereas the plot treated with cupric sulfate showed a decrease in the activity of these microorganisms. The addition of cupric sulfate caused a partial elimination of this competition because of the toxic properties of the heavy metal and enabled the young cembra pine plants to increase their uptake of nitrogen, phosphorus and other bioelements.In the control plot and in the plot treated with copper beginnings of mycorrhiza were observed. It could not be proved that the results were influenced by weed competition.These observations have contributed to explaining the results of previous experiments which have always shown that fertilizers containing cupric sulfate favour the growth of cembra pine seedlings.     

A survey of insects attacking seed cones of Pinus cembra in the Alps, the Pyrénées and Massif Central

The composition and impact of the entomofauna exploiting seed cones of Swiss stone pine ( Pinus cembra L.) was surveyed in native stands and plantations of France, Italy, Switzerland and Austria during 1992–1996. A total of 2785 cones were collected from 25 sites. The cone entomofauna consisted of five species, of which three ( Dioryctria abietella , Eupithecia abietaria , Cecidomyia pini ) were regularly observed. The two other insects, Polydrusus atomarius and Zeiraphera diniana were observed on first-year cones, and only in two stands. No spermatophagous insect was detected. None of these insect species could be considered specific to cones of P. cembra. The overall damage never exceeded 40% of the cone crop. The importance of cone damage varied with year, cone abundance, and the position of the tree in the stand. The cone entomofauna of Swiss stone pine largely deviated from that recorded in cones of other high altitude conifers but resembled that of other stone pines growing in Eurasia. The question: why cones of P. cembra were less damaged by insects than those of other alpine conifers is discussed.     

Maples at the sub-Alpine vegetation belt: a long history

Pollen analysis was carried out on lacustrine sediment of a small hollow (15 m x 25 m) at the treeless sub-Alpine belt (202 m) of the inner Maurienne valley in the northern French Alps. A 2,500-year-long maple settlement was demonstrared. Three AMS dates of terrestrial plant macroremains support the chronology. First, Betula and Salix spread prior to 9,000 C14 BP. The first pollen grains of Acer, Abies and Pinus cembra are quoted at 8,600 C14 BP. High frequencies of Alnus glutinosa/incana (20%) and Acer (10%) show that mixed communities of Acer and Alnus persisted above the mountainous Abies forest between 7,490 and 5,850 C14 BP. After 5,850 C14 BP, the decrease in Acer stands could be attributed to fire as suggested by the strong increase in Betula and by the delayed expansion of Pinus cembra.     

Chloroplast microsatellites and mitochondrial nad1 intron 2 sequences indicate congruent phylogenetic relationships among Swiss stone pine (Pinus cembra), Siberian stone pine (Pinus sibirica), and Siberian dwarf pine (Pinus pumila)

We studied the phylogenetic relationships among the three stone pine species, Pinus cembra, P. sibirica, and P. pumila, using chloroplast microsatellites and mitochondrial nad1 intron 2 sequences. The three chloroplast microsatellite loci combined into a total of 18 haplotypes. Fourteen haplotypes were detected in 15 populations of P. cembra and one population of P. sibirica, five of which were shared between the two species, and the two populations of P. pumila comprised four species-specific haplotypes. Mitochondrial intron sequences confirmed this grouping of species. Sequences of P. cembra and P. sibirica were identical, but P. pumila differed by several nucleotide substitutions and insertions/deletions. A repeat region found in the former two species showed no intraspecific variation. These results indicate a relatively recent evolutionary separation of P. cembra and P. sibirica, despite their currently disjunct distributions. The species-specific chloroplast and mitochondrial markers of P. sibirica and P. pumila should help to trace the hybridization in their overlapping distribution area and to identify fossil remains with respect to the still unresolved postglacial re-colonization history of these two species.     

Similar Susceptibility to Excess Irradiance in Sun and Shade Acclimated Saplings of Norway Spruce [Picea Abies (L.) Karst.] and Stone Pine (Pinus Cembra L.)

Photosynthetica - We compared the responses of sun and shade acclimated saplings of Picea abies and Pinus cembra to excess photosynthetic photon flux density (PPFD) equivalently exceeding the level...     

