Aggregation of Colletotrichum acutatum in Response to Simulated Rain Episodes

The influence of rain splash dispersal on the aggregation and variability of spores of Colletotrichum acutatum, the causal agent of strawberry anthracnose, was studied with simulated rain over a soil surface. In the first experiment, rains with intensities of 15 and 30 mm/h were generated and sampling plates, with a selective medium for Colletotrichum, were positioned under rainshields at both 20 and 60 cm from an inoculum point source (five infected fruit) for 1-min exposure periods. Number of colonies growing in plates were used to measure spore density. For each distance, time, rain intensity, and replication, the mean (m), variance (v), and Lloyd's index of patchiness (LIP) were calculated. Estimated (LIP) exceeded 1, indicating the high variability and clustering of colonies, and hence spores, resulting from splash dispersal. LIP increased with distance from the source and with rain intensity at the greater distance. Aggregation was further quantified by Taylor's power law describing the linear relation between ln (v) and ln (m). The slope (b) of the relation for all data points was 1·67, which was significantly (P < 0·01) greater than 1, a further indication of clustering. Distance form the source had a slight positive effect on b. Colony means (Y) were transformed to Y*=Y^1/5 based on an approximate b of 1·6. Analysis of Y* indicated an expected significant main effect of rain duration and a significant interaction of distance and rain intensity. In another experiment, six infected fruit and the sampling plates were uniformly placed over the surface. LIP again exceeded 1, and Taylor's b equaled 1·63. In a final study, the inoculum source and sampling plates were raised 46 cm above the soil surface. There was a drastic reduction in Y* compared to surface values, supporting the prior hypothesis that surface topography strongly affects splash dispersal. Results indicate that the physical process of splash dispersal produces a degree of aggregation similar to that produced by population dynamic and other ecological processes.     

Patterns of unobstructed splash dispersal

Unobstructed splash dispersal patterns were measured in the absence of rain over mown grass using a fluorescent tracer, and a colorimetric method was used indoors in still air. When drops fell into a thin horizontal water film 0–1 mm deep, the volume of the incident drops dissipated as splash droplets was similar to the volume splashed from the film, irrespective of the distance of fall of the drops. Drop size, angle of inclination and distance of fall had significant effects on the volume of drops splashed from an inclined surface. The effects of rigidity, inclination and nature of surface were found to be significant when drops impacted onto surfaces with or without a wax covering and either rigidly or loosely supported. When splash- and dry-air-dispersed Lycopodium spores were simultaneously released, many more splashed spores were caught close to the source, but the dispersal gradient of splashed spores was steeper than that of dry-air-dispersed spores. Splash-dispersed spores were caught on slides, cylinders and rotorods but trap efficiency could not be evaluated.     

Studies on Mechanisms of Splash Dispersal of Spores, using Pseudocercosporella herpotrichoides Spores

Simulated raindrops, 4 or 5 mm in diameter, fell 13 m onto target water films, with Pseudocercosporella herpotrichoides spores incorporated into either drops or targets. Resulting splash droplets were collected on fixed photographic film and numbers of droplets, spore-carrying droplets and spores determined.
The patterns of dispersal of splash droplets, spore-carrying droplets and spores with distance and droplet size were similar for 4 mm and 5 mm incident drops with spores incorporated into either targets or drops. Numbers of droplets, spore-carrying droplets and spores decreased with increasing distance from targets and none were collected at 1 m. However, more spores were dispersed by 5 mm than by 4 mm drops and with spores in targets than with spores in incident drops. Whereas most splash droplets were in the smallest size category (0–100 μm), most spore-carrying droplets were 200–400 μm and most spores were in droplets with diameter greater than 1000 μm. Regressions of square root (number of spores) on droplet diameter were significant (p < 0.001) in all cases. The slopes of regression lines were greater when spores were in targets than when they were in incident drops. Splash droplets were collected up to a height of 70 cm, with most between 15 and 20 cm. The dye experiment showed that most splash droplets contained liquid from both incident drop and target film.     

