Heterobasidion on Abies nordmanniana in north‐eastern Turkey

Occurrence of Heterobasidion annosum s.l. was investigated in 15 coniferous stands in the Giresun and Pontic Mountains in north‐eastern Turkey. Basidiocarps of the fungus were found in 11 stands. Fifty‐two basidiocarp specimens of Heterobasidion were collected from stumps of Abies nordmanniana ssp. nordmanniana and two from stumps of Picea orientalis. Pure cultures were isolated from the basidiocarps and identified with the aid of mating tests. Forty‐five (90%) of the specimens collected from A. nordmanniana were identified as H. abietinum and five (10%) as H. annosum s.s. The former species was also found twice on P. orientalis. This is the first report of H. annosum s.s. for Turkey and the first report of H. abietinum on P. orientalis. Heterobasidion abietinum seems to be mostly a saprotroph on A. nordmanniana.     

Turkish Heterobasidion abietinum is pathogenic to inoculated Abies nordmanniana ssp. nordmanniana and ssp. bornmülleriana

Four‐year‐old seedlings of Abies nordmanniana ssp. bornmülleriana and 5‐year‐old seedlings of Abies nordmanniana ssp. nordmanniana were inoculated on the lower stem with 28 Heterobasidion abietinum isolates originating from four different regions of Turkey. Replicate seedlings were incubated in greenhouse and growth chamber. After 7 weeks, infection incidence, mortality, lesion length in the inner bark and fungal growth in the sapwood were examined. Infection incidence in different host–incubation combinations ranged between 70.5 and 79.5% and the average mortality rate was 4.2%. Average lesion lengths varied between 8.4 and 33.9 mm, and average fungal growth between 18.7 and 34.8 mm. There was a significant positive correlation between lesion length in the inner bark and fungal growth in sapwood on both hosts and conditions. Clear differences in virulence between H. abietinum isolates originating from different regions were not found. The results indicate that H. abietinum is pathogenic on both investigated subspecies of A. nordmanniana. Both lesion length and fungal growth were greater on ssp. bornmülleriana, indicating that it would be more susceptible than ssp. nordmanniana.     

Genetic variation of Heterobasidion abietinum in Southern Europe and the Mediterranean Basin

Heterobasidion abietinum causes root and butt rot on Abies species occurring in central Europe and the Mediterranean Basin in a distribution extending from Spain in the west to Turkey and Caucasia in the east. To investigate the genetic diversity within H. abietinum, 95 isolates from different regions were analysed with 61 polymorphic molecular markers. The markers showed genetic differentiation amongst the H. abietinum populations. The most diversified population is that on A. pinsapo in southern Spain, which is isolated from gene flow with other populations. Lower but still significant differences were found amongst other European populations. Isolates from Turkey showed little diversification compared with the south European isolates from east of the Alps. The genetic structure of the H. abietinum populations in relation to postglacial recolonization of Abies is discussed.     

Cedrus libani: the most susceptible Turkish conifer species to local Heterobasidion isolates in spring inoculations

Five‐year‐old seedlings of Pinus nigra, Pinus brutia, Abies nordmanniana ssp. bornmülleriana and Cedrus libani, 3‐year‐old seedlings of Pinus sylvestris, and 2‐year‐old seedlings of Juniperus excelsa were inoculated on the lower stem with Turkish Heterobasidion abietinum and Heterobasidion annosum s.s. isolates. In total, 300 seedlings were inoculated in April 2007 with five isolates of each Heterobasidion species and incubated in a glasshouse for 12 weeks. The daily maximum temperature increased gradually from 13°C to 31°C by the end of the incubation period. Infection incidence, mortality, lesion length in the inner bark and fungal growth in the xylem were examined. For H. abietinum, infection incidence ranged from 0% to 36%, and for H. annosum s.s. from 0% to 60%. Mortality was low; only 1% of the inoculated seedlings died. Mean lesion lengths varied between 7.6 and 11.9 mm for H. abietinum and 8.0 and 15.7 for H. annosum s.s. in all species apart from C. libani, in which the corresponding values were 42.2 and 42.6 mm respectively. Similarly, mean fungal growth in sapwood of C. libani was approximately 25 mm for both Heterobasidion species, in contrast to 0–3.2 mm in the other tree species. The H. abietinum isolates were not reisolated from P. nigra, P. sylvestris or J. excelsa. The H. annosum s.s. isolates did not infect Abies seedlings. Only P. brutia and C. libani were susceptible to both pathogens. Control seedlings showed no symptoms. The results indicated that C. libani is highly susceptible to Turkish isolates of Heterobasidion.     

