1. In forest communities, the Janzen-Connell (J-C) hypothesis proposes that species diversity is maintained by non-competitive distance-and/or density-dependent seedling mortality caused by host-specific natural enemies. However, the effects of pathogen associations from nearby conspecifics versus heterospecifics remain unknown in spatially heterogeneous light environments.2. Seeds of hardwood species Cornus controversa (Cornus) and Prunus grayana (Prunus) were sown beneath 6-7 Cornus and Prunus adults in both the forest understory (FU) and in gaps (Gap) created by felling all woody vegetation near the focal adults. Seedling growth, mortality, killing agents (e.g. pathogens that cause damping-off and leaf diseases), and root infection by arbuscular mycorrhizal fungi (AMF) were investigated. 3. We found strong habitat effects on the expression of soil fungi beneath both tree species. Seedling mortality caused by soil-borne damping-off pathogens was greater in FU than in Gap, and AMF infection, which enhanced relative seedling growth rate, was greater in Gap than in FU. Seedling mortality caused by damping-off pathogens did not differ between Cornus and Prunus seedlings beneath the adults of conspecific or heterospecific adults in both FU and Gap, suggesting little distance-dependence or host preference in the fungus. 4. Beneath the adults of Cornus and Prunus, the most prevalent leaf diseases were zonate leaf blight and angular leaf spot caused by the airborne pathogenic fungi Haradamyces foliicola and Phaeoisariopsis pruni-grayanae, respectively. Although these pathogens attacked the seedlings of both species, conspecific seedlings (i.e. home) showed more severe leaf damage, earlier leaf shedding and/or less defensive behaviour (cell wall defence) relative to heterospecific seedlings (i.e. away), suggesting negative distance-dependent attack (i.e. host preference) for these leaf diseases. As a result, greater seedling mortality was observed for conspecific seedlings under both FU and Gap treatments. 5. Synthesis. In the temperate forest, the J-C hypothesis is largely mediated through the strong negative influence of airborne leaf diseases rather than through soil-borne damping-off pathogens. We found that airborne diseases demonstrated distance-dependent host preferences, which led to greater conspecific seedling damage regardless of environmental light conditions.
In forests, negative density/distance-dependent seedling mortality (NDD) caused by natural enemies plays a key role in maintaining species diversity [Janzen-Connell (J-C) model]. However, the relative importance of natural enemies in mediating NDD under heterogeneous light conditions has remained unclear. We examined the relative importance of pathogens (i.e., soil pathogens, leaf diseases) on seedling performance in forest understories (FUs) and gaps (gaps) during a 3-year period (results of first year of our study have been previously reported). For the hardwood, Prunus grayana, we investigated seedling mortality, morbidity agents, growth, and root infection by arbuscular mycorrhizal fungi (AMF) beneath conspecific and heterospecific adults in FUs and gaps. Seedling mortality was higher beneath conspecific than heterospecific adults throughout 3 years at both sites, mainly due to continuous leaf disease (i.e., angular leaf spot), whereas damping-off diseases caused mortality only in the first year. Beneath each adult, seedling mortality was higher in FUs than in gaps until second year, but it did not differ between two habitat types in the third year, because leaf diseases caused severe damage even in gaps. Seedling mass was significantly lower beneath conspecific adults. AMF infection of seedlings was also lower beneath conspecific adults, while it was higher in gaps than in FUs beneath both adults. This study demonstrates that the J-C model in a hardwood tree, P. grayana is mainly driven by high NDD seedling mortality caused by airborne leaf diseases, which continuously attack seedlings in a NDD manner regardless of environmental light conditions.
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