Allozyme polymorphism of Swiss stone pine Pinus cembra L. in mountainous populations of the Alps and the Eastern Carpathians

Swiss stone pine Pinus cembra L. is a species with fragmented range, occurring in the Alpine-East Carpathian mountain system. Seeds of P. cembra are dispersed by nutcrackers, which offers potential possibilities for gene exchange among populations. Using isozyme analysis, we have examined five samples from two parts of the Swiss stone pine range: the Alps (Switzerland and Austria) and the Carpathians (two samples from the northern macroslope of the Gorgany Ridge, Eastern Carpathians, Ivano-Frankovsk oblast and one sample from Trans-Carpathian oblast of Ukraine). The allele frequencies of 30 isozyme loci, coding for enzymes ADH, FDH, FEST, GDH, GOT, IDH, LAP, MNR, MDH, PEPCA, 6PGD, PGI, PGM, SDH, SKDH, SOD, were analyzed using cluster analysis and methods of principal components. Two clusters, corresponding to the isolated Alpine and Carpathian parts of the range, were found. The main contribution to these differences were made by loci Adh-1, Adh-2, Fest-2, Lap-3, Mdh-4, and Sod-4. The interpopulation differentiation proved to be somewhat higher than that typical for pines (F(ST) = 7.4%), but within the limits characteristic for taxonomically close species. Thus, isolation of the populations did not lead to their marked differentiation, which may be explained by gene flow and balancing selection, which equalizes gene frequencies across the fragmented species area. Interlocus F(ST) heterogeneity (from 0.003 to 0.173) suggests adaptive significance of some of the allozyme polymorphisms or linkage of some loci with adaptive genes. The Carpathian populations were shown to have higher gene diversity than the Alpine ones (expected heterozygosities 0.095-0.114 and 0.060-0.080, respectively). A deficit of heterozygotes (as compared to the Hardy-Weinberg proportions), observed in the embryo sample, was probably explained by inbreeding. The reduction in the area of Carpathian pine forests in Holocene, caused by the global climatic changes and the anthropogenic impact, is hazardous for the gene pool of the species. The maintenance of genetic uniqueness of both Carpathian populations of P. cembra in general, and individual stands in particular, requires special measures for protection of Swiss stone pine in the Eastern Carpathians.     

Palaeostructures of vegetation at the upper limit of forests in the inner French Alps

The comparison of six pollen diagrams from French Alps allows us to reconstruct the past changes of vegetation structure at the upper limit of Subalpine range. Dense populations of Pinus cembra developed between 6500 and 2400 cal. BP, both in the southern Alps and the northern ones. Southern Alps seem however to be characterised by higher altitudinal limits, as shown by the past development of fir forests at 2080 m a.s.l. in the Ubaye valley. This study highlights the importance of taking in account local parameters in regional or continental reviews.     

Utilization of a biogeochemical model in dendroecology. Application to the Cembro pine

Dendroecology, which is based on annual growth variable measurements, is in possession of data particularly well adapted to analyse the impact of global change on vegetation production. But the classical statistical methods of quantification of tree ring-climate relationship cannot take into account the effect of CO2 increase. Therefore, a biogeochemistry model (BIOME3) has been adapted to these data and then validated on Pinus cembra. The results indicate that the production is reduced by 14% if only the climatic changes are taken into account. If both climatic changes and CO2 increase are taken into account the production is increased by 62%. The direct fertilisation effect of CO2 increase will have more influence on the productivity than the indirect climatic effect.     

Humus functioning types in evergreen coniferous forests of the French Inner Alps

Three humus forms that are widespread in the Inner Alps, a dysmull below Pinus sylvestris, an oligomull below Abies alba and a mor below Pinus cembra, were studied by following mineral nitrogen production and uptake and by a characterization of the pedofauna. Due to the production and uptake of nitrogen, mainly as nitrates and also due to the dominance of Lumbricids versus Arthropods, the oligomull was the sole humus form that functioned as a mull. Nevertheless relatively weak mineral nitrogen production and the dominance of endogeic soil-dwelling earthworms versus anecic earthworms demonstrated the moderate activity of this mull, which is representative of humus forms of old growth forests and cold climates. There was a discrepancy between the very low biological activity of the dysmull and its status of mull, while an observed bifunctioning between the different layers of this humus also differentiated it from the mor.     

