Diversity and distribution of co-infecting Botryosphaeriaceae from Eucalyptus grandis and Syzygium cordatum in South Africa

Species in the fungal family Botryosphaeriaceae are latent pathogens on woody trees. These fungi often have a wide host range, which can include native and introduced hosts in an area. Multi-locus DNA sequence identification on a recent collection of Botryosphaeriaceae from Eucalyptus grandis and Syzygium cordatum trees in South Africa revealed cross-infectivity of several species, novel host associations and new country reports. Neofusicoccum eucalyptorum, Neofusicoccum kwambonambiense, Neofusicoccum parvum, Neofusicoccum australe and Lasiodiplodia pseudotheobromae were identified from both tree species, with L. pseudotheobromae and N. eucalyptorum isolated for the first time from S. cordatum, similar to N. kwambonambiense from Eucalyptus. This also represents the first report of L. pseudotheobromae from South Africa. Botryosphaeriaceae species on Eucalyptus species and S. cordatum are fairly well known from South Africa. However, this study revealed new associations, indicating that conclusions should not be generalized and that more intensive sampling from different areas and over time is likely to reveal distinct species and host association patterns.     

Ability of a Generalist Seed Beetle to Colonize an Exotic Host: Effects of Host Plant Origin and Oviposition Host

The colonization of an exotic species by native herbivores is more likely to occur if that herbivore is a generalist. There is little information on the life-history mechanisms used by native generalist insects to colonize exotic hosts and how these mechanisms are affected by host properties. We examined the ability of the generalist seed beetle Stator limbatus Horn to colonize an exotic species. We compared its host preference, acceptability, performance, and egg size when ovipositing and developing on two native (Pithecellobium dulce (Roxb.) Benth and Senegalia riparia (Kunth)) and one exotic legume species (Leucaena leucocephala (Lam.)). We also analyzed the seed chemistry. We found that females recognize the exotic species as an unfavorable host for larval development and that they delayed oviposition and laid fewer and larger eggs on the exotic species than on the native species. Survivorship on the exotic host was 0%. Additionally, seeds of the native species contain five chemical compounds that are absent in the exotic species, and the exotic species contains three sterols, which are absent in the native legumes. Genetically based differences between beetles adapted to different hosts, plastic responses toward new hosts, and chemical differences among seeds are important in host colonization and recognition of the exotic host. In conclusion, the generalist nature of S. limbatus does not influence its ability to colonize L. leucocephala. Explanations for the colonization of exotic hosts by generalist native species and for the success of invasive species must be complemented with studies measuring local adaptation and plasticity.     

BIODIVERSITY RESEARCH: Parasites of exotic species in invaded areas: does lower diversity mean lower epizootic impact?

Aim Exotic species may serve as vectors for the introduction of parasites from their native range and may also become infected by parasites already present in invaded areas, but the total number of parasites infecting such exotic species in their invaded areas is typically less than that in their native range. We tested whether the diversity of parasites associated with exotic species in the native and invaded areas is related to the epizootic impact these parasites cause. Location Global. Methods We examined the diversity and epizootic impact of 384 parasite taxa associated with 22 exotic freshwater invertebrate species. The epizootic impact of each parasite was rated based on whether it had been documented to cause a major pathological impact on a large proportion of an infected host population (other than the invader under consideration). Results The total number of parasites associated with an exotic host in its native range was about twice that of all parasites associated with it in its entire invaded range. This was mainly because of the loss in the invaded areas of low impact parasites, whereas the average number of high impact parasites per host in these areas did not differ statistically from that in the native range. Main conclusions Our study suggests similar levels of adverse impact of parasites of exotic species in both their native and invaded areas. In addition to the introduction of highly pathogenic exotic parasites, other mechanisms that may be involved include (1) acquisition by the invaders of new high impact parasites in the invaded ranges, (2) high abundance of the invaders in their new ranges and (3) susceptibility of novel hosts to exotic parasites because of the ‘naive host syndrome’.     

