Ecology of Armillaria species on silver fir (Abies alba) in the Spanish Pyrenees

? We describe the distribution and the ecology of three Armillaria species observed in silver fir (Abies alba) forests of the Pyrenees.

? We surveyed the presence and abundance of Armillaria above and belowground in 29 stands. Isolates were identified by the PCR-RFLP pattern of the IGS-1 region of their ribosomal DNA. We measured several ecological and management parameters of each stand in order to describe Armillaria infected sites.

? Armillaria cepistipes was the most abundant of three species observed. Armillaria gallica was dominant in soils with a higher pH and at lower elevations. Armillaria ostoyae seemed to be more frequent in stands where A. alba recently increased its dominance relative to other forest tree species. Thinning activities correlated with an increased abundance of Armillaria belowground. In 83% of the stands the same Armillaria species was observed above and belowground.

? It seems that in a conifer forest, A. cepistipes can be more frequent than A. ostoyae, a virulent conifer pathogen. Since logging is related to a higher abundance of Armillaria in the soil, the particular Armillaria species present in a given stand could be considered an additional site factor when making management decisions.

     

Ectomycorrhizal communities associated with silver fir seedlings (Abies alba Mill.) differ largely in mature silver fir stands and in Scots pine forecrops

Context

The requirement for rebuilding forecrop stands besides replacement of meadow vegetation with forest plants and formation of soil humus is the presence of a compatible ectomycorrhizal (ECM) fungal community.

Aims

This study aims to assess ectomycorrhizal fungi diversity associated with silver fir (Abies alba Mill.) seedlings regenerating in silver fir stands and Scots pine forecrops.

Methods

One-year-old seedlings were sampled in six study sites: three mature fir forests and three pine forests. ECM fungi were identified by polymerase chain reaction amplification and sequencing of the internal transcribed spacer of rDNA.

Results

The mean mycorrhizal colonization exceeded 90 %. Thirty-six ectomycorrhizal taxa were identified in fir stands and 23 in pine forecrops; ten out of these species were common to both stands. The fungal communities were different between study sites (R?=?0.1721, p?=?0.0001). Tomentella stuposa was the only species present at all sites.

Conclusion

Silver fir seedlings in Scots pine forecrops supported smaller ECM fungal communities than communities identified in mature silver fir stands. Nevertheless, fungal colonization of seedling roots was similar in both cases. This suggests that pine stands afforested on formerly arable land bear enough ECM species to allow survival and growth of silver fir seedlings.     

Effect of knife wear on finger-joint strength of silver fir (Abies alba) wood

Abstract

This study was carried out to determine the effect of knife wear, as measured in hours of use, on the finger-joint strength of silver fir wood (Abies alba), bonded with polyurethane (PU) and polyvinyl acetate (PVAc) adhesives. Wood samples were collected after 1, 32 and 64 h of knife wear. Results showed that as knife wear increased, bending strength or modulus of rupture (MOR) of the finger-jointed samples was reduced by 11.1% (PVAc) to 21.2% (PU) after 32 h of knife wear, and by 19.6% (PVAc) to 27.6% (PU) after 64 h of knife wear, whereas modulus of elasticity (MOE) of the finger-jointed samples was reduced by 10.5% (PVAc) to 12.5% (PU) after 32 h of knife wear, and by 11.7% (PVAc) to 10.0% (PU) after 64 h of knife wear. Tension strength of the finger-jointed samples was reduced by 2.1% (PVAc) to 1.5% (PU) after 32 h of knife wear, and by 5.4% (PVAc) to 16.0% (PU) after 64 h of knife wear.     

An isolated population of Heterobasidion abietinum on Abies pinsapo in Spain

Abies pinsapo, a fir endemic to a small area in southernmost Spain, is frequently attacked in natural stands by Heterobasidion root rot. To identify the Heterobasidion species causing the disease, pure cultures of the fungus were isolated from symptomatic trees in five localities in Sierra de las Nieves Natural Park and identified by pairing tests. In addition, genets of the fungus were identified in two pure stands of A. pinsapo. All the Heterobasidion specimens collected belonged to the species Heterobasidion abietinum. The largest genet found was 57 m long, and had colonized 10 trees. The large size of the main genets implied that H. abietinum had spread via root contacts from old infections generated before the establishment of the Sierra de las Nieves National Park in 1989. Exceptionally dry summers in the last 20 years may have weakened A. pinsapo and favoured spread of the disease.     

