Epidemiology of anther-smut disease (Microbotryum violaceum) and numeric regulation of populations ofSilene dioica

Summary The interaction between the anther smutMicrobotryum violaceum and its hostSilene dioica was studied during 1985–1990 in 47 populations of different ages, sizes and densities, in an archipelago area in northern Sweden. The sizes of these populations had also been surveyed in the early 1970s. Our results indicate that establishment ofMicrobotryum violaceum is host-size and density dependent. Firstly, young populations ofSilene dioica that became diseased during the study were larger and tended to be more dense than young populations that remained healthy. Secondly, populations diseased in both 1985 and 1990 were found to be larger and tended to be more dense than populations healthy in both years. We were able to document that the pathogen actually failed to establish in two small young populations (diseased plants died shortly after they appeared) and did go extinct in one small old population. Disease incidences within populations did not show large fluctuations between years. The highest increases in disease incidence during the study were found in three relatively young populations that were disease-free at the start of the study. Older populations highly diseased at the start showed less of an increase. Our study indicates thatMicrobotryum violaceum acts as a regulatory factor influencing the rate of increase inSilene dioica populations, once they are sufficiently large to maintain the pathogen. Firstly, seedling density decreased with increased incidence of disease and a seed addition experiment indicated seed-limited recuritment in highly diseased populations. Secondly, those populations that reached very large sizes or densities were either healthy or had very low incidences, indicating the potential for populations that for one reason or another escape an epidemic. However, in the comparison of changes in population size over 16–18 years there was no simple correlation between expansion rate and disease incidence.     

VARIATION IN PATHOGENICITY AMONG AND WITHIN POPULATIONS OF THE FUNGUS PHOMOPSIS SUBORDINARIA INFECTING PLANTAGO LANCEOLATA

Pathogenicity of isolates of the fungus Phomopsis subordinaria, sampled in three scarcely or heavily infected populations of Plantago lanceolata, was investigated on three different host genotypes. The expression of the pathogen appeared to be quantitative rather than qualitative in character, which suggests polygenic inheritance of host susceptibility. Significant statistical interaction between pathogen and host pointed to some degree of physiological specialization between them. None of the individual host-pathogen combinations was found to contribute significantly to the interaction. Differences in mean pathogenicity between the pathogen populations could not explain the different intensities of disease observed in the field. As the variation in susceptibility between populations of the host at the same three locations also cannot account for the differences in intensity of disease in the field, it can be concluded that environmental factors (in particular weevils that spread the disease) are important for the development of the disease. In one of the populations, the spatial scale at which variation within the pathogen occurs was determined. It appeared that the pathogen varied in pathogenicity in the field, even among scapes within an individual host plant. The consequences of this scale of variation in the pathogen are discussed for the dynamics and evolution of the pathosystem.     

AN EXPERIMENTAL FIELD STUDY OF ANTHER-SMUT DISEASE OF SILENE ALBA CAUSED BY USTILAGO VIOLACEA: GENOTYPIC VARIATION AND DISEASE INCIDENCE

Twenty cloned genotypes of Silene alba differed greatly (0–100%) in the percentage of flowering plants that became diseased by the anther-smut fungus Ustilago violacea following natural spore dispersal in a two-year field experiment. Male genotypes with the highest percentage of disease had high rates of flower production; this trait may increase the probability of spore deposition on flowers, a common site of infection. Because of this relationship, male genotypes with the highest percentage of disease also produced the most healthy flowers in the two-year period. Flowering early in the season was also a predictor of high disease levels for male genotypes in the first year. Variation among female genotypes in disease levels was not correlated with either flower production or phenology, suggesting that the sexes differ in their interaction with the pathogen. Plants of both sexes that remained nonreproductive the first year but flowered the second year could become diseased due to infection of vegetative tissue. Disease levels of the genotypes following natural spore dispersal were not correlated with disease levels of the genotypes following inoculation of vegetative tissue. This discrepancy points out that the methodology used to investigate genetic variation in disease resistance may affect the results obtained.     