A novel preference for an invasive plant as a mechanism for animal hybrid speciation

Homoploid hybrid speciation--speciation via hybridization without a change in chromosome number--is rarely documented and poorly understood in animals. In particular, the mechanisms by which animal homoploid hybrid species become ecologically and reproductively isolated from their parents are hypothetical and remain largely untested by experiments. For the many host-specific parasites that mate on their host, choosing the right host is the most important ecological and reproductive barrier between these species. One example of a host-specific parasite is the Lonicera fly, a population of tephritid fruit flies that evolved within the last 250 years likely by hybridization between two native Rhagoletis species following a host shift to invasive honeysuckle. We studied the host preference of the Lonicera fly and its putative parent species in laboratory experiments. The Lonicera fly prefers its new host, introduced honeysuckle, over the hosts of both parental species, demonstrating the rapid acquisition of preference for a new host as a means of behavioral isolation from the parent species. The parent taxa discriminate against each other's native hosts, but both accept honeysuckle fruit, leaving the potential for asymmetric gene flow from the parent species. Importantly, this pattern allows us to formulate hypotheses about the initial formation of the Lonicera fly. As mating partners from the two parent taxa are more likely to meet on invasive honeysuckle than on their respective native hosts, independent acceptance of honeysuckle by both parents likely preceded hybridization. We propose that invasive honeysuckle served as a catalyst for the local breakdown of reproductive isolation between the native parent species, a novel consequence of the introduction of an exotic weed. We describe behavioral mechanisms that explain the initial hybridization and subsequent reproductive isolation of the hybrid Lonicera fly. These results provide experimental support for a combination of host shift and hybridization as a model for hybrid speciation in parasitic animals.     

Overlooking the smallest matter: viruses impact biological invasions

Parasites and pathogens have recently received considerable attention for their ability to affect biological invasions, however, researchers have largely overlooked the distinct role of viruses afforded by their unique ability to rapidly mutate and adapt to new hosts. With high mutation and genomic substitution rates, RNA and single‐stranded DNA (ssDNA) viruses may be important constituents of invaded ecosystems, and could potentially behave quite differently from other pathogens. We review evidence suggesting that rapidly evolving viruses impact invasion dynamics in three key ways: (1) Rapidly evolving viruses may prevent exotic species from establishing self‐sustaining populations. (2) Viruses can cause population collapses of exotic species in the introduced range. (3) Viruses can alter the consequences of biological invasions by causing population collapses and extinctions of native species. The ubiquity and frequent host shifting of viruses make their ability to influence invasion events likely. Eludicating the viral ecology of biological invasions will lead to an improved understanding of the causes and consequences of invasions, particularly as regards establishment success and changes to community structure that cannot be explained by direct interspecific interactions among native and exotic species.     

Bursaphelenchus xylophilus: opportunities in comparative genomics and molecular host-parasite interactions

Most Bursaphelenchus species are fungal feeding nematodes that colonize dead or dying trees. However, Bursaphelenchus xylophilus, the pine wood nematode, is also a pathogen of trees and is the causal agent of pine wilt disease. B. xylophilus is native to North America and here it causes little damage to trees. Where it is introduced to new regions it causes huge damage. The most severely affected areas are found in the Far East but more recently B. xylophilus has been introduced into Portugal and the potential for damage here is also high. As incidence and severity of pine wilt disease are linked to temperature we suggest that climate change is likely to exacerbate the problems caused by B. xylophilus and, in addition, will extend (northwards in Europe) the range in which pine wilt disease can occur. Here we review what is currently known about the interactions of B. xylophilus with its hosts, including recent developments in our understanding of the molecular biology of pathogenicity in the nematode. We also examine the potential developments that could be made by more widespread use of genomics tools to understand interactions between B. xylophilus, bacterial pathogens that have been implicated in disease and host trees.     

Use of exotic hosts by Lepidoptera: widespread species colonize more novel hosts

The study of host shifts by herbivorous insects has played an important role in evolutionary biology, contributing to research in coevolution, ecological speciation, and adaptive radiation. As invasive plants become more abundant in many ecosystems, the potential for exotic host use by native insects increases. Graves and Shapiro (2003) have documented exotic host use by 34% of Californian butterflies, suggesting that the plants and butterflies of California might be an important model system for the colonization and utilization of novel resources. In this study, we analyze relationships among geographic range, native diet breadth, and the use of exotic hosts by Californian butterflies and skippers (Lepidoptera). Geographic range and, to a lesser extent, native diet breadth are significant predictors of exotic host use, with positive relationships found both before and after phylogenetic correction. These results give insight into the process of insect host range evolution, as geographically widespread generalists have an apparently greater tendency to use novel, exotic hosts than geographically constrained specialists. Increasing occurrences of exotic host use are expected and those species not capable of shifting to nonnative hosts are likely to have higher vulnerability to extirpation and extinction in the future.     

