Plant-enemy interactions on islands are shaped by ecological and evolutionary processes during the history of the islands. It is, however, rare to be able to observe the foundational phases on islands, but opportunities exist in areas with land-rising coastlines, such as in parts of the Baltic Sea. Using several plant-enemy systems, a pattern appears where low-defended plants are typically dominating on young islands and on shores of older islands. One reason for this pattern is that the main seed sources are plants from sites with low enemy attack rates and with high reproductive output. Later on in succession, once herbivore populations become established and attack rates increase, selection acts to increase plant resistance traits. Even though the number of studied plant species is still low, it is apparent that processes during foundational phases may be complex and simplified explanations for current patterns may be hard to find.

Fungi and substrates undergoing fungal decomposition were collected from forests in northern and southern Sweden and analyzed for chlorinated dimethoxybenzenes (DMBs). Specimens were fungi fruiting bodies, rotting wood, forest litter and underlying humus. Targeted compounds were DAME (1,2,4,5-tetrachloro-3,6-DMB) and related fungal secondary metabolites. A screening procedure was developed which involved soaking the specimens in ethyl acetate followed by analysis by capillary gas chromatography – mass spectrometry with mass selective detection (GC-MSD). DAME was the most frequently found (62% of 47 specimens) and often the most abundant target compound, with range and mean ± SD concentrations of <0.0017–3.81 and 0.21 ± 0.63 mg kg−1 ww. Based on log-log correlations of partition coefficients of hydrophobic compounds between fungal biomass/water (KD) and octanol/water (KOW), five species of fungi are suggested to produce DAME de novo versus bioaccumulation from forest runoff water. Full-scan mass spectra of some high-concentration specimens indicated the presence of a Cl2DMB and a Cl3DMB, which could not be identified further due to lack of standards, and drosophilin A (DA = 2,3,5,6-tetrachloro-4-methoxyphenol), the precursor to DAME. Tetrachloroveratrole (TeCV = 1,2,3,4-tetrachloro-5,6-DMB) was found in only a few specimens. This study supports our hypothesis of fungi as a source of DAME in terrestrial runoff and indicates that other chlorinated secondary metabolites are present. DAME is widely distributed globally, and it would be good to have a better understanding of its sources and pathways as a marker of terrestrial organochlorines and their availability for bioaccumulation.

Introduction Thousands of halogenated natural products (HNPs) are generated in the ocean and on land. A subset of these, halomethoxybenzenes (HMBs), are released from both natural and anthropogenic sources. Here we consider: 1. Brominated anisoles (BAs), transformation products of bromophenols. 2. Drosophilin A methyl ether (DAME: 1,2,4,5-tetrachloro-3,6-dimethoxybenzene), a secondary metabolite of terrestrial fungi. 3. Tetrachloroveratrole (TeCV: 1,2,3,4-tetrachloro-5,6-dimethoxybenzene), a lignin byproduct found in bleached kraft mill effluent. 4. Pentachloroanisole (PeCA), a metabolite of the wood preservative pentachlorophenol. Methods We examined several ecosystem compartments to determine sources and exchange processes for these HMBs: air, precipitation, rivers, forest fungi and litter, and water from northern Baltic estuaries and offshore. Samples were analyzed for HMBs by capillary gas chromatography – quadrupole mass spectrometry. Results and discussion All four types of HMBs were found in air, and BAs, DAME and TeCV were also present in precipitation. BAs and DAME were common in rivers and estuaries, whereas TeCV was low and PeCA was below detection. DAME was identified in several species of fungi and in forest litter; TeCV was occasionally present, but BAs and PeCA were below detection. Concentrations of BAs were higher in estuaries than in rivers or offshore waters, showing that estuaries are hot spots for production. BAs were negatively or not correlated with chlorophyll-a, suggesting contribution by heterotrophic bacteria as well as known production by phytoplankton and macroalgae. DAME was negatively or not correlated with BAs and did not appear to be produced in the estuaries; fungi and forest litter containing fungal mycelia are suggested as sources. HMBs volatilize from sea and land, disperse through the atmosphere, and return via precipitation and rivers. Production and biogeochemical cycles are influenced by climate change and we suggest BAs and DAME for following partitioning and exchange processes.