The effect of different mixtures of NaCl and CaCl2 on the silver fir (Abies alba Miller)

Young potted silver firs were treated with six different mixtures of NaCl and CaCl². Growth was measured as well as the foliar contents of Cl, Na and Ca and the development of chlorosis and necrosis was recorded. At both concentration levels used, growth inhibition and needle necrosis were more pronounced in test conditions rich in CaCl², a fact which is probably due to Cl toxicity.     

Susceptibility of Abies pinsapo and its tree cohort species to Heterobasidion abietinum

Abies pinsapo, a rare fir found in the southernmost mountain region of Spain, is severely affected by Heterobasidion abietinum. It was hypothesized that spread of the disease might be favoured by woody species growing in the same habitat. In order to evaluate this hypothesis, artificial inoculation tests were performed on other potential hosts forming the tree cohort species with A. pinsapo. Inoculation test on seedlings demonstrated that H. abietinum was able to colonize stems of Pinus halepensis, P. pinaster and Quercus ilex subsp. rotundifolia, but failed on Q. faginea. Both Pinus species were colonized to a similar extent as the natural host, but Q. ilex subsp. rotundifolia showed a significantly lower susceptibility than conifers. These results suggest that Pinus spp. growing in mixed stands with A. pinsapo might favour spread of H. abietinum. As a consequence, natural replacement of pure A. pinsapo stands by mixed forest Abies‐Quercus should be facilitated.     

The efficacy of selected biological and chemical control agents against Heterobasidion abietinum on Abies cilicica

The efficacy of 30% aqueous urea solution, borax powder and spore suspensions of Phlebiopsis gigantea and Trichoderma harzianum against establishment of artificially inoculated Heterobasidion abietinum on Abies cilicica was tested both in the field and in a simulated stump treatment experiment carried out in a growth chamber. In the field, in each of the four selected stands 20 fresh stumps per control agent were treated and 20 stumps were left as untreated controls. In two of the stands, the treatments were applied in June and in the two others in November. Stumps were sampled twice, at 6‐ and 12‐months after treatment. In the growth chamber, 10 log pieces per treatment and control were used, and sampling performed after 6‐week incubation. Results of the urea and borax treatments were consistent between the experiments; the mean efficacies were 98.8 and 99.4% in the growth chamber, and 80.2–91.3 and 89.4–90.1% in the two samplings of the field experiment, respectively. Despite the high efficacies of the P. gigantea (85.9%) and T. harzianum (97.5%) treatments in the growth chamber, efficacies of these biological control agents in the field were 47.1–49.2 and 61.3–65.5%, respectively.     

Relationships between local stand density and local species composition and nutrient content in the topsoil of pure and mixed stands of silver fir (Abies alba Mill.)

This study examined topsoil samples from the humic mineral horizon in four pure silver fir (Abies alba Mill.) and five mixed stands with beech (Fagus sylvatica L.), spruce (Picea abies L.; Karst.) and pine (Pinus sylvestris L.). In every stand, 60–70 sampling locations were chosen using a stratified random scheme that represented a gradient in local stand density and local species composition. One working hypothesis was that stand density and species composition affect the content of nutrients in the mineral humic soil horizon. Specifically, the analysis statistically tested for relationships between general and species-specific local stand density measures (basal area, sum of tree diameters weighted by distance, canopy openness) and the contents of total C, total N (in four stands), base cations (Ca²⁺, Mg²⁺, K⁺, Na⁺) and Mn²⁺ (in nine stands). A considerable variation in nutrient concentrations was observed; the coefficients of variation ranged from 0.21 to 0.38 for N, from 0.20 to 0.94 for the sum of base cations and from 0.56 to 0.95 for Mn. However, no clear relationship emerged between local stand density and the concentration of N and base cations; in five stands, manganese showed a negative correlation with local stand density. In the mixed fir-beech stand, significantly higher concentrations of Ca, K and Mn were found in the vicinity of beeches, and species-specific stand density explained up to 15, 29 and 34% of total variation in the content of these elements, respectively. Similar tendencies also appeared in the mixed stands with pine, although the relationships were weak. The results obtained for two fir-spruce stands were ambiguous. In sum, in the stands studied, small canopy gaps and inter-crown openings might not be considered ‘‘nutrient hot spots’’, and the admixed species, except for beech, had an insignificant effect on nutrient content in the humic mineral horizon.     