Ecological determinants of parasite acquisition by exotic fish species

Disease‐mediated threats posed by exotic species to native counterparts are not limited to introduced parasites alone, since exotic hosts frequently acquire native parasites with possible consequences for infection patterns in native hosts. Several biological and geographical factors are thought to explain both the richness of parasites in native hosts, and the invasion success of free‐living exotic species. However, the determinants of native parasite acquisition by exotic hosts remain unknown. Here, we investigated native parasite communities of exotic freshwater fish to determine which traits influence acquisition of native parasites by exotic hosts. Model selection suggested that five factors (total body length, time since introduction, phylogenetic relatedness to the native fish fauna, trophic level and native fish species richness) may be linked to native parasite acquisition by exotic fish, but 95% confidence intervals of coefficient estimates indicated these explained little of the variance in parasite richness. Based on R²‐values, weak positive relationships may exist only between the number of parasites acquired and either host size or time since introduction. Whilst our results suggest that factors influencing parasite richness in native host communities may be less important for exotic species, it seems that analyses of general ecological factors currently fail to adequately incorporate the physiological and immunological complexity of whether a given animal species will become a host for a new parasite.     

Pathogenicity and specificity of Neozygites tanajoae and Neozygites floridana (Zygomycetes: Entomophthorales) isolates pathogenic to the cassava green mite

The cassava green mite (CGM), Mononychellus tanajoa, a native of South America was accidentally introduced into Africa where it causes serious crop losses. The possibility of introducing classical biological agents from the native home of CGM into Africa was investigated. Thus, we conducted a series of laboratory assays of the native fungal pathogens, Neozygites tanajoae from Brazil and Neozygites floridana from Colombia and Brazil, and compared them with N. tanajoae isolates from Benin. Infectivity of both fungal species, was assayed against the twospotted spider mite, Tetranychus urticae, and against the red mite, Oligonychus gossypii. Pathogenicity against CGM and host range studies were conducted by transferring adult females of each mite species to leaf discs containing sporulated cadavers with a halo of conidia of each fungal isolate. All isolates caused some degree of infectivity to CGM. None of the isolates of N. floridana and N. tanajoae tested were pathogenic to O. gossypii, and only two isolates infected T. urticae. Most isolates from Brazil were highly virulent and infected only CGM. Sixteen N. tanajoae isolates caused more than 89% mortality and more than 62% of the CGM became mummified. A mummified CGM is characteristically a swollen, brown fungus-killed mite that has great potential to produce conidia. However, high mortality was not always associated with high mummification. The median mummification time ranged from 4.4 to 6.7 days. Five Brazilian isolates caused >75% mummification with a median mummification time <5 days. Isolates that cause high mummification in a short period of time would be more likely to cause epizootics and to establish in the new environment. Therefore, these isolates would be the best candidates for introduction to Africa.     

Phylogenetic congruence between subtropical trees and their associated fungi

Recent studies have detected phylogenetic signals in pathogen–host networks for both soil‐borne and leaf‐infecting fungi, suggesting that pathogenic fungi may track or coevolve with their preferred hosts. However, a phylogenetically concordant relationship between multiple hosts and multiple fungi in has rarely been investigated. Using next‐generation high‐throughput DNA sequencing techniques, we analyzed fungal taxa associated with diseased leaves, rotten seeds, and infected seedlings of subtropical trees. We compared the topologies of the phylogenetic trees of the soil and foliar fungi based on the internal transcribed spacer (ITS) region with the phylogeny of host tree species based on matK, rbcL, atpB, and 5.8S genes. We identified 37 foliar and 103 soil pathogenic fungi belonging to the Ascomycota and Basidiomycota phyla and detected significantly nonrandom host–fungus combinations, which clustered on both the fungus phylogeny and the host phylogeny. The explicit evidence of congruent phylogenies between tree hosts and their potential fungal pathogens suggests either diffuse coevolution among the plant–fungal interaction networks or that the distribution of fungal species tracked spatially associated hosts with phylogenetically conserved traits and habitat preferences. Phylogenetic conservatism in plant–fungal interactions within a local community promotes host and parasite specificity, which is integral to the important role of fungi in promoting species coexistence and maintaining biodiversity of forest communities.     