Armillaria ostoyae in managed coniferous forests in Kastamonu in Turkey

Although several Armillaria species have been reported in Turkey, there is little information about their ecology in Turkish forests. In this study, we investigated five forest stands, approximately 5–74 ha in size, in Kastamonu province in the Black Sea Region of Turkey for the presence of Armillaria species in stumps and logs. The stands were mixed Abies nordmanniana ssp. bornmülleriana and Pinus sylvestris forests managed using a selective cuttings system; the proportion of fir in the total number of stems and stumps ranged from 36 to 98%. Based on sequence analysis of the internal transcribed spacer and intergenic spacer regions of the rDNA, all rhizomorphs sampled from the stumps and logs were of Armillaria ostoyae. The size of the genets was estimated with random amplified microsatellites analysis of the isolates and ranged from single stumps to approximately 450 m². One to seven genets were found in each stand. These results indicate that the genets had arisen from spores and vegetative spread was limited on most sites.     

Genetic variation in Heterobasidion abietinum populations detected with the M13 minisatellite marker

Amplification profiles of the M13 minisatellite core sequence from 47 isolates of Heterobasidion annosum s.l., collected mostly from Abies alba in different European countries, were analysed in comparison with profiles of four isolates of H. annosum s.s. and three isolates of H. parviporum. Genetic variation within and among groups of populations was studied by analysis of molecular variance. A dendrogram constructed with the Neighbor‐Joining method differentiated the three species from each other. Isolates of H. abietinum from Balkan peninsula, Italian peninsula and French Pyrenees each separated into clusters according to geographical origin. Such clear geographic clustering was not detected among isolates from central Europe. The variations between H. abietinum isolates in relation to the migration history of Abies are discussed.     

Investigations on Heterobasidion annosum s.lat. in central and eastern Asia with the aid of mating tests and DNA fingerprinting

To investigate the taxonomy of Heterobasidion in Eurasia, 49 specimens belonging to H. annosum sensu lato from Asia were identified with the aid of mating tests. Most of the specimens originated from north‐eastern and south‐western China and from the Altai region in southern Siberia, but a few isolates from Kirghizia, Japan and India were also tested. In addition to mating tests, the material from China was investigated with DNA fingerprinting. Heterobasidion annosum sensu stricto was identified only from the Altai region. Homokaryotic isolates from other specimens, except the Indian ones, were sexually compatible with H. parviporum, but they also showed a high degree of compatibility with H. abietinum and with the North American S group. The isolates from SW China (eastern Himalayas) mated with about equal frequency with the European strains of H. parviporum and H. abietinum. However, the DNA fingerprinting showed that these isolates were more closely related to H. parviporum, and hence they were tentatively included in this species. The North American S group was more distant from these Eurasian taxa. Four old isolates from India mated only weakly with the members of the H. annosum s.lat. According to the species concept presented, the distribution of H. parviporum extends from western Europe through southern Siberia to northern China, Japan and the eastern Himalayas. H. annosum s.str. is so far identified only from the Altai region outside Europe, and H. abietinum only from Europe.     

Mortality and growth of dwarf mistletoe‐infected red and white fir and the efficacy of thinning for reducing associated losses

In managed forests dominated by true fir (Abies) species, stands are often restocked using understory trees retained during timber harvest, making the effects of dwarf mistletoe (Arceuthobium spp.) infestation on small true fir a concern. This study examined the response of small red (A. magnifica) and white (A. concolor) fir and their dwarf mistletoes (A. abietinum f.sp. magnificae and A. abietinum f.sp. concoloris, respectively) to precommercial thinning treatments in fir‐dominated stands in the Sierra Nevada Mountains of California. Tree diameters and dwarf mistletoe ratings were monitored from 1981 to 2001, and mortality was observed from 1981 to 2006. Red and white fir survival and radial growth decreased significantly with greater mistletoe ratings and increased with larger diameters and live crown ratios. Thinning significantly increased survival and growth of white, but not red fir. Over the course of the study, mistletoe ratings for both fir species did not change significantly in unthinned stands, but increased in thinned stands. However, while statistically significant, increases in mistletoe ratings in thinned stands were gradual and mistletoe ratings did not statistically differ between treatments 20 years post‐thinning. Additionally, thinning did not significantly influence mistletoe spread to uninfected trees, indicating that precommercial thinning in lightly infested red and white fir stands will not likely result in substantial increases in dwarf mistletoe abundance over typical harvesting intervals. Overall, while severe dwarf mistletoe infection ratings reduced tree survival and growth, because ratings remained low, actual losses resulting from mistletoes during this study were minor and will not likely result in substantial economic losses over normal harvesting intervals. This finding indicates that precommercial thinning treatments designed specifically to compensate for mistletoe‐associated losses may not be necessary when managing red and white fir for timber production.     

