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.     

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.     

Direct Splash Dispersal Prevails over Indirect and Subsequent Spread during Rains in Colletotrichum gloeosporioides Infecting Yams

Plant pathogens have evolved many dispersal mechanisms, using biotic or abiotic vectors or a combination of the two. Rain splash dispersal is known from a variety of fungi, and can be an efficient driver of crop epidemics, with infectious strains propagating rapidly among often genetically homogenous neighboring plants. Splashing is nevertheless a local dispersal process and spores taking the droplet ride seldom move farther than a few decimeters. In this study, we assessed rain splash dispersal of conidia of the yam anthracnose agent, Colletotrichum gloeosporioides, in an experimental setting using a rain simulator, with emphasis on the impact of soil contamination (i.e., effect of re-splashing events). Spores dispersed up to 50 cm from yam leaf inoculum sources, though with an exponential decrease with increasing distance. While few spores were dispersed via re-splash from spore-contaminated soil, the proportion deposited via this mechanism increased with increasing distance from the initial source. We found no soil contamination carryover from previous rains, suggesting that contamination via re-splashing from contaminated soils mainly occurred within single rains. We conclude that most dispersal occurs from direct splashing, with a weaker contribution of indirect dispersal via re-splash.     

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.     

Does proximity to a mature forest contribute to the seed rain and recovery of an abandoned agriculture area in a semiarid climate?

Proximity to forests contributes to the recolonisation of anthropogenic‐disturbed areas through seed input. We evaluated the role of proximity to a mature forest in the recolonisation of an agricultural area that has been abandoned for 18 years and is currently a young forest. Seed rain was monitored at fixed distances from the mature forest. The type of surface recolonisation (germination versus resprouting) and the reproductive season were measured in both forests. The majority of plants recolonising the young forest originated from seed germination. Proximity to the mature forest contributed to the seed rain in the young forest; however, 18 years has not provided sufficient time for the recolonisation of 80 species present in the mature forest. Some species shared between forests differed in their fruiting season and seed dispersal. The seed rain had a total species richness of 56, a total density of 2270 seeds·m^?2·year^?1 and predominance of self‐ and wind dispersal. A significant reduction in seed rain with increasing distance from the mature forest was observed. The young forest contained 35 species not observed in the mature forest, and the floristic similarity between the two forests was 0.5, indicating that the two forests are floristically distinct.     

Vertebrate‐mediated seed rain and artificial perches contribute to overcome seed dispersal limitation in a Mediterranean old field

Natural regeneration of vegetation is a frequent outcome of land abandonment, although the rate and diversity of such regeneration may be severely restricted by seed dispersal limitation, among other factors. In spite of this, studies aiming to quantify seed rain and test methods to enhance it, such as artificial perches, are still underrepresented in the Mediterranean. In our study, we quantified seed rain density and richness and tested the effects of artificial perches on such rain over a distance gradient on seven Mediterranean island old fields. In each of the seven sites, we positioned three sampling stations, each consisting of 1 seed trap under an artificial perch and 1 as a control on the ground, distributed at 30, 60, and 90 m from natural vegetation remnant. All traps received seeds, suggesting no overall dispersal limitation. Of the 11 seed species found, 10 were fleshy‐fruited and dispersed by vertebrates. Seed traps under perches received significantly higher seed rain of fleshy‐fruited species dispersed by birds, while ground traps received significantly more seeds of the species also dispersed by mammals, especially Rubus ulmifolius. The distance from the seed source was nonsignificant in all cases. Our study demonstrates the key role of vertebrate‐mediated seed dispersal services to overcome dispersal limitation in old fields, as well as the effective contribution of even small artificial perches in contrasting such limitation. The lack of differences over the distance gradient reveal that the upper spatial limit of dispersal limitation was not achieved.     

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.

     

Spatiotemporal patterns of seed dispersal in a wind-dispersed Mediterranean tree (Acer opalus subsp. granatense): implications for regeneration