Susceptibility of lady beetles (Coleoptera: Coccinellidae) to entomopathogenic nematodes

We investigated differential susceptibility of lady beetles to entomopathogenic nematodes, for two reasons: (1) to estimate potential nontarget effects on natural lady beetle populations, (2) to compare the susceptibility of exotic versus native lady beetle species. We hypothesize that successful establishment of some exotically introduced arthropods may be due, in part, to a lower susceptibility relative to competing native species. In laboratory studies, we compared the pathogenicity, virulence, and reproductive capacity of Heterorhabditis bacteriophora and Steinernema carpocapsae among two native (Coleomegilla maculata and Olla v-nigrum) and two successfully established exotic (Harmonia axyridis and Coccinella septempunctata) lady beetles, and a known susceptible lepidopteran host, Agrotis ipsilon. After 1 and 2 days of exposure to either nematode species, mortality of A. ipsilon was higher than in all lady beetles. Thus, we predict that nematode field applications would have significantly less impact on lady beetle populations than on a susceptible target pest. Additionally, the impact of soil-applied nematodes may be lower on lady beetles than on soil-dwelling hosts because the former spends relatively less time on the soil. Exotic lady beetles were less susceptible to nematode infection than native species. Reproductive capacity data also indicated lower host suitability in H. axyridis, but not in C. septempunctata. Overall, the hypothesis that low susceptibility to pathogens in certain exotic lady beetles may have contributed to competitive establishment was supported (especially for H. axyridis). Additional studies incorporating different hosts and pathogens from various geographic locations will be required to further address the hypothesis.     

Does the periodical cicada,Magicicada septendecim,prefer to oviposit on native or exotic plant species?

Abstract 1. In 2004, Brood X of the periodical cicada (Magicicada spp.) emerged in Delaware. Extensive suburban development and concomitant planting of exotic species has occurred since the previous emergence of Brood X in 1987. 2. Exotic species could suffer extensive damage during years of cicada emergences if they are preferred for oviposition. Alternately, a shortage of suitable plant hosts may negatively affect remaining cicada populations. 3. We determined if the periodical cicada, Magicicada septendecim, preferred to oviposit on native or exotic woody plant species. Potential hosts were divided into three groups and planted in a randomised design near a likely source of cicadas. The first group, Natives, included 15 species native to Delaware. The second group, Non‐natives, included 15 exotic species that had a native congener represented in the Native group. The final group, Aliens, included 13 exotic species that did not have a native congener. 4. Based on observations of 428 plants, cicadas were more likely to oviposit on Natives or Non‐natives, which did not differ from each other in this measure, than on Aliens. Non‐natives had more oviposition holes per metre than Natives, which had more holes per metre than Aliens. The likelihood of stem flagging was lowest on Alien species; the percentage of the total plant that flagged was greatest for Natives. Plant morphology also influenced host preference and likelihood of flagging. 5. Taking plant morphology into consideration, we speculate that the evolutionary history between periodical cicadas and potential hosts is an important component of host preference.     

Potential invasion of China by exotic insect pests associated with tree seeds

A total of 39 insect species, mostly seed chalcids in the genus Megastigmus (Hymenoptera), but also midges (Diptera), are listed as potential seed-borne invaders of Chinese conifers. Although the number of native seed insects per conifer genus does not differ between China and other biogeographical regions, there are significantly fewer seed insects associated with each conifer genus in China than potential invaders. The eventual success of the invaders is likely to depend on the presence of native Chinese conifers that are congeneric with the original host, or on the presence of the original host as an exotic. When a substantial entomofauna is already associated with cones, competition for seed resources may limit the potential impact of invaders because seed insects are usually the last organisms to colonize the cone. A survey of 26 fir species, both native and introduced to Europe, showed that overall seed infestation by five species of exotic chalcids is negatively correlated to levels of damage by native insects, except on the original hosts of the chalcids. Similar patterns are hypothesized for native firs, spruces, Douglas firs, and larches in China. Uncontrolled importation of seeds and nuts of broad-leaved trees could also facilitate the introduction of seed chalcids, seed bruchids, tortricid moths and nut weevils into China. Only six species of seed chalcids are present in China, out of the 72 known to attack broad-leaved seeds over the world.     