Dispersal of Rhynchosporium secalis conidia from infected barley leaves or straw by simulated rain

Simulated rain (mean drop diameter c. 1 or 3 mm) was allowed to fall for 10 – 15 min on to barley leaves or straw infected by Rhynchosporium secalis (leaf blotch). The leaves were supported on a mesh through which run-off water drained and the straw was supported on a rigid surface on which run-off water collected. The numbers of R. secalis conidia and spore-carrying splash droplets collected by horizontal samplers (microscope slides and pieces of photographic film) decreased rapidly with increasing distance from and increasing height above the sources, with half-distances of 2 – 10 cm. Less than 10% of the spores or droplets reached heights of more than 30 cm. Incident drops 3 mm in diameter produced more spore-carrying droplets and dispersed more conidia than did 1 mm drops. The size category of splash droplets with the greatest proportion of the spore-carrying droplets dispersed by 3 mm drops was 200 – 400 μm, whether the source was infected barley leaves or barley straw. For leaves or straw the greatest proportions of spores were carried in droplets > 1000 μm in diameter. The mean diameter of spore-carrying droplets (478 μm) dispersed from free-draining leaves was less than that of droplets from straw plus run-off water (563 μm). However, the leaf source had more spores cm^-2 and the mean number of spores per droplet was greater (113 as opposed to 6·8) than for the straw source.     

Dispersal of Septoria nodorum Pycnidiospores by Simulated Rain and Wind

The influence of wind on the splash dispersal of Septoria nodorum pycnidiospores was studied in a raintower/wind tunnel complex with single drops or simulated rain falling on spore suspensions or infected stubble with windspeeds of 1.5 to 4 m/sec. When single drops fell on spore suspensions (depth 0.5 mm, concentration 7.8 × 10⁵ spores/ml) most of the spore-carrying droplets collected on fixed photographic film between 0–4 m downwind (windspeed 3 m/sec) were >200 μm in diameter. However, most spores were carried in droplets with diameter > 1000 μm, 70 % of which carried more than 100 spores. When simulated rain fell on infected stubble most of the spore-carrying droplets collected beyond 1 m downwind (windspeeds 1.4 and 4 m/sec) were <200 μm in diameter and none were >600 μm; most of these droplets carried only one spore. The distribution of splash droplets (with diameter >100 μm) deposited on chromatography paper showed a maximum at 40–50 cm upwind of the target but many more droplets were deposited 20–30 cm downwind, when single drops fell on a spore suspension (concentration 1.2 × 10⁵ spores/ ml) containing fluorescein dye with a windspeed of 2 m/sec; droplets were collected up to 3 m downwind but not more than 70 cm upwind. With a windspeed of 3 m/sec, numbers of sporecarrying droplets and spores collected on film decreased with increasing distance downwind; most were collected within 2 m of the target but some were found up to 4 m. When simulated rain fell on infected stubble, increasing the windspeed from 1.5 to 4 m/sec greatly increased the number of spores deposited more than 1 m downwind. At 1.5 m/sec none were collected beyond 2 m downwind, whereas at 4 m/sec some were collected at 4 m. A few air-borne S. nodorum spores were collected by suction samplers at a height of 40 cm at distances up to 10 m downwind of a target spore suspension on which simulated rain fell.     

Secondary seed dispersal of Erodiophyllum elderi,a patchily distributed short‐lived perennial in the arid lands of Australia

We investigated secondary dispersal of propagules of Erodiophyllum elderi (Asteraceae), a short‐lived perennial plant growing in small patches in the arid lands of southern Australia. In spite of its importance for population dynamics, secondary dispersal is a little understood process. We monitored the dispersal of 2280 large woody capitula (seed heads) released in six E. elderi patches for 9 months. Colour‐coded seed heads were located at night using UV light and their distance and direction from the release point were measured. Over the 9‐month period, more seed heads moved, and those that did, moved further in areas with high herbivore activity. Overall dispersal distance across the ground was limited to less than 30 m. Dispersal patterns were related to the topographical slope at the release site: seed heads moved further, and more dispersed on steeper slopes unless the steep slopes had sandy soil in which case seed heads were buried, caught or there was reduced sheet water flow limiting their dispersal potential. After several months, seed head dispersal virtually ceased as seed heads became stuck in the debris and soil after heavy rains or further dispersal became unlikely when seed heads reached locally low‐lying areas. Secondary dispersal patterns suggest two distinctly different influences associated with the presence of herbivores: the direct movement of seed heads by trampling from sheep (an introduced herbivore) and the indirect effect of a reduced standing biomass from grazing. Reduced vegetation cover allows seed head redistribution via sheet water flow during large rainfall events.     