Halomethoxybenzenes (HMBs) are a group of compounds with natural and anthropogenic origins. Here we extend a 2002–2015 survey of bromoanisoles (BAs) in the air and precipitation at Råö on the Swedish west coast and Pallas in Subarctic Finland. New BAs data are reported for 2018 and 2019 and chlorinated HMBs are included for these and some previous years: drosophilin A methyl ether (DAME: 1,2,4,5-tetrachloro-3,6-dimethoxybenzene), tetrachloroveratrole (TeCV: 1,2,3,4-tetrachloro-5,6-dimethoxybenzene), and pentachloroanisole (PeCA). The order of abundance of HMBs at Råö was ΣBAs > DAME > TeCV > PeCA, whereas at Pallas the order of abundance was DAME > ΣBAs > TeCA > PeCA. The lower abundance of BAs at Pallas reflects its inland location, away from direct marine influence. Clausius-Clapeyron (CC) plots of log partial pressure (Pair)/Pa versus 1/T suggested distant transport at both sites for PeCA and local exchange for DAME and TeCV. BAs were dominated by distant transport at Pallas and by both local and distant sources at Råö. Relationships between air and precipitation concentrations were examined by scavenging ratios, SR = (ng m−3)precip/(ng m−3)air. SRs were higher at Pallas than Råö due to greater Henry's law partitioning of gaseous compounds into precipitation at colder temperatures. DAME is produced by terrestrial fungi. We screened 19 fungal species from Swedish forests and found seven of them contained 0.01–3.8 mg DAME per kg fresh weight. We suggest that the volatilization of DAME from fungi and forest litter containing fungal mycelia may contribute to atmospheric levels at both sites.

Marine macroalgae are used worldwide for human consumption, animal feed, cosmetics and agriculture. In addition to beneficial nutrients, macroalgae contain halogenated natural products (HNPs), some of which have toxic properties similar to those of well-known anthropogenic contaminants. Sixteen species of red, green and brown macroalgae were collected in 2017-2018 from coastal waters of the northern Baltic Sea, Sweden Atlantic and Norway Atlantic, and analyzed for bromoanisoles (BAs) and methoxylated bromodiphenyl ethers (MeO-BDEs). Target compounds were quantified by gas chromatography-low resolution mass spectrometry (GC-LRMS), with qualitative confirmation in selected species by GC-high resolution mass spectrometry (GC-HRMS). Quantified compounds were 2,4-diBA, 2,4,6-triBA, 2'-MeO-BDE68, 6-MeO-BDE47, and two tribromo-MeO-BDEs and one tetrabromo-MeO-BDE with unknown bromine substituent positions. Semiquantitative results for pentabromo-MeO-BDEs were also obtained for a few species by GC-HRMS. Three extraction methods were compared; soaking in methanol, soaking in methanol-dichloromethane, and blending with mixed solvents. Extraction yields of BAs did not differ significantly (p > 0.05) with the three methods and the two soaking methods gave equivalent yields of MeO-BDEs. Extraction efficiencies of MeO-BDEs were significantly lower using the blend method (p < 0.05). For reasons of simplicity and efficiency, the soaking methods are preferred. Concentrations varied by orders of magnitude among species: ∑2BAs 57 to 57 700 and ∑5MeO-BDEs < 10 to 476 pg g-1 wet weight (ww). Macroalgae standing out with ∑2BAs >1000 pg g-1 ww were Ascophyllum nodosum, Ceramium tenuicorne, Ceramium virgatum, Fucus radicans, Fucus serratus, Fucus vesiculosus, Saccharina latissima, Laminaria digitata, and Acrosiphonia/Spongomorpha sp. Species A. nodosum, C. tenuicorne, Chara virgata, F. radicans and F. vesiculosus (Sweden Atlantic only) had ∑5MeO-BDEs >100 pg g-1 ww. Profiles of individual compounds showed distinct differences among species and locations.