INFECTIOUS DISEASES OF MAZZAELLA LAMINARIOIDES (RHODOPHYTA): CHANGES IN INFECTION PREVALENCE AND DISEASE EXPRESSION ASSOCIATED WITH SEASON,LOCALITY, AND WITHIN-SITE LOCATION1

This study addresses the issues of infection prevalence and disease expression in two wild populations of the red algal host Mazzaella laminarioides and their variability associated with locality, season, and spatial location of the host in the intertidal zone. Our results demonstrated that Endophyton ramosum is the most frequent infective pathogen affecting M. laminarioides in Matanzas and Pucatrihue. This situation prevailed through the year and across the high-to-low intertidal gradient. Although there was a general trend for lower levels of infection in late winter and early spring, only in a few, cases was well-defined seasonality detected. Furthermore, clear seasonal patterns, as displayed by deformative disease in the high intertidal zone of Pucatrihue, were attenuated in the middle and lower intertidal zones. Differences in levels of infection in M. laminarioides between the high intertidal zones of Matanzas and Pucatrihue diminished toward the low intertidal zone. Thus, effects of seasonality and locality on infection prevalence may be influenced, at least in part, by the position of the hosts an the intertidal zone. Spatial distribution of the diseased individuals also varied along the beach. This pattern was consistent between the two sites and seemed related to wave exposure and the specific pathogen. Comparisons of the size distribution of noninfected fronds with their infected counterparts showed that infections by Endophyton ramosum and Pleurocapsa sp. more frequently affected medium-and large-sized fronds. This pattern was consistent temporally and similar in the two localities. Finally, a clear association between maturity and prevalence of infection was detected. This association resulted in most fronds of the noninfected segment of the host population being immature, whereas most mature fronds were infected. In conclusion, infectious diseases affecting the red alga Mazzaella laminarioides are a persistent phenomenon in wild populations of the host, although only a small segment of the infected populations displays the full expression of the disease. In spite of the suggested role of factors such as season, latitude, and spatial location of the host on disease prevalence and expression, additional studies are needed to understand fully the dynamics of infectious diseases in wild populations of algal hosts.     

HOST-PATHOGEN INTERACTIONS IN NATURAL POPULATIONS OF LINUM MARGINALE AND MELAMPSORA LINI: II. LOCAL AND REGIONAL VARIATION IN PATTERNS OF RESISTANCE AND RACIAL STRUCTURE

Spatial variation in the resistance structure of Linum marginale (wild flax) populations to the rust fungus Melampsora lini, and in the racial structure of this pathogen, was investigated by sampling 10 populations distributed throughout the Kosciusko National Park, New South Wales, Australia. Considerable differences were found among populations in the structure of both host and pathogen. Host populations were divided into three broad categories: (1) populations susceptible to all testing races; (2) populations containing a strictly limited number of resistant phenotypes; and (3) populations with a considerable diversity of resistant phenotypes. The pathogen populations also showed a range of diversity. The major differences between these populations were determined by the occurrence and frequency of four common races of pathogen (races A, E, K, and N). These differences were apparent both at a regional spatial scale (over the 100 km separation of the most distant populations) and at a local scale where major differences were detected between two populations only 300 m apart. The distribution of the four common races of M. lini was consistent with the hypothesis that a fitness cost was associated with unnecessary virulence. In general, however, differences in the structure of pathogen populations from genetically very similar host populations implied that, in addition to host resistance genes, other evolutionary forces are also important in determining the genetic structure of individual pathogen populations.     

ANTHER SMUT DISEASE IN DIANTHUS SILVESTER (CARYOPHYLLACEAE): NATURAL SELECTION ON FLORAL TRAITS

Mating opportunities, pollination intensity, and pollen dispersal ability may vary with variation in floral traits such as color, size, and shape. Where these traits are selected by pollinators for enhanced elaboration, they should evolve toward the equilibrium between selection for further elaboration and selection against this through reduced fecundity or vitality. Here we show that pollinator-borne fungal diseases of plants may be a factor influencing the position of this equilibrium. Populations of the rock pink, Dianthus silvester often contain individuals infected with the anther smut fungus Microbotryum violaceum (= Ustilago violacea). In a naturally infected population in the Alps of eastern Switzerland we investigated how intrapopulation variation in flower size and nectar rewards influenced spore deposition and how floral traits varied with disease status. We found that spore deposition increased with increasing petal size, suggesting that large-flowered plants were at a greater risk of disease. Spore deposition was also higher for plants growing in patches with many or a high proportion of diseased neighbors. Multiple regression analyses showed that petal size or nectar reward influenced spore deposition when the effects of neighborhood disease abundance were controlled statistically. In sequential analyses, after removing the effects of disease density or frequency and plant gender, petal length explained significant variation in spore deposition. Diseased plants had reduced female reproductive organs, but calyx size was intermediate between that of healthy perfect and female flowers of this gynodioecious-gynomonoecious species, and diseased plants bore flowers with the largest petals. This may reflect a symptom of this disease or the cause, if larger-flowered plants are more likely to become infected. We conclude that investment to pollinator attraction may bring an enhanced risk of contracting this sterilizing pollinator-borne disease, so natural selection by the fungus M. violaceum acts to lower attractiveness to pollinators.     