Dispersal may limit the occurrence of specialist wood decay fungi already at small spatial scales

We used species‐specific spore traps to measure airborne dispersal of the wood decay fungus Phlebia centrifuga (spore size 6.5–9 × 2.5–3 μm) up to 1000 m distance from a point source. We fitted two simple dispersal models, an empirical power law model and a semi‐mechanistic diffusion model to the data using the Bayesian approach. The diffusion model provided a better fit than the power law model which underestimated deposition at 3–55 m and overestimated deposition at longer and shorter distances. Model fit improved by allowing overdispersion, suggesting that spores are not dispersed independently but wind can transport spores in groups inside discrete air packages up to considerable distances. Using the fitted diffusion model and available information on the establishment rates of wood‐decay fungi, we examine the distance up to which colonisation from a single fruit body is likely to occur. We conclude that the diluting effect of distance and low establishment success make the occurrence of P. centrifuga dispersal limited possibly already at the distance of tens of metres and very probably at a few hundred metres from the nearest fruit body, despite the fact that under favourable conditions a high proportion of the spores can disperse considerably further. This conclusion is likely to hold generally for those fungal species that inhabit fragmented landscapes, have specialised resource and habitat requirements, and have similar spore size and other dispersal traits as P. centrifuga.     

Splash seed dispersal by raindrops

Splash seed dispersal by raindrops was investigated for plants in southern Japan. Nine families, 10 genera and 19 species were confirmed as raindrop-dispersed plants. The 10 genera were Gentiana, Gratiola, Chrysosplenium, Mazus, Mitella, Ophiorrhiza, Sagina, Sedum, Trigonotis and Veronica. The method of splash rain dispersal in these species was clarified. Raindrop-dispersed species were all small herbaceous plants with a vertical pedicel and an apically opening fresh capsule when the seeds mature. Open capsules were cup-shaped or boat-shaped and can accommodate raindrops easily. The raindrops splashed the seeds from the capsule. In general, the seeds weighed very little, but they were heavier than powder or dust seeds dispersed by wind. A strong negative correlation was found between seed weight and the number of seeds per capsule. In the case of Trigonotis brevipes (Maxim.) Maxim., raindrops were received into the cup-shaped calyx-tube and dispersed the fruitlets. Some species, such as Gentiana thunbergii (G. Don) Griseb., Gentiana zollingeri Fawcett and Ophiorrhiza japonica Blume, had hydroscopic movement capsules that opened widely only when wet. Raindrop-dispersed plants were found in various habitats. For example, some plants grew together on rocks along the mountain torrents where splash water could easily be caught. The results of the laboratory and field experiments indicated that the dispersal distance of seeds by raindrops was 1m or less. For small herbaceous plants, splash dispersal by rain might be an effective and advantageous method of seed dispersal because dispersal is not affected by plant height.     

Detection of arbuscular mycorrhizal fungal spores in the air across different biomes and ecoregions

Aerial dispersal of fungal spores is common, but the role of wind and air movement in dispersal of spores of arbuscular mycorrhizal (AM) fungi is largely unknown. Several studies have examined the possibility of AM fungal spores being moved by wind vectors without observing spores taken from the air environment. For the first time this study observed the presence of AM fungal spores in the air. The frequency of AM fungal spores in the air was determined in six North American biomes composed of 18 ecoregions. Multiple samples were taken from both the air and the soil at each location. AM fungal spores were found in high abundance in the soil (hundreds of spores per gram of soil), however, they were rarely found in the air (most samples contained no AM fungal spores). Furthermore, only the Glomus morphotype was found in the air, whereas spores in the soil were taxomomically more diverse (Glomus, Acaulospora, Gigaspora, Scutellospora morphotypes were observed). The proportion of Glomus spores in the air relative to Glomus spores in the soil was highest in more arid systems, indicating that AM fungi may be more likely to be dispersed in the air in such systems. Nonetheless, the results indicate that the air is not likely a dominant mode of dispersal for AM fungi.     