Summary A long‐term study of disease dynamics caused by the rust Uromyces valerianae in 31 discrete populations of Valeriana salina provides a rare opportunity to explore extended temporal patterns in the epidemiology of a natural host–pathogen metapopulation. Over a 28‐year period, pathogen population dynamics varied across the metapopulation with disease incidence (presence/absence), prevalence (% plants infected) and severity (% leaf area covered by lesions) all showing strong population and year effects, indicative of heterogeneity among years and host populations in the suitability of conditions for the pathogen. Disease incidence within individual host populations was significantly affected by host population size, disease prevalence the previous year and the proximity of neighbouring populations infected in the current year. After accounting for these variables there was still a marked temporal component with winter sea level having a significant effect; as did summer rainfall in the second part of the study period (1997–2011). Disease prevalence was also effected by host population size and disease prevalence in the previous year. However, it was less affected by spatial aspects of disease spread than was disease incidence. Winter sea level and June rainfall significantly affected disease prevalence. Assessment of disease impact on plant performance found strong variation in disease severity associated with the aspect and positioning of host populations. Plants growing in lower disease environments produced significantly more seeds than those growing in high disease sites. Significant variation in reaction to infection by U. valerianae was detected among plants within four populations and between these different populations. Synthesis. The epidemiology of Uromyces valerianae was highly influenced by host population size, previous disease and distance. After accounting for these factors, there was a clear temporal signal of change in disease incidence linked to winter sea level and summer rainfall. These patterns reinforce the importance of considering interactions in multiple populations over long periods of time in order to obtain a clear picture of the variability in disease‐induced selection pressures across time and space. The behaviour of the pathogen fitted that predicted for a metapopulation with considerable asynchrony in epidemiological patterns among demes.

The importance of disease in modulating ecosystem responses to climate change remains poorly understood. A seven-year study of the effects of increased snow cover on tundra plant communities in Sweden found that, although plant growth was favoured by increased snow, biomass and carbon-balance trends were reversed by a pathogen outbreak.

Modern forestry practices have decreased the abundance of coarse woody debris (CWD), and as a result many species that depend on dead wood are now threatened. This implies a need to develop forestry practices that maintain biodiversity. We examined the conservation value of experimental spruce logs (control logs, logs placed in natural shade, and cut tree tops) for wood-inhabiting fungi in two forest stands, one nature reserve and one mature managed forest, in each of seven forest areas in northern Sweden. Here we report the initial findings of the experiment that was established in winter 2001-2002 and data were collected in 2002, 2003 and 2006. A pre-inventory of the local species composition in 2002 revealed a higher per area species richness, including red-listed species, in reserves than in managed forests. Ordination analyses of the experimental logs showed a significant effect of area, while not of stand type in 2003. ANOVA analysis showed no significant effect of stand type on species richness and abundance. In 2006, the species assemblage started to differentiate between the two stand types and forest age, forest site type (moisture), and distance to forest reserves, all explained part of the variation, whereas the amount of CWD, and species composition at the start of the experiment only showed a marginal effect.The early successional fungal community was dominated by two functional groups, humus-decayers and white-rot species, both characterized by a rapid, early colonization and fruit-body formation on the competition-free new substrate. A similar positive response to the new substrate was also observed for the mycorrhizal species in 2006. The high frequency and early appearance of humus-decayers and mycorrhizal species that do not primarily depend upon CWD for their nutrition suggest that their formation of fruiting bodies is limited by substrate availability. Thus some mycorrhizal fungi are apparently rare due to lack of suitable substrate for fruit-body formation.Evidence of dispersal-limitation was observed in 2006. Fomitopsis pinicola, an early colonizer in boreal forests, playing a key role for other wood-inhabiting organisms, colonized significantly more logs in the reserve stands compared with the managed stands. Our data demonstrate that lack of CWD strongly affects both species that depend upon wood for nutrition and species that depend upon wood for fruit-body formation. Thus some species may show an apparent rarity due to lack of suitable substrate. We conclude that creation of CWD appears to be a useful method to maintain or restore fungal diversity in boreal coniferous forests. (C) 2010 Elsevier Ltd. All rights reserved.

1. Both evolutionary theory and empirical evidence from agricultural research support the view that asexual, vertically transmitted fungal endophytes are typically plant mutualists that develop high infection frequencies within host grass populations. In contrast, endophyte–grass interactions in natural ecosystems are more variable, spanning the range from mutualism to antagonism and comparatively little is known about their range of response to environmental stress.