Zur Taxonomie,Verbreitung und Ökologie des Filzigen Schmierlings Chroogomphus helveticus (Sing.) Moser

Chroogomphus helveticus is an european, dealtomontanous species with a south-hercynian-alpian-carpathian and illyrian-macedonian distribution. It grows in association with various conifers. Picea abies, Pinus cembra, Pinus peuce, Pinus strobus and Abies alba probably are partners of the mycorrhiza. The area is similar to that of Abies alba, Hygrophorus marzuolus and various dealpine plants. Distribution centres are districts, where 5-needled pines (Pinus cembra, Pinus peuce) are to be found naturally. Chroogomphus helveticus is to be found in an altitude from 300 to over 2000 m NN. The species is acidophilous, but not strongly bound to acid soils. A subdivion of the species into subspecific taxa is not possible. The closest relative of Chroogomphus helveticus occurs in Asia and Northern America. An other related european species is only known from England.     

Phylogenetic and biogeographic implications of chloroplast DNA variation in Picea

Purified chloroplast DNA ( cp DNA) extracts from 31 species of Picea and two species of Pinus (P. sylvestris and P. cembra ) were digested with eight restriction endo-nucleases, separated by electrophoresis and scored for restriction fragment length polymorphisms. The resulting data was analyzed phenetically and cladistically. The phenetic analysis indicated lower levels of cpDNA differentiation within Picea than within Pinus and lower levels of differentiation among Eurasian than among North-American Picea species. The cladistic analysis, using Pinus sylvestris as an outgroup, suggested monophyly for Picea and resolved several monophyletic groups among the 31 species of Picea . An assessment of biogeographic events, based on the cladogram, suggests that Picea originated in North-America and that the colonization of Eurasia occurred through separate, intercontinental migrations.     

The relationship between vegetation and pollen deposition in soil and in biological traps

The patterns of pollen deposition in the upper layers of the soil and in biological traps (mosses and lichens) in relation to the widespread and well known vegetational types of the Alps is investigated. The communities studied are beach woods, pine woods (Pinus cembra, P. sylveslris) and alpine pastures. Quantitative correlations between the pollen spectra and the vegetation have been calculated and over, under and equi-represented species delimited. Since representativity differs, in some cases, from that recorded in the literature it is suggested that it would be useful to continue the study in order to improve our understanding of actual pollen deposition and, as a consequence, of the interpretation of fossil pollen data.     

Effects of resource availability and distribution on autumn movements of the Nutcracker Nucifraga caryocatactes in the Alps

Radiotracking was used to study the ranging behaviour and movements of the Nutcracker Nucifraga caryocatactes in the Alevé forest (northwest Italian Alps), from September to December 1994-96, when Nutcrackers collected seeds of Arolla Pines Pinus cembra. Home range size and overlap decreased from September to December. Autumn ranging behaviour was not consistent with territoriality. Nutcrackers stored seeds throughout the Alevé; many apparently stored seeds in scree at high altitudes (above 2200 m) where the forest comes to an end, and also in Larch forests opposite the Alevé (probably to avoid competition with rodents and conspecifics). Estimates of Nutcracker abundance, derived from transect counts, varied significantly among habitats, years and months, which possibly reflected differences in seed availability and abundance. The ranging behaviour observed at the Alevé was compared with that monitored at Mont Avic, another area in the northwest Italian Alps, where resources (in this case Hazel Corylus avellana nuts) were concentrated in a few zones. Home range size was significantly larger at Mont Avic, especially during late autumn. There, the home range size increased and overlap decreased from September to November, when birds abandoned Hazels and dispersed to the coniferous woods. Differences between Alevé and Mont Avic implied that the availability and distribution of resources are the major determinants of the ranging behaviour of the Nutcracker during autumn. However, differences between adults and juveniles were also found.     

