Comparing Chilean and Swedish rust populations: possible impact of long‐distance transport of Melampsora rust

As willow (Salix) cultivation grows globally, worldwide monitoring of pathotypes for Melampsora rust, the most serious disease in willow, becomes increasingly important. We compared the pathotype composition of the Melampsora rust population in Chile with that in Sweden and assessed inocula exchange for the rust between the continents. For pathotyping rust isolates, a willow differential was used (a standardized set of willow test clones) to obtain clone‐virulence patterns. These patterns were used to identify virulence components involved in determining which willow clones a given individual pathogen isolate (genotype) can successfully attack. Each virulence component detected earlier in Sweden's rust population was present in Chile's rust population. Using isoenzyme analysis to classify willow species in Chile, a low genetic variation of Salix viminalis (an introduced willow species) was found, compared with the endemic species Salix humboldtiana. Comparison of rust populations for the two countries supports the present hypothesis that intercontinental inocula exchange can be a significant determinant of local pathotype structure, and consequently can be important for willow‐resistance breeding.     

Effect of plantation design on stem‐infecting form of rust in willow biomass coppice

Planting of genotype mixtures is a major strategy for the non‐chemical control of willow rust in short‐rotation coppice willows grown for biomass. In 1997 and 1998, the relative severity and spatiotemporal distribution of stem canker damage by Melampsora spp. on Salix viminalis `Bowles Hybrid' in a field trial containing monocultures and two design mixtures (random or regular) of three or five genotypes were assessed. Sphaerellopsis filum, a hyperparasite of Melampsora spp. was detected in all plots during the summer (1998) between the time of the two canker assessments. The percentage of rust pustules with this hyperparasite was initially greater in monoculture plantings than in mixtures in June, but by July, more than 92% of pustules were parasitized, irrespective of plantation design. The stem rust assessments showed that, in general, there were fewer cankers per stool in plots containing mixtures of willow genotypes compared with monoculture plantings. The distribution of rust cankers was different between monocultures and willow mixture designs, with differences between years for monocultures.     

Rust severity in bioenergy willow plantations treated with additional nutrients

A 3‐year field study was carried out to determine the effect of wastewater irrigation and previous differences in mineral fertilization on the occurrence of willow leaf rust (Melampsora epitea). The experiment was conducted in two energy forest plantations: one designed for wastewater purification and the other as a mineral fertilization experiment. The severity of leaf rust on different clones and sites with different treatments was assessed by counting the number of uredinia per leaf unit area. Generally, plants irrigated with wastewater consistently had more leaf rust, irrespective of the study years or willow clones. Previous mineral fertilization had mixed effects on different clones 2 years after the last application. Three years after the last fertilizer application, however, no impact of the treatment on rust disease development was detected. In general, the rust levels differed from year to year probably due to climate. In this study, no correlation was detected between shoot age and rust severity, whereas climate and treatments strongly influenced leaf rust levels on some willow clones.     

Interactions between Melamspora larici‐epitea pathotypes and the mycoparasite Sphaerellopsis filum from willow rusts

Five pathotypes of the willow rust Melamspora larici‐epitea were inoculated with 12 isolates of Sphaerellopsis filum derived from Melampsora species/forms occurring on willows. On average, 20.5% uredinial pustules produced S. filum pycnidia and rust spore production was reduced by 38.4% on leaf discs inoculated with S. filum. Some rust isolates were more readily infected by S. filum than others while some S. filum isolates caused higher levels of infections than other S. filum isolates. In general, the suppressive effects of these S. filum isolates on rust spore production were similar on the majority of rust pathotypes tested. There appeared to be a positive link between the rust pustule area and the rate of infection by S. filum. Sphaerellopsis filum inoculum densities were positively correlated with the reduction in rust spore production but not with the number of rust pustules. Implications from the results were discussed in relation to the deployment of S. filum in biological control of willow rust in willow mixture plantations which harbour more diverse rust pathotypes compared with monocultures.     