Pathogen genes involved in interactions with their plant hosts are expected to evolve under positive Darwinian selection or balancing selection. In this study a single copy avirulence gene, AvrP4, in the plant pathogen Melampsora lini, was used to investigate the evolution of such a gene across species. Partial translation elongation factor 1-alpha sequences were obtained to establish phylogenetic relationships among the Melampsora species. We amplified AvrP4 homologues from species pathogenic on hosts from different plant families and orders, across the inferred phylogeny. Translations of the AvrP4 sequences revealed a predicted signal peptide and towards the C-terminus of the protein, six identically spaced cysteines were identified in all sequences. Maximum likelihood analysis of synonymous versus non-synonymous substitution rates indicated that positive selection played a role in the evolution of the gene during the diversification of the genus. Fourteen codons under significant positive selection reside in the C-terminal 28 amino acid region, suggesting that this region interacts with host molecules in most sequenced accessions. Selection pressures on the gene may be either due to the pathogenicity or avirulence function of the gene or both.

A major challenge in conservation biology is to understand species’ responses to habitat loss. In Fennoscandia, the ongoing decline in aspen in forests is of particular concern, since aspen is the boreal forest tree species that supports the most host-specific species of cryptogams and invertebrates. In order to predict the potential effects of aspen decline we compared the occurrence of three epiphytic cyanolichens in old-growth stands of the same habitat quality, in four aspen-rich and four aspen-poor landscapes. Collemacurtisporum and Collemafurfuraceum were, on average, five and six times more frequent, respectively, in the aspen-rich than in the aspen-poor landscapes. Leptogiumsaturninum was not affected by the abundance of aspen stands at the landscape level. Our data suggests that lichen species with poor dispersal abilities may be more sensitive to habitat loss than more easily dispersed species and that species with broader habitat amplitude may be less sensitive to habitat loss than more specialized species, even if they have inferior dispersal ability. We conclude that (i) predictions of species occurrences at the stand level have to take account of the amount of suitable habitat at the landscape level, and (ii) predicting the responses of individual species based on life-history traits can be crucial, but cannot be based on single traits. Thus our study shows that biological value cannot be assessed on the basis of habitat quality alone and that a landscape perspective is needed for the sustainable management of specialist species.

Although phenological matching between two and three trophic interactions has received some attention, it has largely been disregarded in explaining the lack of strong cascade dynamics in terrestrial systems. We studied the response of the specialist seed predator, Paroxyna plantaginis (Tephritidae) and associated generalist parasitoids (Chalcidoidea) to controlled fertilisation of individuals of naturally growing Tripolium vulgare (Asteraceae) on four island populations (Skeppsvik Archipelago, Sweden). We consistently found evidence of nutrient limitation: fertilised plants increased their biomass, produced more capitula (the oviposition units for tephritid flies), were more at risk of attack by the tephritids, and puparia were heavier in fertilised plants. During some parts of the season tephritids became more heavily parasitized, supporting the presence of cascade dynamics, however net parasitism over season decreased in response to nutrient addition. We found no evidence that capitulum size complicated parasitoid access to the tephritids, however the extended bud production prolonged the flowering season. Thus, tephritids utilized the surplus production of capitula throughout the entire season, while parasitoids did not expand their oviposition time window accordingly. Implications for top down regulation and cascade dynamics in the system are discussed.

Partial beta-tubulin 1 sequence data were obtained for 80 taxa of Pucciniaceae, with hosts from 33 angiosperm families, covering all major ordinal groups in the Angiosperm Phylogeny Group classification. As in previous studies, most species of Puccinia and Uromyces fell into two main clades (I and II), with P. glechomatis and P. psidii excluded from Pucciniaceae. Results suggest two processes; a coevolution of and hosts in each clade, as well as associated frequent jumps to ecologically close, but taxonomically distant, hosts. Clade I contained all rusts on Cyperaceae and Juncaceae, and most rusts on host orders from rosids to euasterids II. Clade II contained all rusts on Poaceae, and most on host orders from monocots to core eudicots. In both main clades, several well-supported subclades were identified. The grouping in clade I, subclade E of rusts of Cyperaceae and Asteraceae and, in particular, of an Australian isolate of P. dioicae with rusts on Australian families of Asterales, suggested a local radiation, and supported the coevolutionary relationship between rusts on these two families seen with a different range of asteraceous rusts in the Northern Hemisphere. In clade I, two clades contained only rusts of Asteraceae and Fabaceae, respectively, and in clade II, subclade F contained only rusts of pooid hosts. Rusts on non-pooid hosts were separated from them in subclade G. Other subclades contained a range of rusts on distantly related angiosperm families. Urediniospore morphology was often, but not always, correlated with the molecular phylogeny. Most rusts with urediniospores having few (1-5) equatorial germ pores were in clade I, whereas most with spores having several (5-14) scattered pores were in clade II. The distribution of telial host families on the beta-tubulin rust phylogeny was not random. Aecial hosts of heteroecious rusts played an important role in the evolutionary process. Possible examples of host jumps were seen in rusts on Geraniaceae, Polygonaceae, and Apiaceae. Despite such jumps obscuring past host associations, possible ancestral hosts were identified by the pattern of host distribution at higher taxonomic levels along the ss-tubulin phylogeny. Results suggest that clade I diverged with Cyperaceae, Juncaceae, and the more advanced core eudicot orders (rosids and asterids), whereas clade II diversified with earlier angiosperm groups, such as monocots, Poaceae, and Ranunculales. Qualified support was given to the hypothesis that rusts can reveal taxonomic relationships between their hosts, at genus, family, and ordinal levels.