Do Native Parasitic Plants Cause More Damage to Exotic Invasive Hosts Than Native Non-Invasive Hosts? An Implication for Biocontrol

Field studies have shown that native, parasitic plants grow vigorously on invasive plants and can cause more damage to invasive plants than native plants. However, no empirical test has been conducted and the mechanism is still unknown. We conducted a completely randomized greenhouse experiment using 3 congeneric pairs of exotic, invasive and native, non-invasive herbaceous plant species to quantify the damage caused by parasitic plants to hosts and its correlation with the hosts' growth rate and resource use efficiency. The biomass of the parasitic plants on exotic, invasive hosts was significantly higher than on congeneric native, non-invasive hosts. Parasites caused more damage to exotic, invasive hosts than to congeneric, native, non-invasive hosts. The damage caused by parasites to hosts was significantly positively correlated with the biomass of parasitic plants. The damage of parasites to hosts was significantly positively correlated with the relative growth rate and the resource use efficiency of its host plants. It may be the mechanism by which parasitic plants grow more vigorously on invasive hosts and cause more damage to exotic, invasive hosts than to native, non-invasive hosts. These results suggest a potential biological control effect of native, parasitic plants on invasive species by reducing the dominance of invasive species in the invaded community.     

Experimental assemblage of novel plant–herbivore interactions: ecological host shifts after 40 million years of isolation

Geographic isolation is the first step in insect herbivore diet specialization. Such specialization is postulated to increase insect fitness, but may simultaneously reduce insect ability to colonize novel hosts. During the Paleocene‐Eocene, plants from the order Zingiberales became isolated either in the Paleotropics or in the Neotropics. During the Cretaceous, rolled‐leaf beetles diversified in the Neotropics concurrently with Neotropical Zingiberales. Using a community of Costa Rican rolled‐leaf beetles and their Zingiberales host plants as study system, we explored if previous geographic isolation precludes insects to expand their diets to exotic hosts. We recorded interactions between rolled‐leaf beetles and native Zingiberales by combining DNA barcodes and field records for 7450 beetles feeding on 3202 host plants. To determine phylogenetic patterns of diet expansions, we established 20 experimental plots in the field, in which we planted plots five exotic Zingiberales, recording beetles feeding on these exotic hosts. In the laboratory, using both native and exotic host plants, we reared a subset of insect species that had expanded their diets to the exotic plants. The original plant–herbivore community comprised 24 beetle species feeding on 35 native hosts, representing 103 plant–herbivore interactions. After exotic host plant introduction, 20 percent of the beetle species expanded their diets to exotic Zingiberales. Insects only established on exotic hosts that belong to the same plant family as their native hosts. Laboratory experiments show that beetles are able to complete development on these novel hosts. In conclusion, rolled‐leaf beetles are preadapted to expand their diets to novel host plants even after millions of years of geographic isolation.     

Phylogenetic determinants of potential host shifts in fungal pathogens

Understanding what determines the host range of pathogens and the potential for host shifts is of critical importance to controlling their introductions into new environments. The phylogeny of the hosts has been shown to be important: pathogens are more likely to be infectious on hosts closely related to their host‐of‐origin because of the similar host environments that is shared by descent. The importance of pathogen phylogenies for predicting host range has never been investigated, although a pathogen should also be able to exploit a new host that its close relative can infect. We performed cross‐inoculations using a plant–fungal association and showed that both host and pathogen phylogenies were significant predictors of host range, with at least partly independent effects. Furthermore, we showed that some pathogens were better at infecting novel hosts. Our results should have implications in the context of biological invasions and emergences of new diseases due to globalization.     