DISEASE IMPACT AND LOCAL GENETIC DIVERSITY IN THE CLONAL PLANT PODOPHYLLUM PELTATUM

Genotypic diversity is restricted within local colonies of mayapple (Podophyllum peltatum), due to extensive asexual reproduction. Transplant experiments were used to examine whether disease impact from a specialist fungal pathogen (Puccinia podophylli) was affected by the local frequency of host genotypes within colonies. In each of six large mayapple colonies, I measured infection intensity on 1) ramets replanted in their native colony (which were thus surrounded mostly by identical genotypes) and 2) transplants from two foreign colonies (surrounded by different genotypes). Disease incidence during the pathogen's first generation did not vary significantly between native (11% infected) and foreign host genotypes (6% infected). In the pathogen's second generation, significant variation in infection intensity occurred among ramets from different source populations. However, at five of the six transplant sites, mean infection intensity was higher on some nonnative plants (locally rare host genotypes) than on natives (locally common host genotypes). In this system, pathogen attack does not act in a frequency-dependent manner to promote local genetic diversity among hosts.     

PATTERNS OF RUST INFECTION AS A FUNCTION OF HOST GENETIC DIVERSITY AND HOST DENSITY IN NATURAL POPULATIONS OF THE APOMICTIC CRUCIFER,ARABIS HOLBOELLII

It is often assumed that genetic diversity contributes to reduced disease incidence in natural plant populations. However, little is known about the genetic structure of natural populations affected by disease. Here I present data from three apomictic (asexual) populations of Arabis holboellii infected by the rusts Puccinia monoica and P. thlaspeos. An average of 300 host individuals per population were genotyped (using seven variable allozyme loci) and scored for disease presence. Arabis holboellii populations are genetically diverse; the number of clones detected per population ranged from 6 to 27. There was substantial variation in frequency of host clones within and among sites, and significant variation among clones in susceptibility to the different rusts. Contrary to predictions based on frequency-dependent selection theory there was not a consistent positive relationship between clone frequency and disease incidence within any of the populations (Spearman's r = -0.096, P > 0.5). In addition, clonally diverse populations did not necessarily have decreased disease incidence. The population with the lowest overall (both pathogens combined) disease incidence (7.5 ± 1.9%) had the smallest number of clones (6), the lowest spatial variability, and the highest Arabis density. By comparison, another population had 22 clones, high spatial variability, low Arabis density and significantly more disease overall (16.8 ± 2.7%). Although this study does not eliminate the possibility of frequency-dependent pathogen attack in these populations, the results suggest that it is likely to be weak or intermittent.     

The spatial distribution of plant populations, disease dynamics and evolution of resistance

Empirical studies of the interaction between the anther smut fungus Microbotryum violaceum and its host plant Lychnis alpina were combined with modelling approaches to investigate how variation in the spatial distribution of host populations influences disease dynamics and variation in resistance. Patterns of disease incidence and prevalence were surveyed in three contrasting systems of natural L . alpina populations where there is substantial variation in spatial structure, ranging from large continuous populations through to small isolated patches. Disease incidence (fraction of populations where disease was present) was highest in the continuous situation, and lowest in the most isolated populations. The reverse was true for prevalence (fraction of individuals diseased). To better understand the long-term ecological and evolutionary consequences of differences in among population spatial structure, we developed a two-dimensional spatially explicit simulation model in which host-population spacing was modelled by varying the percentage of sites suitable for the host. The general patterns of disease incidence and prevalence generated in the simulations corresponded well with the patterns observed in natural populations of L. alpina and M. violaceum ; i.e. the fraction of sites with disease increased while the average disease prevalence in diseased populations decreased when host populations became more connected. One likely explanation for the differences in disease incidence and prevalence seen in natural populations is that the evolution of host resistance varies as a function of the degree of fragmentation. This is supported by simulation results that were qualitatively similar to the survey data when resistance was allowed to vary, but not when hosts were assumed to be uniformly susceptible. In the former, the frequency of resistance increased markedly as host populations became more connected.     