Ethylene production and premature senescence of needles from fir trees (Abies alba)

Measurements of ethylene production, free and conjugated ACC concentrations, chlorophyll content, and length of needles from fir trees (Abies alba Mill.) growing in stands affected by “Waldsterben” showed that during the development of the disease two consecutive stages can be distinguished: a. long-term, low ethylene production and accumulation of conjugated ACC (MACC) in response to chronic stress, associated with reduced needle growth, and b. high ethylene production associated with chlorophyll breakdown, followed by premature abscission of the needles.     

Variability of stomatal conductance in a small and isolated population of silver fir (Abies alba Mill.)

We analyzed the response to drought of 420 individuals from eight half-sib families from a small and isolated population of silver fir (Abies alba Mill.): 105 of them were kept in well-watered conditions as control while the remaining 315 were exposed to drought for 27 days. A model describing stomatal behavior derived from Monteith and developed in beech by Leonardi et al. was fitted to experimental transpiration data obtained simply from the difference between two daily pot weighings. The estimated parameters were maximum stomatal conductance, maximum transpiration in well-watered conditions and sensitivity to soil water deficit. The model worked well: convergence for all but four individuals and concordance between experimental and fitted data were good (R(2)=0.86). Inter-individual variability for all three estimated parameters was high and two of them (maximum stomatal conductance and sensitivity to soil water deficit) were significantly different among families, suggesting genetic control. Our results validate the simplified method used to evaluate individual stomatal parameters. We also show that in the small and isolated population of our study substantial adaptive variability remains, a crucial prerequisite to endure environmental conditions determined by climatic change foreseen for the next decades.     

Spatial and seasonal variation in amino compounds in the xylem sap of a mistletoe (Viscum album) and its hosts (Populus spp. and Abies alba)

In a field study, the composition and concentrations of amino compounds in the xylem sap of the mistletoe, Viscum album L., and in the xylem sap of two host species, an evergreen conifer (Abies alba Mill.) and a deciduous broad-leaved tree (Populus x euramericana), were analyzed. The xylem sap of both hosts and mistletoe contained large, but similar amounts of total organic nitrogen in low molecular weight amino compounds (TONLW). Nevertheless, individual amino compounds accumulated in the xylem sap of mistletoe relative to the host xylem sap, indicating selective uptake. In the xylem sap of Populus, major amino compounds (asparagine (Asn) and glutamine (Gln)) and the bulk parameters, TONLW and proteinogenic amino acids, showed significant seasonal variation. In Abies and in mistletoe on either host, variation of amino compounds in xylem sap was largely explained by inter-annual differences, not by seasonal variation. In both hosts, TONLW in the xylem sap was dominated by Gln. There was a steady decrease in relative abundance of Gln from the host xylem sap to the mistletoe xylem sap and to the stems and leaves of mistletoe. Simultaneously, the abundance of arginine (Arg) increased. Arginine was the predominant amino compound in the stems and leaves of mistletoe, occurring at concentrations previously observed only in leaves of trees exposed to excess nitrogen. We conclude that Gln (2 mol N mol(-1)) delivered by the host xylem sap is converted, in mistletoe, to Arg (4 mol N mol(-1)) and that the organic carbon liberated from Gln contributes significantly to the parasite's heterotrophic carbon gain. Statistical analyses of the data support this conclusion. Accumulation of Arg in mistletoe is an indication of excess N supply as a result of the uptake of amino compounds from the host xylem sap and a lack of phloem uploading.     