Rust scorings in a plantation of Salix viminalis clones during ten consecutive years

Among pests and diseases occurring in coppiced willow plantations established to produce biofuel, Melampsora rust is the most widespread and severe. In the present study, the variation in rust attacks in a set of Salix viminalis clones was investigated. The aim was to look for trends possibly arising from changes in rust genotype composition, to get some idea about the risks of rust overcoming resistance bred into new willow varieties. Rust levels on 92 S. viminalis clones belonging to 18 different families were scored once each year during 1987–96 in a plantation in southern Sweden. Rust scorings in 1993 deviated most from the average scoring pattern of the clones. Furthermore there was a successive change, such that the longer the time interval between scorings the lower was the correlation between rust scores. Willow families were identified that deviated from the others over time. Various explanations for inconsistency in rust scores over time are discussed.     

Genetic diversity of the hyperparasite Sphaerellopsis filum on Melampsora willow rusts

The non‐specific rust hyperparasite Sphaerellopsis filum occurs naturally on Melampsora rusts of many species of the genus Salix as well as on a large range of other rust genera worldwide. To study the genetic diversity of the hyperparasitic fungus 77 S. filum isolates collected from rusts on willow clones from plantations, clone collections and natural habitats of different sites were investigated using polymerase chain reaction ‐ restriction fragment length polymorphism (PCR‐RFLP) analysis of the rDNA internal transcribed spacer regions including 5.8S rDNA and sequence analysis. Additionally, strains from Melampsora poplar rusts (4) and strains of Puccinia abrupta from Parthenium hysterophorus (5) and of P. obscura from Bellis perennis (1) were used for comparisons. Results of genetic analysis demonstrated distinct variation within the S. filum isolates. Two main groups with more than 32% difference between their nucleotide sequences were distinguished, indicating two taxa within S. filum. Within the first main group three profiles (I, II, III) were detected. The differences between these profiles were about 12%. The variation within each profile was very low (less than 2%). The second main group comprised two profiles (IV, V), which differed in 12 to 16% of their nucleotide positions. The isolates of group IV possessed a higher variation (up to 5%) within the group than those of the first main group (I, II, III). Group V was only represented by a single isolate. Neither interrelations between the S. filum profiles and the Melampsora genotypes nor a spatial distribution could be detected. It is remarkable that the six strains of S. filum from Puccinia rusts belong to one subgroup.     

A new species of Melampsora rust on Salix elbursensis from Iran

A rust fungus was found causing stem cankers on 1‐ to 5‐year‐old stems of Salix elbursensis in the north west of Iran. The rust also forms uredinia on leaves and flowers of the host willow. Light and scanning electron microscopy revealed that the new rust is morphologically distinct from several Melampsora species occurring on the willows taxonomically close to S. elbursensis, but indistinguishable from Melampsora larici‐epitea. Examination of the internal transcribed spacer (ITS) region of the ribosomal DNA suggested that the rust fungus is phylogenetically close to Melampsora allii‐populina and Melampsora pruinosae on Populus spp. Based on both the morphological characteristics and the ITS sequence data, the rust is described as a new species –Melampsora iranica sp. nov.     

Compatible and Incompatible Reactions of Melampsora Rust on Willow Leaves

Spore germination, growth and appresssoria formation for Melampsora willow rust were studied for compatible and incompatible hosts to elucidate penetration specificity and prehaustorial events in the infection process. Rust originating from Salix dasyclados was inoculated on a compatible (S. dasyclados) and an incompatible (Salix daphnoides) clone and, for comparison, on the non-host plants tomato, tobacco and poplar. The same experimental procedure was also carried out for rust originating from poplar. Rust development was followed for 5 days at 20 h intervals using a microscope with fluorescent equipment. The study showed that the Melampsora rust can develop on, and penetrate, the leaf regardless of whether the rust was inoculated on compatible or incompatible willow plants. However, the fungus was able to infect and reproduce itself only in compatible interactions. For willow rust in the prehaustorial stage, the study indicated that specific recognition mechanisms were unnecessary to start the infection process.     

Life cycle of Melampsora larici‐epitea on short‐rotation cultures of Salix burjatica

The host alternation of Melampsora larici‐epitea occurring on Salix burjatica in experimental short‐rotation cultures in Finland was studied by inoculation tests. Larix sibirica was tested as the most probable alternate host. It was inoculated with rust basidiospores, and the life cycle of the pathogen was succesfully completed by inoculating the willow with aeciospores developed on larch needles.     