Seed dispersal can severely limit the quantity of plant recruits and their spatial distribution. However, our understanding of the role of dispersal in regeneration dynamics is limited by the lack of knowledge of seed deposition patterns in space and time. In this paper, we analyse the spatiotemporal variability of seed dispersal patterns in the Mediterranean maple, Acer opalus subsp. granatense, by monitoring seed rain along two years at a broad spatial scale (2 mountain ranges, 2 populations per range, 4 microhabitats per population). We quantified seed limitation and its components (source and dispersal limitation), and explored dispersal limitation in space by analysing dispersal distances, seed aggregation, and microhabitat seed distribution. Acer opalus subsp. granatense was strongly seed‐limited throughout the gradients explored, being always dispersal limitation much higher than source limitation. The distribution of seeds with distance from adult individuals was leptokurtic and right‐skewed in all populations, being both kurtosis and skewness higher the year of the highest seed production. Dispersal distances were shorter than expected by random in the four populations, which suggests distance‐limited dispersal. Dispersal patterns were highly aggregated and showed a preferential direction around adults. At the microhabitat scale, most seeds accumulated under adult maples. However, there were no more seeds under trees and shrubs other than maple than in open interspaces, implying that established vegetation does not disrupt patterns of seed deposition by physically trapping seeds. When compared with patterns of seedling establishment, limited dispersal ability and inter‐annual spatial concordance in seed rain patterns suggest that several potentially safe sites for recruitment have a very low probability of receiving seeds in most maple populations. These findings are especially relevant for rare species such as Acer opalus subsp. granatense, and illustrate how dispersal studies are not only crucial for our understanding of plant population dynamics but also to provide conservation directions.     

CULTIVATION OF ARTHROSPIRA (SPIRULINA) PLATENSIS (CYANOPHYCEAE) BY FED‐BATCH ADDITION OF AMMONIUM CHLORIDE AT EXPONENTIALLY INCREASING FEEDING RATES1

Arthrospira (Spirulina) platensis (Nordstedt) Gomont was cultivated under light‐limited conditions in 5‐L open tanks by daily supplying NH₄Cl as nitrogen source. Exponentially increasing feeding rates were adopted to prevent ammonia toxicity. The total feeding time (T) was varied between 12 and 20 days, and the starting (m₀) and total (m_T) quantities of the nitrogen source per unit reactor volume were varied in the ranges 0.19–1.7 mM and 2.3–23.1 mM, respectively. This intermittent addition of the nitrogen source prevented ammonia from reaching inhibitory levels and ensured final cell concentrations (X_m) and cell productivities (P_x) comparable with those of batch runs with KNO₃. Moreover, the lower nitrogen addition due to the use of NH₄Cl rather than KNO₃ allowed for higher nitrogen‐to‐cell conversions (Y_x/n). These results were evaluated using three‐factor, five‐level, central composite experimental planning, combined with the response surface methodology, selecting T, m₀, and m_T as the independent variables and X_m, P_x, and Y_x/n as the response variables. This approach allowed us to identify, through the simultaneous optimization of the variables, T=16 days, m₀=1.7 mM, and m_T=21.5 mM as the best conditions for A. platensis cultivation at 72 μmol photons·m^?2·s^?1. Under these conditions, a maximum cell concentration of 1239 mg ·L^?1 was obtained, which is a value comparable with that obtained using KNO₃ as nitrogen source and nearly coincident with the theoretical one estimated by the response surface methodology.     

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.     

Dispersal of the conidia of Colletotrichum gloeosporioides by rain and the development of anthracnose on onion

The conidia of Colletotrichum gloeosporioides were found to be dispersed during rainfall by wash-off and splash mechanisms. The initiation and development of onion anthracnose was found to depend on the frequency of rainfall and the movement of conidial inoculum during rainfall. Experiments conducted under controlled conditions in the laboratory employing splash and wash-off assemblies showed that impacting incident water drops (splash) and flowing water (wash-off) liberated the conidia from the anthracnose lesions of the onion leaf/peduncle. Peak liberation of conidia occurred with 3 to 5 water drops and most of the conidia were removed from the source within 90 seconds. A possibility of the dispersal of conidia of C. gloeosporioides from soil to lower leaf by splash mechanisms and then from the leaves to the neck of the onion bulb and to the bulb by wash-off mechanisms is indicated.     

Xylitol conversion by fermentation using five yeast strains and polyelectrolyte-assisted ultrafiltration

Batch fermentations for xylitol production were conducted using Candida boidinii (BCRC 21432), C. guilliermondii (BCRC 21549), C. tropicalis (BCRC 20520), C. utilis (BCRC 20334), and P. anomala (BCRC 21359) together with a mixture of sugars simulating lignocellulosic hydrolysates as the carbon source. C. tropicalis had the highest bioconversion yield (Y_P/S) of 0.79 g g⁻¹ (g xylitol·g xylose⁻¹) over 48 h. Additional fermentations with C. tropicalis achieved Y_P/S values of 0.6 and 0.39 g g⁻¹ after 96 and 72 h using urea and soybean meal as the nitrogen sources, respectively. Ethanol and arabitol were also produced in all fermentation. Xylitol in the fermentation broth was recovered by cross-flow ultrafiltration. With prior application of 2 mg polydiallyl dimethylammonium chloride l⁻¹ on the membrane surface, protein in the permeate was reduced from 7.1 to 1.5 mg l⁻¹ after 2 h.     