EXPERIMENTAL EVIDENCE FOR GENETIC VARIATION IN COMPATIBILITY BETWEEN THE FUNGUS ATKINSONELLA HYPOXYLON AND ITS THREE HOST GRASSES

Variation in compatibility has been documented within and among several natural plant populations infected by fungal pathogens. In this study, seeds and isolates of the fungus Atkinsonella hypoxylon (Ascomycetes, Clavicipitaceae) were collected from three populations of the grass Danthonia spicata, two populations of D. compressa, and four populations of Stipa leucotricha. Each fungal strain was reciprocally inoculated into seedlings grown in aseptic culture from its original host population, into seedlings from other conspecific host populations, and into seedlings from the other two host species. There were three distinct patterns of compatibility, as evidenced by the ability of the fungus to grow on the seedling and to colonize new tillers. Fungal strains from one host genus were incompatible with seedlings from the other host genus. Strains from the two Danthonia species were broadly compatible among Danthonia populations and had very high rates of infection, while strains from Stipa also were broadly compatible among Stipa populations but had relatively low rates of infection. Literature surveys indicate that, in contrast to pathogenic microorganisms, mutualistic microorganisms typically exhibit broad patterns of compatibility among hosts, which lack resistance to infection. The effect of A. hypoxylon on host fitness is most detrimental in Stipa, where the fewest seedlings became infected, and most beneficial in Danthonia, where most seedlings became infected.     

ROLE OF MYCORRHIZAL FUNGUS ORIGIN IN GROWTH AND NUTRIENT UPTAKE BY GERANIUM ROBERTIANUM

Prior field studies have shown that populations of forest herbs on relatively nutrient poor soils have higher vesicular-arbuscular mycorrhizal (VAM) infection intensity than plants on rich soils. However, the growth responses and ability to take up P against the soil nutrient gradient are often not linearly related to infection intensity. To determine if intraspecific differences among populations of the common VAM fungus Glomus occultum could differentially affect growth and nutrient uptake, Geranium robertianum seedlings were inoculated with Glomus occultum isolated from four forest types along a gradient of soil fertility, and grown in a greenhouse at P levels typical of the extremes of that gradient. Plants given inoculum from relatively infertile forest sites generally produced greater root, shoot, and total mass than plants given inoculum from fertile sites or uninoculated plants, especially at the low P supply rate. Total P uptake and both P and N uptake efficiency were also highest in plants given inocula from low fertility sites. These results indicate that local adaptation and intraspecific variations in the ability of VAM fungi to induce growth and nutrient uptake effects on host plants may be as important as interspecific differences among VAM fungus species.     

Relationship of phenotypic and genetic variation in Plantago lanceolata to disease caused by Fusarium moniliforme var. subglutinans

Summary Naturally established individuals of Plantago lanceolata with the inflorescence disease caused by Fusarium moniliforme var. subglutinans had more inflorescences and were more likely to be male-sterile than healthy plants. Half-sib families planted in the field varied in the percentage of diseased plants, the number of inflorescences per plant, the incidence of male-sterility, and the pattern of inflorescence phenology. The rankings of families with respect to disease incidence was, however, not simply related to their reproductive phenotypes. Plants derived from field genotypes with a history of disease were slightly more likely to become diseased than plants derived from healthy genotypes. Inflorescence infection was more severe on plants derived from genotypes with a known history of disease. Since the fungus reduces seed production in the plants it infects, differential incidence of disease based on plant phenotype and genotype may have ecological and evolutionary consequences for the host population.     

SONG POPULATIONS AND DISPERSAL IN STEELBLUE AND PURPLE WIDOWFINCHES

Payne, R. B. 1985. Song populations and dispersal in Steelblue and Purple Widowfinches. Ostrich 56:135-146.

Songs of Steelblue Widowfinches Vidua chalybeata were recorded over eight years at Lochinvar National Park, Zambia. Neighbouring males share their song-type repertoires. Individual birds changed the structural details of all song types in their repertoire from year to year, and over 4–5 years the songs accumulated changes so that a song type could scarcely be recognized as the same. A few males in addition switched their song repertoires when they dispersed from one song population to another, or when their old neighbours disappeared and new neighbouring males sang a different song repertoire. Individual marked adult birds moved from one song neighbourhood to another. Purple Widowfinches had a similar song behaviour but only one song dialect was found in the study area.