Induction of tumours by various strains of Agrobacterium tumefaciens on Abies nordmanniana and Picea abies

Abies nordmanniana and Picea abies seedlings were inoculated with a wild‐type (C 58) or an attenuated strain (rooter or shooter mutants) of Agrobacterium tumefaciens. Large tumours were formed on A. nordmanniana in response to the wild‐type and rooter strains, 60–65 % of the seedlings being susceptible. Smaller tumours were formed on Picea abies in response to the wild‐type strain, at least 12% of the seedlings being susceptible. Tumours from both species induced by the wild‐type strain grew in vitro on medium without added phytohormones.     

Gene flow and inter‐sterility between allopatric and sympatric populations of Heterobasidion abietinum and H. parviporum in Europe

The tree pathogenic fungi Heterobasidion parviporum and H. abietinum are recently described sibling species that show marked host preference for Picea abies and Abies alba, respectively. H. parviporum shows a higher reproductive character displacement (i.e. higher inter‐sterility) towards H. abietinum in sympatry than in allopatry. We inspected whether this pattern was also present in H. abietinum by studying the inter‐sterility and gene flow amongst sympatric central European populations of H. abietinum and H. parviporum, and two isolated allopatric populations, Pyrenean H. abietinum and Scandinavian H. parviporum. Inter‐ and intra‐specific fixation index values were calculated from DNA sequence data of the GST‐1 locus and four microsatellite loci. Present allopatric and sympatric populations of H. abietinum were equally inter‐sterile towards H. parviporum, not suggestive of relaxed reinforcement. Several hypotheses explaining the observed pattern are presented. Genetic differentiation was observed between H. abietinum in the Pyrenees and in the Alps, suggesting geographical structure of H. abietinum in Europe.     

Testing species of genus Abies for Christmas tree production in Norway

Ten exotic Abies species were tested in two field trials, where the aim was to find suitable species and provenances for Christmas tree production in coastal and fjord areas in Norway. The material included 14 provenances of Abies nordmanniana, 3 provenances of Abies bornmuelleriana, 3 provenances of Abies koreana, 2 provenances of Abies amabilis, and 1 provenance each of Abies equi-trojani, Abies alba, Abies procera, Abies homolepis, Abies nephrolepis, and Abies forrestii. Field trials were established at Gulen in Sogn og Fjordane County and at Verdal in Nord-Trøndelag County. Christmas tree classification was done seven and eight growing seasons after establishment. The Christmas tree yield was higher in Gulen (64%) than in Verdal (45%), which is situated further north. Also, the survival and the height growth were higher in Gulen than in Verdal. A more humid climate and a longer growth season in Gulen may explain some of the differences. Of the 10 species, A. nordmanniana, A. homolepis, and A. bornmuelleriana produced the highest Christmas tree yield at Gulen, while A. homolepis and A. koreana had the highest yield at Verdal. Due to early bud burst, A. equi-trojani and A. bornmuelleriana are only suited for Christmas tree growing in the best climatic areas.     

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.     

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.     

Untersuchungen über Fichtengallenläuse, Dreyfusia spp. (Hom., Adelgidae) und deren Predatoren in der Nord-Türkei

In the East Pontic mountains at Giresun (Black Sea shore) three species of the genus Dreyfusia have been established: D. nordmannianae Eckst. and D. prelli Grosm. on Abies nordmanniana Link. and Picea orientalis Link. and Dreyfusia nov. spec. on A. nordmanniana. The latter species, earlier known from Greece and the Balkans, but not yet described is most probably unholocyctic on fir.In the Zigana mountains between Trabzon and Gümüsane a species producing plenty galls on young P. orientalis trees and closely resembling D. merkeri Eichh. has been observed. Further investigations are necessary to evaluate whether or not it lives unholocyclic on spruce.In the West Pontic mountains near Bolu D. nordmannianae was intensively studied in twig infestations on A. bornmülleriana Mattf. The pattern of attack differs greatly from the one experienced in central Europe in that almost exclusively the current year's shoots are attacked whereas the leader and the older twig axis remain free of attack. In Europe all parts of the trees are colonized.Considering the known facts about generation- and morph sequences, morph determination and phenological behaviour of D. nordmannianae the characteristics of their population dynamics in Turkey and the interactions of the curious mode of attack and the extraordinary efficiency of their special predators have been completely evaluated.     