Spore rain in relation to regional sources and beyond

While patterns of spore dispersal from single sources at short distances are fairly well known, information about ‘spore rain’ from numerous sources and at larger spatial scales is generally lacking. In this study, I sampled spore rain using a novel method consisting of 0.25–0.5 m² cotton cloth traps at nine sites in the boreo‐nemoral vegetation zone in eastern Sweden during two seasons, using Sphagnum spores as a model. Traps were located in various landscapes (mainland, islands). Additional trapping was done in an arctic area (Svalbard) without spore production. Spore densities were tested against distance from the nearest source and area of sources (open peatlands) within different radii around each site (5, 10, 20, 50, 100, 200, 300, 400 km). The cloth method appeared reliable when accounting for precipitation losses, retaining approximately 20–60% of the spores under the recorded amounts of precipitation. Estimated spore densities ranged from 6 million m^?2 and season within a large area source, via regional deposition of 50 000–240 000 spores m^?2, down to 1000 m^?2 at Svalbard. Spore rain for all sites was strongly related to distance from the nearest source, but when excluding samples taken within a source peatland, the amount of sources within 200 km was most important. Spores were larger at isolated island sites, indicating that a higher proportion originated from distant, humid areas. Immense amounts of Sphagnum spores are dispersed across regional distances annually in boreal areas, explaining the success of the genus to colonise nutrient poor wetlands. The detectable deposition at Svalbard indicates that about 1% of the regional spore rain has a trans‐ or intercontinental origin. The regional spore rain, originating from numerous sources in the landscape, is probably valid for most organisms with small diaspores and provides a useful insight in ecology, habitat restoration and conservation planning.     

Long-Distance Wind-Dispersal of Spores in a Fungal Plant Pathogen: Estimation of Anisotropic Dispersal Kernels from an Extensive Field Experiment

Given its biological significance, determining the dispersal kernel (i.e., the distribution of dispersal distances) of spore-producing pathogens is essential. Here, we report two field experiments designed to measure disease gradients caused by sexually- and asexually-produced spores of the wind-dispersed banana plant fungus Mycosphaerella fijiensis. Gradients were measured during a single generation and over 272 traps installed up to 1000 m along eight directions radiating from a traceable source of inoculum composed of fungicide-resistant strains. We adjusted several kernels differing in the shape of their tail and tested for two types of anisotropy. Contrasting dispersal kernels were observed between the two types of spores. For sexual spores (ascospores), we characterized both a steep gradient in the first few metres in all directions and rare long-distance dispersal (LDD) events up to 1000 m from the source in two directions. A heavy-tailed kernel best fitted the disease gradient. Although ascospores distributed evenly in all directions, average dispersal distance was greater in two different directions without obvious correlation with wind patterns. For asexual spores (conidia), few dispersal events occurred outside of the source plot. A gradient up to 12.5 m from the source was observed in one direction only. Accordingly, a thin-tailed kernel best fitted the disease gradient, and anisotropy in both density and distance was correlated with averaged daily wind gust. We discuss the validity of our results as well as their implications in terms of disease diffusion and management strategy.     

Splash dispersal of fungus spores and fungicides in the laboratory and greenhouse

A laboratory technique is described for the production of drops of simulated rain in which fungal spores were suspended. When such drops containing conidia of Botrytis fabae impacted on a target leaf the secondary droplets produced infections on receptor broad bean leaves. The capacity of fungicides applied to the target leaf to redistribute in secondary splash droplets was examined in terms of the infectivity of the spores in the droplets. The extent to which a copper fungicide reduced infection on the receptor leaves was related to the level and tenacity of the fungicide deposit on the target leaf. The effect of wetting agents on the redistribution of this fungicide could probably be explained by their influence on the tenacity of the initial deposit. In general the capacity of different fungicides to inhibit infection by the secondary droplets was related to the inherent toxicity of the fungicides to B. fabae. Implications of the dispersal of spores and fungicides by rain splash are briefly considered with reference to field conditions.     