European pear rust on Juniperus excelsa L. in south‐western Turkey

In this study, the frequency of European pear rust (Gymnosporangium fuscum) lesions on Juniperus excelsa was investigated both on seedlings in a nursery and on forest trees in six juniper stands located in south‐western Turkey. In the nursery, lesions occurred on 0.7% of 5510 1‐year‐old seedlings and 29% of 200 2‐year‐old potted seedlings. In each of the juniper forests, 50 trees were chosen within a square of approximately 400 m² in size. In general, rust lesions were common on the trees. On average, 49.0% of the trees had lesions; on 17.3% of the trees, the lesions were located on the trunk, on 43.3% on the branches and on 11.7% on both. In lesion‐bearing trees, the average number of lesions was 1.47–3.59 and the maximum number of lesions per tree were six to 11. Most of the lesions (85.1%) were on branches. There were significant differences between the stands, which can be attributed to different age of the trees, site factors and the presence of alternate hosts. The rust identified from stem lesions in forest trees and from nursery seedlings was European pear rust (G. fuscum), but possible co‐occurrence of other Gymnosporangium spp. cannot be excluded.     

川东北地区杨树与柳树锈病发生与病理学观察

杨树和柳树均属于杨柳科植物,两者在经济林及绿化行道树上的运用越来越广泛。锈病在杨树和柳树高栽植密度苗圃地和成片用材林的发生极为严重。本研究比较杨树和柳树锈病的发病症状、流行特点及病原菌特征,为该病的防治提供理论和实际指导依据。以川东北地区4区(县)(南充市高坪区和西充县,广安市前锋区和武胜县)患发锈病的杨树和柳树为研究对象,采用流行病学调查和显微观察技术等方法。杨树锈病和柳树锈病大都侵害叶和幼苗,引起树叶早期脱落,幼苗枯萎。但杨树锈病的病原菌为锈菌科栅锈菌属松杨栅锈菌(Melampsora larici-populins),柳树锈病的病原菌为锈菌科栅锈菌属拟鞘锈栅锈菌(Melampsora coleosporioides Diet.)。杨树锈病和柳树锈病虽有共同的发病特征,但在为害症状、病原类型和发生规律方面均有很大的不同。建议在防治时根据寄主的不同选择针对性强的防治方法。     

Phylogeographic patterns of Armillariaostoyae in the western United States

Nuclear ribosomal DNA regions (i.e. large subunit, internal transcribed spacer, 5.8S and intergenic spacer) were sequenced using a direct‐polymerase chain reaction method from Armillaria ostoyae genets collected from the western USA. Many of the A. ostoyae genets contained heterogeneity among rDNA repeats, indicating intragenomic variation and likely intraspecific hybridization. Intragenomic variation was verified by visually editing base‐sequence offsets in regions with insertions/deletions, and using sequence‐specific internal primers to resequence heterogeneous regions. Phylogenetic analyses with Bayesian Inference methods were used to define groups within A. ostoyae. Analysis of A. ostoyae from outside the western USA indicated the presence of a Circumboreal group of A. ostoyae that also occurs in Utah; two other phylogeographic groups were associated with the Rocky Mountain and Pacific Northwest regions of the USA. Mixed sequence types, an indication of intraspecific hybrids, were common in some geographic regions. Hybridization events may have influenced species evolution, contributing to variation in pathogenicity and virulence. The occurrence of these groups and intraspecific hybrids also indicates that paleogeography and paleoclimate may have influenced the phylogeography of A. ostoyae. In addition, other Armillaria species were examined for evolutionary relationships with the groups of A. ostoyae. These findings will provide a basis for future research relating ecological function to genetic diversity within A. ostoyae.     