Retrospective analysis of tree architecture in silver fir (Abies alba Mill.): ontogenetic trends and responses to environmental variability

Context

Understanding the effects of exogenous factors on tree development is of major importance in the current context of global change. Assessing the structure development of trees is difficult given that they are large and complex organisms with lifespan of several decades.

Aims

We used a retrospective analysis to derive the ontogenetic trends in silver fir development and assess the effects of climate or light environment on tree architecture.

Methods

Thanks to the identification of relevant growth markers (bud cataphylls and pseudo-whorl branches), a retrospective analysis allowed to record annual shoot extension and to date them on silver firs of various sizes under different environmental conditions.

Results

The length of successive annual shoots located on different axes clearly show gradual trends related to the physiological age of meristems. Within- and between-tree variations are noted due to the plasticity of development and growth induced by light environment and climate.

Conclusion

Retrospective analysis is an efficient method for getting information on the history of trees architecture and subsequently to relate it to environmental factors.     

Individual vulnerability factors of Silver fir (Abies alba Mill.) to parasitism by two contrasting biotic agents: mistletoe (Viscum album L. ssp. abietis) and bark beetles (Coleoptera: Curculionidae: Scolytinae) during a decline process

? Context

In recent decades, there have been increasing reports of forest decline, especially in Mediterranean forest ecosystems. Decline in tree vitality is usually due to complex interactions between abiotic factors and biotic agents that attack weakened trees.

? Aims and methods

Estimating dendrometrical characteristics [basal area increment (BAI), age at DBH from tree ring counting, social status, height, and diameter], tree health status, and a competition index, we investigated the individual vulnerability of a French declining silver fir forest to both mistletoe (Viscum album L. ssp. abietis) and bark beetles (Pityophthorus pityographus Ratz., Pityokteines vorontzovi Jac., and Pityokteines spinidens Reitt.).

? Results

BAI was negatively correlated with both mistletoe infection (via mistletoe biomass) and bark beetle attack (number of insects per square meter), but there was evidence of divergence in tree choice between two groups of parasites. Mistletoe preferentially infected isolated and dominant trees that showed higher past growth rates than non-infected ones. Conversely, bark beetles mainly attacked defoliated and preferably declining trees with diameter (DBH) lower than 44.5 cm and slower past growth.

? Conclusion

While successive severe drought periods are thought to greatly weaken southern silver fir populations, mistletoe and bark beetles may contribute actively to their decline processes as inciting and contributing factors, respectively.     

Susceptibility of red fir and white fir to fir dwarf mistletoe (Arceuthobium abietinum) in California

Fir dwarf mistletoe (Arceuthobium abietinum, Viscaceae) is a damaging parasite of true firs (Abies spp.) in California. In the Sierra Nevada Mountains, fir dwarf mistletoe consists of two special forms (formae speciales, f. sp.) which are morphologically similar, but exhibit a high degree of host specificity: A. abietinum f. sp. concoloris whose principal host is Sierra white fir (Abies lowiana) and A. abietinum f. sp. magnificae which only parasitizes red fir (Abies magnifica). Although field observations and cross‐inoculation studies have consistently supported the extreme host specialization demonstrated by these special forms of fir dwarf mistletoe, little quantitative data from mixed red fir/white fir stands infested with fir dwarf mistletoe have been available. Therefore, I sampled 20 mixed conifer stands dominated by red and white fir and infested with f. sp. concoloris (8 stands) or f. sp. magnificae (12 stands) from near Mount Shasta to the southern Sierra Nevada Mountains near Sherman Pass. Infection of red and white fir was evaluated using temporary circular plots (0.012 ha) established around large, severely infected trees and with rectangular transects placed within fir dwarf mistletoe‐infested stands. In each plot and transect, live trees >5 cm in diameter at breast height were assigned a dwarf mistletoe rating (6‐class system). A total of 55 plots and 51 transects were established in the 20 stands, and over 1,900 red firs and 1,700 white firs were sampled. Ninety‐nine per cent of the red firs and 98% of the white firs were infected in stands infested with f. sp. magnificae or f. sp. concoloris, respectively. In contrast, no infection of red fir was observed in stands infested with f. sp. concoloris and no infected white firs were observed in stands infested with f. sp. magnificae. These data provide further support that the special forms of fir dwarf mistletoe are extremely host specific in California.     