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.     

Evaluating Dispersal Limitation in Passive Bottomland Forest Restoration

Dispersal limitation can retard natural establishment of desirable species on restoration sites, especially where landscapes are fragmented, but dispersal limitation is assumed to become less critical with time as early colonists become reproductively mature. Distribution patterns of recruiting trees in a 20‐year‐old passively restored bottomland in northeast Louisiana suggested persistent dispersal limitation in some bottomland hardwood species and influence of dense shrub patches on colonization. To test these hypotheses, we measured seed rain as a function of distance to seed source and association with shrub cover. Seed rain of the wind‐dispersed Fraxinus pennsylvanica was highest near the forest edge, except where mature recruits occurred. Although shrub presence did not influence dispersal of F. pennsylvanica, its negative influence on probability of occurrence in the sapling layer suggests that shrub cover may limit its regeneration. The bird‐dispersed Crataegus viridis and Ilex decidua were found in the seed rain and as reproductive individuals within the field; neither had a positive relationship with shrub presence. Dispersal of heavy‐seeded Quercus spp. and Carya aquatica was limited to within 20 m of the forest edge. These results imply that dispersal limitation is diminishing in wind‐ and bird‐dispersed species with maturation of in‐field recruits and that shrub patches may influence these patterns. Heavy‐seeded species, however, remain restricted to field edges that directly abut a seed source. If canopy closure by wind‐ and bird‐dispersed species precedes dispersal of heavy‐seeded species into the field, establishment of Quercus and Carya spp. may remain low for the foreseeable future.     

Dispersal of conidia of Dothidella ulei from Hevea brasiliensis

South American leaf blight caused by Dothidella ulei occurs only in tropical America, on both indigenous and cultivated Hevea spp. The conidium (Fusicladium macrosporum) is a 1-septate, dry, air-borne spore about 40 × 7 μ, occurring on the abaxial surface of dry leaves in dense, powdery, olive-green masses, and with one or both cells collapsed. The conidia adhere to the surface of water droplets, becoming turgid, and are disseminated in splash droplets. A Hirst volumetric trap, placed within a prepared source in north-west Trinidad, showed a diurnal periodicity of conidial production, with a maximum at 10.00 h and minima at night or in the early morning. On rainless days there was also a minor peak at 20.00 h. Transient increases occurred after rain, most of which fell around noon. On wet days almost equal numbers of conidia were dispersed between 10.00 and 12.00 h. Large increases occurred in 87% of all rain showers between 09.00 and 13.00 h. After 13.00 h fewer rain showers caused such increases; the lowest (36%) was between 21.00 and 01.00 h. Twice as many were trapped on sunny days (> 9 h sun) at 09.00 h when there was full sunshine, compared with overcast days (< 5 h sun). A more clearly defined morning maximum occurred on relatively windy days, compared with calmer ones. Conidial sporulation became very low, or ceased, where rain fell below a mean of 3–4 mm per day for at least 20 days. Abundant sporulation occurred with a daily rainfall about twice this amount. The results support the belief that if Dothidella ulei appeared in Malaysia its spread would be rapid and its effects damaging.     

Seed dispersal of the non-native invasive tree <Emphasis Type="Italic">Ailanthus altissima</Emphasis> into contrasting environments

Ailanthus altissima has a long history of invasion in urban areas and is currently spreading into suburban and rural areas in the eastern U.S. The objectives of our study were to (1) determine whether A. altissima seed dispersal distance differed between populations on the edges of open fields and intact deciduous forest, and (2) determine whether dispersal differed for north and south winds. We also assessed the relationship between seed characteristics and distance from source populations in fields and forests, and whether seeds disperse at different rates throughout the dispersal season. Using two fields, two intact forest stands, and one partially harvested stand, we sampled the seed rain at 10 m intervals 100 m into each site from October to April 2002–2003. We compared seed density in field and intact forests using a three-way ANOVA with distance from source, wind direction, and environmental structure as independent variables. To assess the accuracy of common empirical dispersal models, mean seed density data at each site were fitted with alternative regression models. We found that mean seed dispersal distance depended on environmental structure and wind direction, a result driven in large part by dispersal at a single site where seed density did not decline with distance. The two alternative regression models fit each site’s dispersal curve equally well. More seeds were dispersed early than in mid- or late-season. Large, heavy seeds traveled as far as small light seeds. Turbulent winds appear to be necessary for seed release, as indicated by a wind tunnel experiment. A. altissima is able to disperse long distances into fields and into mature forests, and can reach canopy gaps and other suitable habitats at least 100 m from the forest edge. It is an effective disperser and can spread rapidly in fragmented landscapes where edges and other high light environments occur. These conditions are increasingly common throughout the eastern U.S. and in other temperate regions worldwide.     