Terpenes from the Soft Corals Litophyton arboreum and Sarcophyton ehrenbergi

The new cembrane diterpene (3E,11E)‐cembra‐3,8(19),11,15‐tetraene‐7α‐ol ( 1 ), nephthenol ( 2 ), and all‐trans‐peridinin ( 3 ) have been isolated from the soft coral Litophyton arboreum. The tetraterpene 3 , (+)‐7,8‐epoxy‐7,8‐dihydrocembrene C ( 4 ), emblide ( 5 ), sarcophytoxide ( 6 ), sarcoglaucol‐16‐one ( 7 ), guajacophine ( 8 ), and 1,4‐peroxymuurol‐5‐ene ( 9 ) have been obtained from the soft coral Sarcophyton ehrenbergi. The compounds were characterized primarily by NMR spectroscopy. Some of the terpenes were tested for their antiproliferative activity against the cell lines HUVEC and K‐562 and for cytotoxicity against the cell line HeLa, and they showed moderate activities.     

Population structure,gene diversity,and differentiation in natural populations of Cedar pines (Pinus subsect.Cembrae,Pinaceae) in the USSR

Twelve natural populations of four cedar pine species,Pinus sibirica, P. cembra, P. pumila, andP. koraiensis, occurring in the Soviet Union were investigated by starch-gel electrophoresis. Frequencies of 55 alleles at 19 loci were determined. Interpopulation genetic diversity inP. sibirica andP. pumila was only 2–4 per cent of the total genetic diversity. Nei's distance coefficient (Dn) was used to estimate the level of genetic differentiation among conspecific populations and among species. Dn values among populations ranged from 0.006 to 0.038. A dendrogram constructed using Dn values divided cedar pines species into 2 clusters:sibirica-cembra (Dn = 0.030) andpumila-koraiensis (Dn = 0.143). Nei's distance between these clusters was 0.232. On the basis of the data obtained it was possible to draw the following conclusion:P. sibirica, P. pumila, andP. koraiensis are distinct species, whileP. cembra should apparently be regarded as geographicalP. sibirica race.     

The establishment of Holocene vegetation belts: quite near to a complete model of a solid mass

Pollen and macro-remains were analysed in a sixth site (La Gouille 1,800 m) of the Cha?ne des Hurtières (northern French Alps). Nine A.M.S. dates support the chronology. Thus, the establishment of the vegetation belt of a massif can be modelled in the northern French Alps. Betula invaded sub-Alpine grasslands as early as 10,000 14C BP. Around 9,600 14C BP shrublands with Corylus, Alnus and Sorbus were established before the spread of Abies at the site approximately 8,200 14C BP. A decrease in Abies prior to 8,100 14C BP occurred during the Venediger climatic oscillation. At around 2,940 14C BP, a strong regression of Abies due to human action is noted with the expansion of Alnus viridis. Recently, a second Abies retraction led to the present sparce P. cembra and Alnus viridis vegetation cover.     

Home range and habitat selection by the Nutcracker Nucifraga caryocatactes during autumn in the Alps

In the Mont Avic Natural Park (NW Italian Alps), the Nutcracker Nucifraga caryocatactes occurred in coniferous forests where Arolla Pines Pinus cembra, its preferred habitat, were absent. A radiotelemetric study carried out during autumn 1992 and 1993 showed that, from September, Nutcrackers concentrated in the few Hazel Corylus avellana hedges of the rural zones of the park to collect nuts, which were eaten or stored. Individual home ranges overlapped to a high degree until October, when birds started to abandon the Hazel areas. From October onward, individual home ranges significantly expanded and their degree of overlap diminished compared to September. The home range analysis and the observations in the field suggested that the population of Nutcrackers in the park during autumn was made up of at least three different categories of birds, which displayed different ranging behaviour patterns. A few untagged birds probably stored nuts in their breeding territories, whereas all the radio-tagged birds ( n = 15) gave no indication of territorial use, since from October they either stored nuts in the coniferous woods near the Hazel hedges or abandoned the park to fly to other Hazel areas. Hence, during autumn the ranging behaviour of the Nutcrackers seems to be largely independent of their reproductive territorial system. Habitat selection analysis and transect counts revealed that birds positively selected Hazels, especially in the coniferous woods around Hazel hedges, and avoided the other coniferous forests. Nevertheless, from October some individuals reversed their habitat choices and positively selected coniferous forests (especially Mountain Pine Pinus uncinata woods) while avoiding Hazel.     