Experimental testing of rust fungus‐mediated herbivory resistance in Betula pendula

Correlative evidence suggests that pathogenic leaf rust fungus, Melampsoridium betulinum, in late summer may negatively affect folivorous insects in the following summer. Correlative association does not necessarily, however, reveal causality. Alternatively, other interconnected plant characters may determine rust densities and herbivore performance. In this study, we used birch clones and rust fungus inoculations to manipulatively test the effects of birch rust and birch genotypes on the growth performance of folivorous moth larvae of Epirrita autumnata (Lep. Geometridae) in the subsequent year. The inoculation treatment increased rust densities (three‐ to 60‐fold) compared with natural infection levels. E. autumnata performance varied among birch clones and showed 4% lower growth performance on rust‐inoculated trees. However, the larval performance did not differ between rust‐treated shoots and untreated control shoots and the use of tree‐specific rust densities as a covariant in statistical analyses failed to reveal any negative association between rust fungus and larval performance. As the slight difference in larval growth performance also levelled until pupation, we propose that rust infection has biologically insignificant importance to the performance of E. autumnata.     

The risk to Myrtaceae of Austropuccinia psidii,myrtle rust,in Mexico

Austropuccinia psidii is a biotrophic rust fungus that affects species from the Myrtaceae family. In Mexico, Myrtaceae is widely distributed in temperate, tropical and semi‐arid ecosystems, and includes 20 genera and 192 endemic and exotic species. Austropuccinia psidii has been present in Mexico for the last four decades; however, little is known about the distribution of this rust or the vulnerability of native and exotic Myrtaceae to infection. In this study, we used global occurrence records for the pandemic biotype of myrtle rust to model its current and future suitable habitat using a species distribution model, Maxent. We identified regions that are highly suitable for myrtle rust establishment, now and in the future (2050). Additionally, we identified the Myrtaceae species known to be susceptible to rust infection and that are currently distributed in areas with high rust habitat suitability. Thirty‐six susceptible plant species and 142 untested species are distributed within areas of suitable rust habitat and are considered potentially at risk of rust infection. Current suitable habitat is mainly restricted to the east coast of Mexico, with Veracruz, Puebla, Chiapas, Tabasco and Oaxaca being the most vulnerable regions to the rust under current and future climates. We encourage monitoring within these regions by surveying locations where the rust occurs and within areas with high suitable habitat to determine the threat to native ecosystems and industries reliant on Myrtaceae. We also recommend screening to test the susceptibility of Myrtaceae species with no known susceptibility rating.     

Short rotation coppice willow in Northern Ireland since 1973: development of the use of mixtures in the control of foliar rust (Melampsora spp.)

A review is given of the use of clonal mixtures in the development of disease control strategies in short rotation coppice willow in Northern Ireland. Salix burjatica‘Korso’ had been grown successfully for over 10 years when, in 1985, Melampsora epitea var. epitea caused serious problems. Although fungicides were effective, their use was not practical for environmental and economic reasons. Therefore, in 1987 large scale field experiments were initiated to investigate the use of clonal mixtures as a disease control strategy. Increased yields were consistently recorded from mixed stands when compared to either the mean yield of component clones or the individual yields of any of the component clones grown in monoculture. Part of this increased yield was due to a reduction in the impact of rust disease. Investigations are currently being conducted on the effect of number of clones within a mixture, the best clones to use and the optimum planting density. It is essential that there is a range of susceptibilities to M. epitea var. epitea pathotypes within the components of the mixture.     

Co‐occurrence of the ascomycete Lophodermium piceae and the rust fungus Chrysomyxa abietis in Norway spruce needles

The frequency of needles and the proportion of needle segments infected by Lophodermium piceae were compared in symptomless and Chrysomyxa abietis‐infected, 1‐year‐old needles of Picea abies. In late spring, symptomless needles from both rust‐infected and healthy saplings were sampled. In addition, rust‐infected, totally chlorotic needles and needles with chlorosis along about half their length from the diseased trees were examined. In all three stands, the proportion of segments infected by L. piceae was larger in the rust‐infected half of the needle than in the symptomless half; but the difference was statistically significant in only one of the stands. The proportion of L. piceae‐infected segments among the nonrust‐infected needles was the same as that found for the uninfected half of rust‐infected needles (after correction for size differences). No differences in the proportion of L. piceae‐infected segments were found between the totally chlorotic, rust‐infected needles and the green needles of diseased or healthy trees.     