Genet dynamics and ecological functions of the pioneer ectomycorrhizal fungi Laccaria amethystina and Laccaria laccata in a volcanic desert on Mount Fuji

To understand the reproduction of the pioneer ectomycorrhizal fungi Laccaria amethystina and Laccaria laccata in a volcanic desert on Mount Fuji, Japan, the in situ genet dynamics of sporocarps were analysed. Sporocarps of the two Laccaria species were sampled at fine and large scales for 3 and 2 consecutive years, respectively, and were genotyped using microsatellite markers. In the fine-scale analysis, we found many small genets, the majority of which appeared and disappeared annually. The high densities and annual renewal of Laccaria genets indicate frequent turnover by sexual reproduction via spores. In the large-scale analysis, we found positive spatial autocorrelations in the shortest distance class. An allele-clustering analysis also showed that several alleles were distributed in only a small, localised region. These results indicate that Laccaria spores contributing to sexual reproduction may be dispersed only short distances from sporocarps that would have themselves been established via rare, long-distance spore dispersal. This combination of rare, long-distance and frequent, short-distance Laccaria spore dispersal is reflected in the establishment pattern of seeds of their host, Salix reinii.     

Cheek-pouch dispersal of seeds by Japanese monkeys (Macaca fuscata yakui) on Yakushima Island, Japan

Seed dispersal by Japanese monkeys (Macaca fuscata yakui) via cheek-pouch was studied in a warm temperate evergreen forest on Yakushima Island. Plant list was compiled based on a study during 1986–1995, of which troops of monkeys have been habituated without artificial feeding. We followed the well-habituated monkeys in 1993 and 1994 to observe the feeding behavior and their treatments of fruits and seeds, and collected seeds dispersed by monkeys to record the distance carried from the mother trees. We checked the difference of germination ratio between seeds dispersed via cheek-pouch and seeds taken from mother trees by sowing experiments. Seeds and acorns of 22 species were observed to be dispersed via cheek-pouch of monkeys. Among them, three species with acorns were never dispersed via feces, and 15 species with drupes were seldom dispersed via feces. Plant species of which seeds are dispersed only via cheek-pouch had larger seeds than those of dispersed both via cheek-pouch and via feces, and typically had only one or two seeds in a fruit. As for one of cheek-pouch dispersal species,Persea thunbergii, the mean distance when seeds were carried from the mother trees via cheek-pouch was 19.7 m, and the maximum distance was as long as 105 m although more than 80% of seeds were dispersed within 30 m from mother trees. And 82% of seeds dispersed via cheek-pouch germinated. The easy separation of seeds from other parts of the fruit seems to facilitate cheek-pouch dispersal more than dispersal via feces. Cheek-pouch dispersal by monkeys has possibly enhanced the natural selection for larger seeds which bring forth larger seedlings with high shade-tolerance. In conclusion, cheek-pouch dispersal by monkeys is quite an important mode for trees in the mature stand in a warm temperate evergreen forest on Yakushima Island.     

Seed dispersal distance,seed morphology,and recruitment in the Chilean sclerophyllous tree Quillaja saponaria: implications for passive restoration in a semiarid ecosystem