Inter- and intraspecies variations in host plant traits are presumably involved in many host shifts by insect herbivores, and elucidating the mechanisms involved in such shifts has been a crucial goal in insect-plant research for several decades. Here we propose that herbivore-induced evolutionary increases in host plant resistance may cause oligophagous insect herbivores to shift to other sympatric plants as currently preferred host plants become increasingly unpalatable. We tested this hypothesis in a system based on the perennial herb Filipendula uhnaria (Rosaceae), whose herbivory defense has become gradually stronger due to prolonged selection by Galerucella tenella (Coleoptera: Chrysomelidae) herbivory in a boreal archipelago. We. show that Galerucella gradually increases its use of the alternative host plant Rubus arcticus (Rosaceae) in parallel to gradually increased resistance in Filipendula. Our results imply that, by driving the evolutionary increase in Filipendula resistance, Galerucella is also gradually making the original host species more unpalatable and thereby driving its own host-breadth enlargement. We argue that such self-inflicted "rent rises" may be an important mechanism behind host plant shifts, which in turn are believed to have preceded the speciation of many phytophagous insects.

The ForSAFE model, designed for modelling biogeochemical cycles (water, acidity, base cation, nitrogen and carbon) in terrestrial ecosystems, was modified with a vegetation response module (VEG), incorporating the effects of: nitrogen pollution, acidification, soil moisture, temperature, wind chill exposure, light and shading by trees, grazing by animals, competition between plants, above ground for light and below ground for water and nutrients. The model calculates the response of number ground vegetation plant groups. The integrated model was tested and validated at integrated level II forest monitoring sites across Sweden, four have been shown here, and used to assess the effect of acidification and nitrogen pollution in relation to factors such as climate change, forest management and changing grazing pressure. The response functions have been derived from single-factor experiments and integrated through the model structure for use on whole systems. The tests with the model suggest that the ground vegetation composition is reasonably well predicted, that much research remains before the model is fully tested and operational, and that the model may serve as a tool for assessing impacts of climate change, acid rain and forest management on plant biodiversity in forested areas.

GM trees hold promises of increased quality and yield and reduced use of herbicides and pesticides but could also have ecological consequences. We investigated whether modification of a non-defensive trait unintentionally influenced plant traits important for plant-herbivore interactions. We found that over-expression of sucrose phosphate synthase (SPS), which is known to increase mesophyll sucrose content and biomass production in GM aspens, also unintentionally induced changes in the concentration of plant phenolics and nitrogen. One of the GM lines, SPS33A, had higher concentrations of salicin, tremodulin, condensed tannins and nitrogen and tower concentrations of coumaric acid and four ftavonoids compared with the isogenic wild type. Line SPS33A was also utilized less by the leaf-beetle Phratora vitellinae than the isogenic wild type. Ecological consequences such as this are not specific to GM trees or GM plants but can occur as a result of the introduction of all introduced new varieties of crops or trees. Nevertheless, the results underline the need to consider these unexpected effects when evaluating both the potential benefits and the potential risks with GM plants, and highlight the need to establish and implement comprehensive product-by-product evaluation protocols for GM plants. (c) 2006 Gesellschaft fur Okologie. Published by Elsevier GrnbH. All rights reserved.