An estimate of the proportion of immigrants was made from the proportion of birds with songs unlike their neighbours, and from birds that had a mixed song repertoire or that switched their song repertoires. Song populations were sampled at several localities in the Transvaal, Botswana, Zambia, Kenya, Tanzania, Cameroon, and Nigeria. An overall minimal estimate of dispersal between song neighbourhoods was 18 %. The incidence of observed dispersal and the movements indicated by the song differences among local widowfinches were considerable and more than sufficient to prevent genetic isoloation and differentiation of local song populations.     

The possible role of pathogenic fungi in local degeneration of Salvinia auriculata Aublet on Lake Kariba

A patchy degeneration of weed mats of Salvinia auriculata in the sheltered mouths of rivers draining into Lake Kariba was reported in 1962 to be associated with a fungal attack. Laboratory isolations from degenerating plants yielded a saprophytic Alternaria sp. and a parasitic Spicariopsis sp. Degenerating plants failed to transmit the disease when placed among healthy plants, and the results of inoculation experiments with spores of Spicariopsis sp. were inconsistent with this fungus being a primary pathogen. The characteristic feature of degeneration in the field was a mosaic of alternating ‘islands’ of diseased and healthy plants. Although the diseased plants showed the symptoms associated with Alternaria sp. and Spicariopsis sp., their characteristic distribution provided circumstantial evidence that degeneration was physiological in origin. This view was strengthened by the fact that sudd plants growing on degenerating areas of the Salvinia‘carpet’ were usually chlorotic. The presence of symptoms caused by the parasitic Spicariopsis sp. on degenerating plants could be explained by postulating that, after the plants have begun to degenerate, they are predisposed to infection by this fungus. The association of degeneration with a particular type of riverine site suggests that a seasonal investigation of the hydrobiological conditions accompanying the development of weed mats at such sites would indicate the type of factor responsible for the disease. The original hope for a biological control of Salvinia auriculata by means of a parasitic fungus remains a theoretical possibility.     

HOST-PARASITE RELATIONSHIPS IN SOOTY BLOTCH DISEASE OF WHITE CLOVER

Leaflets of Trifolium repens L. parasitized by Cymadothea trifolii (Pers.) Wolf were examined by light and electron microscopy to ascertain cellular aberrations induced by the fungus. The lamellar organization of the chloroplasts became disrupted, with a concomitant increase in the amount of osmophilic material. Many of the cells in diseased tissues exhibited tubule-like aggregations, which often were arranged in a highly ordered pattern. Curious extracytoplasmic, dumbbell-shaped structures extending between host cells and contiguous hyphae were also observed in diseased tissue.     

Preliminary Investigations about the Mode of Transmission and Spread of Xanthomonas campestris pv. graminis on Forage Grasses

Extensive studies showed that no disease was caused when seeds of different forage grasses were inoculated with Xanthomonas campestris pv. graminis. The disease could easily be induced by infecting the plants in the root system, leaves or flower. The inoculation site in the leaf proved to be of vital importance for the development of the disease. Wilting symptoms were quickly induced when the pathogen was inoculated near the leaf base. Plants in root-contact with diseased plants showed disease symptoms. It is not known whether these symptoms were caused by the bacteria or by toxins released by nearby diseased plants. Cross inoculation trials on different grass varieties revealed that different pathovars exist in the group of xanthomonads, pathogenic to forage grasses. Some have a broad host range whereas others are more limited to a single plant genus. Field trials suggest that in Belgian climatic conditions, the losses caused by bacterial wilt are rather limited.     

Anther-smut infection of Silene alba caused by Ustilago violacea: factors determining fungal reproduction

Summary The anther-smut fungus Ustilago violacea sporulates in flowers of the dioecious host plant Silene alba. Growth chamber comparisons of healthy and diseased plants, with the genetic background of host and pathogen controlled, revealed that fungal infection increases the number of flowers produced per plant and decreases the size of individual flowers. There were few consistent effects of plant genotype or fungal isolate on diseased flower traits, but differences between the plant sexes were apparent. Stimulation of flower production is proportionally greater in females than males: thus, although healthy male plants produce many more flowers than healthy females, sexual differences in diseased flower number are reduced. Sexual differences in diseased flower size also exist, with male flowers smaller than females. A field inoculation study confirmed dimorphism in diseased flower size and demonstrated that spore production per flower was greater for males than females for all flower size classes.     