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.     

Epidemiology of Heterobasidion abietinum and Viscum album on silver fir (Abies alba) stands of the Pyrenees

In the last two decades, stand decline and increased mortality has affected silver fir (Abies alba) forests in the Spanish Pyrenees. Simultaneously severe occurrences of the root rot fungus Heterobasidion annosum s.l. and of the mistletoe Viscum album have been reported. We aimed to improve the understanding of the epidemiology of both pathogens in our region. All H. annosum isolates found on silver fir were typed as H. abietinum. H. abietinum was more frequently observed where cuttings had targeted fir trees rather than other species. H. abietinum fruiting bodies were observed in the most recently cut stumps. V. album was more abundant on more dominant fir trees, and in southern aspect stands. The number of V. album colonies in the stand correlated (R² = 0.40) with silver fir mortality. Stands with a high level of V. album infection tended to have a smaller percentage of basal area in species other than silver fir, and they tended to be located on more south‐facing slopes. H. abietinum was widespread in silver fir forests of the Pyrenees. Our data suggest that, in the Pyrenees, the observed H. abietinum incidence may represent a combination of both primary and secondary spread of the pathogen. Favouring mixed forests should be tested as a potential control method for V. album. The correlation between silver fir mortality and V. album infection warrants further study, as the observed tree mortality could have occurred due to other factors than V. album, such as drought damage.     

Tree mortality and habitat shifts in the regeneration trajectory underneath canopy of an old-growth subalpine forest

To better understand tree regeneration trajectories and the resultant coexistence of Abies with co-dominants, Picea jezoensis var. hondoensis, Tsuga diversifolia and Betula ermanii, in an old-growth subalpine forest, we investigated spatial mortality patterns during the regeneration of Abies mariesii and A. veitchii, which are abundant in the understory reflecting their shade tolerance. Regeneration of these Abies spp. from shaded understory to canopy status is affected by other canopy co-dominants. Snags of understory Abies spp. were common, suggesting that the primary mortality agent is suppression by the overstory. Although live, small Abies trees in the understory were positively associated with a Picea canopy, the long-term survival was reduced among Abies trees close to the canopy, suggesting that shading by large Picea in the overstory negatively affects understory Abies plants. The existence of shade-intolerant canopy co-dominants such as Picea and also Tsuga, which are larger and longer lived than the shade-tolerant Abies, may play an important role in preventing the Abies spp. from competitively displacing these other tree species, which are much rarer in the understory, though common in the canopy. Moreover, in spite of the fact that Betula canopies fostered recruitment and growth of Abies saplings, Abies showed no association with Betula canopy and their survival at later-stage was rather reduced near or beneath Betula canopies at the subsequent understory small tree stage. Based on spatially significant events related to tree death, this study detected such “habitat shifts” in the trajectory of tree regeneration. Accordingly, it can be concluded that careful consideration of the regeneration habitat is required for a fuller understanding of ecological processes in spatially complex old-growth forest systems.     

Fossil history and modern distribution of the genus Abies (Pinaceae)

The plants of the genus Abies are dominant and key species in the dark coniferous forest in the Northern Hemisphere. There are 52 species, one subspecies and 12 varieties in the genus Abies in the world. The history and modern distribution of Abies were discussed. The genus has three modern distribution centers: South Europe, North America and East Asia. These areas are also rich in fossil records. The vertical distribution regions of Abies are from sea level to an elevation of 4,700 m, concentrated in 1,000–2,000 m (15 species). In China, the genus distributes in 20 provinces, especially abundant in the Hengduan Mountains. Meanwhile, endemic and relic phenomena are obvious in this genus. There are seven relic species with both limited individuals and limited distributed regions. Based on the fossil records and the latest phylogenetic data, the following hypothesis was proposed: Abies originated from the middle and high altitudes of the Northern Hemisphere in the middle Cretaceous and it was dispersed southward in the Eocene due to global climate cooling down. The distribution of Abies was deeply influenced by geological events such as the uplift of Himalaya, the Alps, the Rocky Mountains, the occurrence of the Asian Monsoon as well as the Quaternary glaciers. Finally, the current distribution pattern appeared in the Quaternary. The genus Abies has a fossil history and modern distribution pattern similar to that of Cathaya and Pseudolarix. __________ Translated from Acta Botanica Yunnanica, 2006, 28(5): 439–452 [译自: 云南植物研究]