Movement of apterous Sitobion avenae (Homoptera: Aphididae) in response to leaf disturbances caused by wind and rain

The effect of wind and rain on initiation of dispersal and subsequent distances moved by apterous Sitobion avenae was quantified in five laboratory experiments. Wind duration and type (steady and gusting), rain duration and intensity, and a combination of wind and rain were investigated. With increased duration of wind there was an increase in the average distance aphids moved. There was less movement when aphids were exposed to gentle gusting, than to steady wind or strong gusting. With increased duration of rain there was an increase in the proportion of aphids lost from plants. Under heavy rain, the proportion of aphids off the release plant, the distance moved by them and the proportion of aphids lost, was greater than for drizzle or light rain. When wind and rain were compared, rain had the greater impact on aphid dispersal. It was concluded that leaf disturbances caused by strong gusts of wind or large rain droplets are of considerable importance in the initiation of aphid dispersal, but that wetness alone is not.     

Predicting the time to colonization of the parasitoid Diadegma semiclausum: The importance of the shape of spatial dispersal kernels for biological control

The time at which natural enemies colonize crop fields is an important determinant of their ability to suppress pest populations. This timing depends on the distance between source and sink habitats in the landscape. Here we estimate the time to colonization of sink habitats from a distant source habitat, using empirical mark-capture data of Diadegma semiclausum in Broccoli. The data originated from experiments conducted at two locations and dispersal was quantified by suction sampling before and after a major disturbance. Three dispersal kernels were fitted to the dispersal data: a normal, a negative exponential, and a square root negative exponential kernel. These kernels are characterized by a thin, intermediate and a fat tail, respectively. The dispersal kernels were included in an integro-difference equation model for parasitoid population redistribution to generate estimates of time to colonization of D. semiclausum in sink habitats at distances between 100 and 2000 m from a source. We show that the three dispersal kernels receive similar support from the data, but can produce a wide range of outcomes. The estimated arrival time of 1% of the D. semiclausum population at a distance 2000 m from the source ranges from 12 days to a length of time greatly exceeding the life span of the parasitoid. The square root negative exponential function, having the thickest tail among the tested functions, gave the fastest spread and colonization in three of the four data sets, but it gave the slowest redistribution in the fourth. In all four data sets, the rate of accumulation at the target increased with the mean dispersal distance of the fitted kernel model, irrespective of the fatness of the tail. This study underscores the importance of selecting a proper dispersal kernel for modelling spread and colonization time of organisms, and of the collection of pertinent data that enable kernel estimation and that can discriminate between different kernel shapes.     

Population genetic structure and larval dispersal potential of spottedtail goby Synechogobius ommaturus in the north‐west Pacific

Larval dispersal may have an important effect on genetic structure of benthic fishes. To examine the population genetic structure of spottedtail goby Synechogobius ommaturus, a 478 base pair (bp) fragment of the hypervariable portion of the mtDNA control region was sequenced and used to interpret life‐history characteristics and larval dispersal strategy. Individuals (n = 186) from 10 locations on the coasts of China and Korea were analysed and 44 haplotypes were obtained. The levels of haplotype and nucleotide diversity were higher in East China Sea populations than in other populations. Both the phylogenetic tree and the minimum spanning tree showed that no significant genealogical structures corresponding to sampling locations existed. AMOVA and pair‐wise F_ST revealed significant genetic differentiation between populations from Korea and China. A significant isolation by distance pattern was observed in this species (r = 0·53, P < 0·001). Both mismatch distribution analysis and neutrality tests showed S. ommaturus to have experienced a recent population expansion. These results suggest that the Pleistocene ice ages had a major effect on the phylogeographic pattern of S. ommaturus, that larvae might avoid offshore dispersal and that dispersal of larvae may maintain a migration–drift equilibrium.     

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.