RECOMBINATION AND MIGRATION RATES IN NATURAL POPULATIONS OF BACILLUS SUBTILIS AND BACILLUS MOJAVENSIS

We have investigated the rates of recombination and migration in native populations of two closely related, naturally competent Bacillus species. Native soil isolates of Bacillus subtilis and Bacillus mojavensis were obtained from three continents and, within North America, from populations at a range of geographical distances from one another. The rate of recombination within populations of each species was estimated from restriction-site data for three genes. Recombination was shown to occur within each species at about the same rate as neutral mutation, whatever the geographical scale or phylogenetic scale over which strains were sampled. The rate of migration between populations was estimated by a cladistic analysis and was shown to be high (i.e., Nm > 1), even among populations on different continents. The level of migration within each species is sufficient to prevent neutral geographical divergence within species.     

Do virus‐resistant plants pose a threat to non‐target ecosystems? I. Evidence from an Australian pathosystem based on glasshouse challenge experiments

One key environmental risk associated with the release of novel disease‐resistant plants is the potential for non‐target host populations to acquire resistance genes and undergo enemy release, leading to damage to associated native plant populations in high conservation‐value ecosystems. Unfortunately, the dynamics of most natural pathosystems are poorly understood, and risk assessment of disease‐resistant plants remains a challenge. Here we describe the first stage of a multi‐tiered risk assessment strategy aimed at quantifying potential ecological release in a model pathosystem (the weedy pasture species Trifolium repens infected with Clover yellow vein virus; ClYVV) in order to assess the level of risk posed by genetically modified and conventionally bred disease‐resistant host genotypes to non‐target plant communities in south‐eastern Australia. Glasshouse inoculation and growth experiments using 14 ClYVV isolates and 20 wild T. repens lines collected from high conservation‐value montane grassland and woodland communities show that viral infection reduces the survival and growth of host plants by on average 10–50%. However, T. repens lines exhibited variable levels of resistance and tolerance to virus infection and ClYVV isolates differed in infectivity and aggressiveness, with grassland isolates having a greater pathogenic effect on associated host plants than woodland isolates. We conclude that ClYVV potentially plays an important role in limiting the size of T. repens populations in some at‐risk non‐target ecosystems and that second‐tier field experiments are required to adequately quantify the risk associated with the commercial release of V‐R T. repens genotypes in Australia.     

The influence of plant spacing on density-dependent versus frequency-dependent spore transmission of the anther smut Microbotryum violaceum

The anther smut fungus Microbotryum violaceum is a pollinator-transmitted plant disease. As for other vector-borne diseases, frequency-dependent transmission patterns are predicted, in contrast to the density-dependent transmission of passively spread diseases. Frequency dependence will, however, only arise if vectors compensate for varying plant spacings. To test this assumption, we set up experimental populations of the host plant, Silene latifolia, with varying disease density (number of diseased plants per plot) and frequency (proportion of plants diseased), and three different plant spacings. We measured spore deposition on healthy flowers in these plots on two dates. Spore deposition decreased considerably from the first to the second census, perhaps related to the concomitant decrease in inflorescence sizes of diseased plants. At our first census, spore deposition rates varied with disease frequency, and the effect of frequency depended on plant spacing. While spore deposition was positively frequency dependent at the 1.5-m inter-plant spacing, no effect of disease frequency was found at a spacings of 0.5 m or 3 m. Nor was there an effect of disease density on spore deposition at the first census. At the later census, on the other hand, spore deposition increased almost significantly with increasing disease density (P = 0.08). This difference in deposition pattern together with a significant decrease in spore receipt indicates changes in pollinator spectrum and/or activity. The correlation of spore numbers among flowers within plants, an indication for intra-plant moves by vectors, was significant at 0.5 m and 1.5 m but not at 3 m. Floral traits and sex of individual plants influenced the number of spores they received. On the first census date, spore deposition increased with increasing inflorescence size in female but not in male plants. On the second census date, neither sex nor number of open flowers had an effect on spore receipt. None of the experimental plants became infected, however, probably because of the unusually hot and dry weather. Received: 14 May 1998 / Accepted: 19 November 1998