Ectomycorrhizal fungi of exotic pine plantations in relation to native host trees in Iran: evidence of host range expansion by local symbionts to distantly related host taxa

Introduction of exotic plants change soil microbial communities which may have detrimental ecological consequences for ecosystems. In this study, we examined the community structure and species richness of ectomycorrhizal (EcM) fungi associated with exotic pine plantations in relation to adjacent native ectomycorrhizal trees in Iran to elucidate the symbiont exchange between distantly related hosts, i.e. Fagales (Fagaceae and Betulaceae) and Pinaceae. The combination of morphological and molecular identification approaches revealed that 84.6 % of species with more than one occurrence (at least once on pines) were shared with native trees and only 5.9 % were found exclusively on pine root tips. The community diversity of ectomycorrhizal fungi in the pine plantations adjacent to native EcM trees was comparable to their adjacent native trees, but the isolated plantations hosted relatively a species-poor community. Specific mycobionts of conifers were dominant in the isolated plantation while rarely found in the plantations adjacent to native EcM trees. These data demonstrate the importance of habitat isolation and dispersal limitation of EcM fungi in their potential of host range expansion. The great number of shared and possibly compatible symbiotic species between exotic Pinaceae and local Fagales (Fagaceae and Betulaceae) may reflect their evolutionary adaptations and/or ancestral compatibility with one another.     

Release from native root herbivores and biotic resistance by soil pathogens in a new habitat both affect the alien <Emphasis Type="Italic">Ammophila arenaria</Emphasis> in South Africa

Many native communities contain exotic plants that pose a major threat to indigenous vegetation and ecosystem functioning. Therefore the enemy release hypothesis (ERH) and biotic resistance hypothesis (BRH) were examined in relation to the invasiveness of the introduced dune grass Ammophila arenaria in South Africa. To compare plant–soil feedback from the native habitat in Europe and the new habitat in South Africa, plants were grown in their own soil from both Europe and South Africa, as well as in sterilised and non-sterilised soils from a number of indigenous South African foredune plant species. While the soil feedback of most plant species supports the ERH, the feedback from Sporobolus virginicus soil demonstrates that this plant species may contribute to biotic resistance against the introduced A. arenaria, through negative feedback from the soil community. Not only the local plant species diversity, but also the type of plant species present seemed to be important in determining the potential for biotic resistance. As a result, biotic resistance against invasive plant species may depend not only on plant competition, but also on the presence of plant species that are hosts of potential soil pathogens that may negatively affect the invaders. In conclusion, exotic plant species such as A. arenaria in South Africa that do not become highly invasive, may experience the ERH and BRH simultaneously, with the balance between enemy escape versus biotic resistance determining the invasiveness of a species in a new habitat.Plant nomenclature follows Arnold and De Wet (1993)     

Controls on pathogen species richness in plants' introduced and native ranges: roles of residence time, range size and host traits

Introduced species escape many pathogens and other enemies, raising three questions. How quickly do introduced hosts accumulate pathogen species? What factors control pathogen species richness? Are these factors the same in the hosts' native and introduced ranges? We analysed fungal and viral pathogen species richness on 124 plant species in both their native European range and introduced North American range. Hosts introduced 400 years ago supported six times more pathogens than those introduced 40 years ago. In hosts' native range, pathogen richness was greater on hosts occurring in more habitat types, with a history of agricultural use and adapted to greater resource supplies. In hosts' introduced range, pathogen richness was correlated with host geographic range size, agricultural use and time since introduction, but not any measured biological traits. Introduced species have accumulated pathogens at rates that are slow relative to most ecological processes, and contingent on geographic and historic circumstance.     

Factors affecting host range in a generalist seed pathogen of semi-arid shrublands

Generalist pathogens can exhibit differential success on different hosts, resulting in complex host range patterns. Several factors operate to reduce realized host range relative to potential host range, particularly under field conditions. We explored factors influencing host range of the naturally occurring generalist ascomycete grass seed pathogen Pyrenophora semeniperda. We measured potential host range in laboratory experiments at high inoculum loads with 26 grass species, including the primary host Bromus tectorum, and developed models to predict susceptibility and tolerance based on host traits, including germination speed, seed hardness, seed size, and phylogenetic relations. We also examined pathogen and host density effects on infection and mortality. All species tested were at least somewhat susceptible to the pathogen at high inoculum loads, but both infection and mortality varied widely. Species more closely related to the original host (B. tectorum) were more susceptible to infection, whereas species with slower germination were less tolerant and therefore more likely to suffer mortality. Infection and mortality were sharply reduced as inoculum load was reduced. Intermediate loads had major negative impacts on dormant B. tectorum seeds but generally minimal effects on native species. In addition, field seed bank studies determined that P. semeniperda rarely exploits native grass species as hosts. This marked reduction in realized host range relative to potential host range indicates that laboratory host range studies are potentially a poor predictor of either the current or possible future realized host range for wildland plant pathogens.