Recolonization of wind-dispersed tree species in degraded areas may decline with distance from remnant forest fragments because seed rain frequently decreases with distance from the seed source. However, regeneration of these species may be even more limited to sites close to the seed source if dispersal distance is negatively affected by seed mass, and germination probability is positively affected by seed mass. We evaluated these hypotheses in a Mediterranean-type ecosystem of central Chile, using the wind-dispersed tree species Quillaja saponaria. We assessed the seed rain curve in a degraded open area adjacent to a remnant forest fragment of this species, and related seed mass with dispersal distance from the seed source. Then, we evaluated the relationship between seed mass, germination, and seedling growth, and if seeds that fall nearer the seed source have greater germination probability. We found a decreasing seed rain with the distance from the seed source. Seed mass was not related to dispersal distance, although seeds with higher wing area dispersed further. Germination probability was significantly and positively related to the seed mass. We observed no significant relationship between distance and germination probability. We conclude that germination probability of this species does not vary along the seed rain curve, and that the recruitment density would be greater near the seed source only due to decreasing seed rain with distance. Our results suggest that this species has the potential to be passively restored in degraded areas, especially within the first 70 m from the remnant forest fragments.

     

Distribution and decline of human pathogenic bacteria in soil after application in irrigation water and the potential for soil-splash-mediated dispersal onto fresh produce

Aims: To improve our understanding of the survival and splash‐mediated transfer of zoonotic agents and faecal indicator bacteria introduced into soils used for crop cultivation via contaminated irrigation waters. Methods and Results: Zoonotic agents and an Escherichia coli marker bacterium were inoculated into borehole water, which was applied to two different soil types in early‐, mid‐ and late summer. Decline of the zoonotic agents was influenced by soil type. Marker bacteria applied to columns of two soil types in irrigation water did not concentrate at the surface of the soils. Decline of zoonotic agents at the surface was influenced by soil type and environmental conditions. Typically, declines were rapid and bacteria were not detectable after 5 weeks. Selective agar strips were used to determine that the impact of water drops 24–87 μl could splash marker bacteria from soil surfaces horizontal distances of at least 25 cm and heights of 20 cm. Conclusions: Soil splash created by rain‐sized water droplets can transfer enteric bacteria from soil to ready‐to‐eat crops. Persistence of zoonotic agents was reduced at the hottest part of the growing season when irrigation is most likely. Significance and Impact of the Study: Soil splash can cause crop contamination. We report the penetration depths and seasonally influenced declines of bacteria applied in irrigation water into two soil types.     

Dispersal of spores and pollen from crops

Fungal spores and pollens can be dispersed in a number of ways: by animals and insects; by water; by wind or by rain. This paper concentrates on the effects of wind on the dispersal of spores and pollen grains and the effects of rain on spore dispersal. For dispersal to be successful particles must complete three phases: removal, dispersal through the air and deposition. The biology of the organism and its environment can affect all three phases, however, once released the fate of all airborne particles largely depends on the laws of physics which govern the motion of the air. Many types of spore are actively ejected into the air while others are simply blown from the host surface. Particle size and shape affects dispersal and deposition phases. Local environmental factors such as temperature, humidity and light, as well as wind or rain, can play a key role in the removal of spores. Wind speed and turbulence or rainfall, largely determine spore dispersal, but, the size and shape of the particle, the nature of the plant canopy and the way the particles are released into the air may also be important. Particle deposition depends on both environmental and biological factors. This paper briefly considers these processes using examples and how they can be modelled.     

Environmental filtering affects soil fungal community composition more than dispersal limitation at regional scales

The relative importance of dispersal limitation versus environmental filtering for community assembly has received much attention for macroorganisms. These processes have only recently been examined in microbial communities. Instead, microbial dispersal has mostly been measured as community composition change over space (i.e., distance decay). Here we directly examined fungal composition in airborne wind currents and soil fungal communities across a 40 000 km² regional landscape to determine if dispersal limitation or abiotic factors were structuring soil fungal communities. Over this landscape, neither airborne nor soil fungal communities exhibited compositional differences due to geographic distance. Airborne fungal communities shifted temporally while soil fungal communities were correlated with abiotic parameters. These patterns suggest that environmental filtering may have the largest influence on fungal regional community assembly in soils, especially for aerially dispersed fungal taxa. Furthermore, we found evidence that dispersal of fungal spores differs between fungal taxa and can be both a stochastic and deterministic process. The spatial range of soil fungal taxa was correlated with their average regional abundance across all sites, which may imply stochastic dispersal mechanisms. Nevertheless, spore volume was also negatively correlated with spatial range for some species. Smaller volume spores may be adapted to long-range dispersal, or establishment, suggesting that deterministic fungal traits may also influence fungal distributions. Fungal life-history traits may influence their distributions as well. Hypogeous fungal taxa exhibited high local abundance, but small spatial ranges, while epigeous fungal taxa had lower local abundance, but larger spatial ranges. This study is the first, to our knowledge, to directly sample air dispersal and soil fungal communities simultaneously across a regional landscape. We provide some of the first evidence that soil fungal communities are mostly assembled through environmental filtering and experience little dispersal limitation.     