Effects of host plant hybridization on resistance to willow leaf rust caused by Melampsora sp.

Patterns of resistance to willow leaf rust caused by Melampsora sp. were measured on Salix eriocephala, Salix sericea, and their interspecific hybrids in natural field populations in 1991–94 and on potted plants in common garden experiments in 1994 and 1995. Testing was carried out to establish if the resistance of hybrids was intermediate between pure parental species (additive), equal to one of the parents (dominant), lower than both parents (susceptible), higher than both parents (resistant), or not different than either parent (null hypothesis). Support was generally found for hybrid susceptibility in the field and common garden experiments, with the null hypothesis of no difference having some support. The results from the common garden study suggest that the susceptibility of hybrids seen in the field is due to genetic differences. Hybridization between these species of willows results in increased susceptibility to this pathogen. The variation among years in the relative susceptibility of hybrid and parental willows may be due to environmental effects on the willows or to shifts in the abundance of Melampsora races.     

Climatic favourability zones for Eucalyptus rust in Brazil

Brazilian forest‐based industries are supported by more than 5.5 million hectares of Eucalyptus growing under different climatic conditions with different degrees of favourability for rust Puccinia psidii, including both traditional and expanding areas, where such disease is of major concern for the industry, foresters and scientists. The main objective of this study was to define favourable climatic zones for Eucalyptus rust in Brazil with the following aims: (i) to develop a spatial method for estimating the mean night‐time temperature (T_ng); (ii) to assess and validate a Eucalyptus rust model; and (iii) to map Eucalyptus rust favourability zones in Brazil based on the proposed model. A straightforward method, based only on latitude, day of the year, maximum and minimum air temperatures, was developed to estimate T_ng, which is a key variable for a proper application of Ruiz rust model. Based on 37 field experiments with natural rust occurrence, it was observed that climatic conditions are determining factors for disease severity. Significant correlations between disease severity observed in the field and climatic conditions were found: a negative correlation with temperature (r = .50, p < .01) and a positive correlation with relative humidity (r = .89, p < .0001). A significant correlation (r = .81, p < .0001) between the normalized infection index, produced from Ruiz model, and the average rust score was also observed. Once the model was validated under field conditions, it was applied using historical average data of air temperature and leaf wetness duration to obtain monthly Eucalyptus rust favourability maps for the whole country. These final maps show that favourable climatic zones for Eucalyptus rust are extremely dynamic, with high temporal and spatial variability in Brazil and that climatic conditions should be considered for expansion of Eucalyptus to new areas, in breeding programmes, and for defining the most suitable seasons for forest establishment in each climate of the country. These results provide forestry managers with practical tools to reduce uncertainty about the expected severity of Eucalyptus rust in Brazil.     

Effect of Betula pendula clone and leaf age on Melampsoridium betulinum rust infection in a field trial

Ten 5‐year‐old Betula pendula clones were studied for their rust resistance in the field. The trees were treated by inoculating 10 leaves on a shoot with Melampsoridium betulinum urediniospore suspension or spraying the control leaves with water. The birch clones differed significantly in their resistance to M. betulinum leaf rust fungus and the clones also varied in their responses to the local rust strain and the inoculated rust strains. However, natural rust infections and inoculation treatment were positively correlated. The older leaves had fewer infections than the younger ones on the tip of the shoot in the control trees, but in the inoculation treatment no significant correlation was found between the leaf ages and rust infection. The factors behind the different leaf susceptibilities are discussed.     

Blister rust in maritime pine

Maritime pine (Pinus pinaster Ait.) seedlings from four stands of central Italy were inoculated with a suspension of 75,000 basidiospores/ml. Significant differences among the stands were found for the following traits: percentages between nonspotted and inoculated seedlings; and percentages between seedlings with spermagonia of Cronartium flaccidum and spotted seedlings. Genetic variation in blister rust resistance among the stands and among the families within a stand confirms the validity of the idea to select the stands which are exposed to heavy rust infection and then within the selected stands to choose trees able to supply the highest possible number of nonspotted seedlings and seedlings with needle spots but without spermagonia.