Effects of ozone on gas exchange,frost resistance,flushing and growth of different provenances of European silver fir (Abies alba Mill.)

Photosynthesis, respiration, and transpiration were studied in different Abies alba provenances during and alter ozone exposure at three different levels (0/control, 125, and 250 ppb O₃, respectively). The after-effects of ozone on frost resistance and growth rate were also analyzed. Pronounced reductions of photosynthesis and transpiration rates and an increased respiration were registered as a response to O₃. Five months after exposure the winter frost resistance was significantly reduced, and growth depressions were registered during the following year. Differences in ozone sensibility and regeneration ability between provenances could be demonstrated.     

Climatic signal in annual growth variation of silver fir (Abies alba Mill.) and spruce (Picea abies Karst.) from the French Permanent Plot Network (RENECOFOR)

This paper explores the growth/climate relationships in earlywood, latewood and total ring-width chronologies of five Norway spruce (Picea abies) and six silver fir (Abies alba) stands sampled in the French permanent plot network (RENECOFOR) (327 trees). The relationships between climate and ring widths were analyzed using extreme growth years, simple correlations and response functions analysis (bootstrapped coefficients). Monthly climatic regressors were derived by a physiological water balance model that used daily climatic data and stand parameters to estimate soil water deficits. Pointer years underline the high sensitivity to winter frosts (1956, 1986) and exceptional annual droughts (1962, 1976, 1991) for both species. For those years, growth variations were higher for Abies alba than for Picea abies. For each species, the climate information of tree ring series is not modified by local site characteristics (altitude, slope, aspect, soil water reserve). Moreover, strong specific differences appear among species. Earlywood and latewood spruce growth mainly depends on current summer soil water deficit conditions. For silver fir, winter and early summer temperatures, as well as the water supply of the previous year (August to October) play a major role for the production of earlywood, after which the current early summer water supply influences mainly latewood width.     

Is Silver fir (Abies alba) less vulnerable to extraction damage than Norway spruce (Picea abies)?

Inspection of Norway spruce and Silver fir on experimental plots in south-western Germany showed that Silver fir had suffered significantly less bark injuries than Norway spruce. Data from both federal forest inventories (1987, 2002) showed a similar species-specific vulnerability. Additional visual inspections of the basal cross-sections of trees removed from the experimental plots showed rather high proportions of butt rot in uninjured Norway spruce (51%). The proportion further increased to 93% in trees, which had sustained bark injuries. In contrast, decay symptoms were almost absent in uninjured Silver fir and less enhanced in trees with bark injuries (27%). Management implications for risk rating of tree species, as well as the necessity of implementing low-damage harvesting regimes, are discussed.     

Terpene response of Picea abies and Abies alba to infection with Heterobasidion s.l.

The monoterpene composition of Picea abies and Abies alba resin was analysed in relation to growth by Heterobasidion spp. Fifteen‐year‐old P. abies and A. alba trees were inoculated on branches with three species of Heterobasidion annosum s.l. After 4 months of incubation, each host was colonized to a significantly greater degree by the pathogen specific to that host (H. parviporum on P. abies, H. abietinum on A. alba) than by the other fungi. Analysis of the enantiomeric monoterpene profiles in the spruce and fir showed that the response in terms of the relative proportions of the monoterpene compounds in the resin differed between tree species. Following challenge with Heterobasidion spp., A. alba trees did not show changes in monoterpene composition in addition to those in the wounding response (increase in (−)‐α‐pinene and (−)‐camphene, and decrease in β‐phellandrene). In P.abies, (−)‐α‐pinene, (+)‐α‐pinene and δ‐3‐carene increased following Heterobasidion attack but not after wounding alone.