The two rust genera with the largest number of species are Puccinia Pers. ex Pers. and Uromyces (Link) Unger in the family Pucciniaceae (Uredinales). The hosts of these pathogens include representatives from almost all major angiosperm orders. Despite their ecological and economic importance, the status of Puccinia and Uromyces as distinct genera has been disputed, and little is known about relationships within and among these groups. Here we present phylogenetic analyses based on sequence data from the translation elongation factor 1 alpha gene for over 60 species in the family Pucciniaceae. In particular, we investigate evolutionary relationships between Puccinia and Uromyces. A relatively smaller phylogeny using the beta-tubulin 1 gene was generated to test support for this phylogeny. Two main phylogenetic clades were identified and indicate at least two radiations within the Pucciniaceae. As expected neither Puccinia s. lat. nor Uromyces s. tat. are supported as monophyletic groups by either of the protein coding genes. However, both Puccinia sensu stricto (type P. graminis), and Uromyces sensu stricto (type U. appendiculatus) constitute distinct clades. In general, members of Uromyces spp. occurred scattered throughout the phylogeny suggesting that they represent more recent radiations. Several host families are found in both of the two main clades while two families, Poaceae and Cyperaceae, are separated, with one in each of the two main clades. (c) 2006 The British Mycological Society. Published by Elsevier Ltd. All rights reserved.

In the boreal forest landscape, aspen has been effectively selected against in favour of conifers. The decrease in aspen is of particular concern, since it has more host-specific species associated with it than any other boreal tree species. Recently, forest management systems have begun to include green-tree retention in order to maintain structural diversity. Earlier studies have focused on the importance of remnant aspen trees for lichen species prevalence. We have focused on the occurrence of free-living photobionts, i.e. cyanobacteria and green-algae, since a successful establishment of sexually dispersed lichens will depend upon the presence of the photobiont. Our study shows that the abundances of Gloeocystis, Nostoc, Scytonema and Trentepohlia increased with stand age, while the abundance of Trebouxia decreased. The response to clear-felling differed between genera. The two cyanobacterial genera were able to persist in clear-cuts, although they were more abundant on the northern side of the remnant trees. The green-algae showed no consistent pattern, Trentepohlia was affected while Trebouxia was unaffected. Our study indicates that the prerequisites for new-establishment for spore dispersed lichen species, on remnant aspen, may be fulfilled in terms of availability of free-living photobionts on the northern side of the trunks. In support of this interpretation we found that the occurrence of cyanolichens was positively correlated with the occurrence of free-living cyanobacteria in the clear-cuts. We conclude that tree retention is likely to provide a useful tool for increasing biodiversity in managed forest landscapes provided that source populations still exist in the surrounding landscape.

It is generally assumed that specialist insect herbivores utilize plant odours to find their particular host plants and that visual cues are of minor importance in the host-finding process. We performed Y-tube olfactometer bioassays and small-scale field experiments to determine whether, under laboratory and field conditions, the monophagous herbivore Altica engstroemi J. Sahlberg (Coleoptera: Chrysomelidae: Alticinae) is guided to its host plant Filipendula ulmaria (L.) Maxim. (Rosaceae) by visual or olfactory cues. The olfactometer tests showed that A. engstroemi was never attracted to odours, either from undamaged or from damaged plants. Even starvation for 24 h did not change this behaviour. However, the field experiment showed that visual cues alone were sufficient to attract a significant number of starved beetles when offered a choice between bagged host plants and bagged green plastic control 'plants'. Our findings contrast with the general view that plant odours constitute the major cue in the host-finding process among specialized phytophagous insects. A review of the literature for the period 1986-2006 inclusive, relating to host-plant finding in Chrysomelidae, identified studies of 19 chrysomelid species, all of which were guided by olfactory cues. No species were guided to their host by visual cues. Although some studies demonstrated that chrysomelids may exhibit orientation responses to colour or contrast, our study on A. engstroemi is the only one demonstrating that visual cues affect host-plant selection in a chrysomelid species. We suggest that the use of visual cues in host-finding may evolve among chrysomelids with limited dispersal ability in persistent habitats and may be found among species monophagous on abundant host plants that dominate the structure of the plant community, that is, where the host plant's presence is predictable in time and space.

Testing the effect of genetically modified hybrid aspen Populus tremula x tremuloides on the interaction with the non-target pathogen Melampsora pinitorqua.