Age-dependent xylogenesis in timberline conifers

Neither anatomical change nor physiological abnormalities have been observed in the cambia of older trees. However, different sensitivity and period of significant responses to climate suggest the existence of some age-related change in the patterns of cambial activity and/or wood cell formation. Here, weekly cambial activity and timing and duration of xylem cell enlargement and wall thickening were compared in adult (50-80 yr) and old (200-350 yr) trees of Larix decidua, Pinus cembra and Picea abies at the Alpine timberline during 2004 and 2005. Timings and durations of xylogenesis differed between adult and old trees, with 2-3 wk shorter cambial activity found in the latter. The delayed onset of cambium division and lower cell production in old trees, with respect to adult trees, led to reductions of 15-20% in the overall duration of xylem differentiation. These results demonstrate that cambial dynamics change during the tree lifespan and that the time window of tree-ring production shortens with age. Variations in the period of xylem growth may be the cause of age-dependent responses to climate. The observed shorter xylogenesis in older plants at the Alpine timberline could be related to a size effect and not just to age per se.     

Aerial decay influence on the stable oxygen and carbon isotope ratios in tree ring cellulose

Sub-fossil wood is often affected by the decaying process that introduces uncertainties in the measurement of oxygen and carbon stable isotope composition in cellulose. Although the cellulose stable isotopes are widely used as climatic proxies, our understanding of processes controlling their behavior is very limited. We present here a comparative study of stable oxygen and carbon isotope ratios in tree ring cellulose in decayed and non-decayed wood samples of Swiss stone pine (Pinus cembra) trees. The intra-ring stable isotope variability (around the circumference of a single ring) was between 0.1 and 0.5‰ for δ¹⁸O values and between 0.5 and 1.6‰ for δ¹³C values for both decayed and non-decayed wood. Observed intra-tree δ¹⁸O variability is less than that reported in the literature (0.5–1.5‰), however, for δ¹³C it is larger than the reported values (0.7–1.2‰). The inter-tree variability for non-decayed wood ranges between 1.1 and 2.3‰ for δ¹⁸O values, and between 2 and 4.7‰ for δ¹³C values. The inter-tree differences for δ¹⁸O values are similar to those reported in the literature (1–2‰ for oxygen and 1–3‰ for carbon) but are larger for δ¹³C values. We have found that the differences for δ¹⁸O and δ¹³C values between decayed and non-decayed wood are smaller than the variation among different trees from the same site, suggesting that the decayed wood can be used for isotopic paleoclimate research.     

Larch- and pine-feeding host races of the larch bud moth (Zeiraphera diniana) have cyclic and synchronous population fluctuations

Population cycles of many forest-defoliating insects often show synchronous fluctuations at both intra-specific (spatial synchrony) and inter-specific levels. However, population dynamics of different host-associated biotypes of the same species, such as those of the larch budmoth (LBM), Zeiraphera diniana (Lepidoptera: Tortricidae), have never been compared. This species causes extensive defoliation of larch forests every 8 to 9 years in the Alps, but it consists of two genetically-differentiated host races, the first one developing on European larch, Larix decidua , and the other one developing on Swiss stone pine, Pinus cembra . The dynamics of Zeiraphera populations have been extensively studied on larch, whereas little is known about larval density and possible population fluctuations on sympatric pines. A larval census on Swiss stone pine was conducted in the Swiss Alps intermittently between 1958 to 2004 and in the French Alps from 1992 to 2004. Population density of Zeiraphera on pine varied up to 5000-fold and showed periodic oscillations, with five peaks in Switzerland and one in France. Because the feeding activity of the pine race is restricted to the elongating shoot of the current year, no conspicuous defoliation of pine trees was noted during years of high larval densities. Zeiraphera populations on pine oscillated in significant synchrony with larch-associated populations, and peak densities were observed either the same year or shifted by±one year. Our results did not allow any explanation for cyclic fluctuations of LBM on pine, but the synchrony with the larch race's cycle suggests that studies on genetics as well as on parasitism should be intensified.