Spore dispersal of a resupinate ectomycorrhizal fungus, Tomentella sublilacina, via soil food webs

Patterns of fungal spore dispersal affect gene flow, population structure and fungal community structure. Many Basidiomycota produce resupinate (crust-like) basidiocarps buried in the soil. Although spores are actively discharged, they often do not appear to be well positioned for aerial dispersal. We investigated the potential spore dispersal mechanisms of one exemplar of this growth form, Tomentella sublilacina. It is a widespread ectomycorrhizal fungus that sporulates in the soil organic horizon, can establish from the spore bank shortly after disturbance, but also can be a dominant species in mature forest stands. We investigated whether its spores could be dispersed via spore-based food webs. We examined external surfaces, gut contents and feces from arthropod fungivores (mites, springtails, millipedes, beetles, fly larvae) and arthropod and vertebrate predators (centipedes, salamanders) from on and around T. sublilacina sporocarps. Spore densities were high in the guts of many individuals from all fungivore groups. Centipede gut contents, centipede feces and salamander feces contained undigested invertebrate exoskeletons and many apparently intact spores. DAPI staining of spores from feces of fungivores indicated that 7-73% of spores contained intact nuclei, whereas spores from predators had lower percentages of intact nuclei. The spiny spores often were lodged on invertebrate exoskeletons. To test the viability of spores that had passed through invertebrate guts we used fecal droppings of the millipede Harpaphe haydeniana to successfully inoculate seedlings of Pinus muricata (Bishop pine). These results indicate the potential for T. sublilacina spore dispersal via invertebrates and their predators in soil food webs and might help to explain the widespread distribution of this species. It is likely that this is a general mechanism of dispersal for fungi producing resupinate sporocarps, indicating a need to develop a fuller understanding of the linkages of soil food webs and spore dispersal.     

Frugivory and Seed Dispersal by Northern Pigtailed Macaques (Macaca leonina), in Thailand

Tropical rain forest conservation requires a good understanding of plant–animal interactions. Seed dispersal provides a means for plant seeds to escape competition and density-dependent seed predators and pathogens and to colonize new habitats. This makes the role and effectiveness of frugivorous species in the seed dispersal process an important topic. Northern pigtailed macaques (Macaca leonina) may be effective seed dispersers because they have a diverse diet and process seeds in several ways (swallowing, spitting out, or dropping them). To investigate the seed dispersal effectiveness of a habituated group of pigtailed macaques in Khao Yai National Park, Thailand, we examined seed dispersal quantity (number of fruit species eaten, proportion in the diet, number of feces containing seeds, and number of seeds processed) and quality (processing methods used, seed viability and germination success, habitat type and distance from parent tree for the deposited seeds, and dispersal patterns) via focal and scan sampling, seed collection, and germination tests. We found thousands of seeds per feces, including seeds up to 58 mm in length and from 88 fruit species. Importantly, the macaques dispersed seeds from primary to secondary forests, via swallowing, spitting, and dropping. Of 21 species, the effect of swallowing and spitting was positive for two species (i.e., processed seeds had a higher % germination and % viability than control seeds), neutral for 13 species (no difference in % germination or viability), and negative (processed seeds had lower % germination and viability) for five species. For the final species, the effect was neutral for spat-out seeds but negative for swallowed seeds. We conclude that macaques are effective seed dispersers in both quantitative and qualitative terms and that they are of potential importance for tropical rain forest regeneration.