Seed Removal and Predation as Factors Affecting Seed Availability of Tree Species in Degraded Habitats and Restoration Plantings in Rainforest Areas of Queensland,Australia

The post‐dispersal removal or predation of seeds of native tree species was investigated in Queensland, Australia, at degraded habitats and rainforest restoration sites where direct seeding might be used to facilitate tree regeneration (old fields or open habitats, lantana thicket, rainforest edge, and 5‐ and 10‐year‐old restoration plantings). Seed removal/predation was assessed in relation to tree seed weight and canopy density of the habitats during the wet season period. Results indicated that seed removal/predation imposes limitations on seed availability, particularly for small seeded species. In most situations, larger seeds were less removed/predated, most likely due to the more limited range of large seed consumers. The use of large, hard‐coated seeds may potentially reduce seed loss in open situations (from both seed removal and desiccation), unless large seed consumers frequent the site. Canopy cover exerted an influence on seed removal/predation, though trends varied in relation to site and the time of season. Broadcast sowing of seed under planted tree canopies at the more advanced stages of closure may in some areas result in higher seed removal/predation. Likewise, seeding in areas dominated by woody weeds may result in high seed losses to consumers such as rodents. Results suggested that undertaking direct seeding to coincide with the maximal period of fruit production may in some situations be beneficial to minimize seed loss. Overall, site context, canopy cover, and species selection appear to be important considerations when aiming to reduce loss of seeds to animal seed consumers in restoration work.     

Seed Dispersal by Avian Frugivores: Non‐random Heterogeneity at Fine Scales

Seed dispersal studies have primarily examined dispersal as a function of distance from the parent tree and/or heterogeneity in dispersal due to animal use of nesting, roosting and sleeping sites. However, non‐random heterogeneity in seed dispersal is also likely to result from the post‐foraging behavior and movement of frugivores which prefer certain trees. To characterize variation in seed rain at fine scales, we studied the dispersal curve of Prunus ceylanica, a primarily bird‐dispersed species. We compared seed rain at conspecifics, heterospecific fruiting trees with similar frugivore assemblages, emergent trees, and the landscape surrounding these trees. Seed rain of P. ceylanica was found to peak globally under the canopy of conspecifics but to peak locally under the canopy and immediate neighborhood of heterospecific fruiting trees. Our results demonstrate that seed rain is highly clumped even at fine spatial scales. A large proportion of seeds are dispersed in specific, localized regions. This variation can have important implications for plant population dynamics and might significantly alter the impact of post‐dispersal processes. Seed dispersal models may need to incorporate this heterogeneity to explain manifestations of spatially explicit dynamics like mixed species ‘orchards’.     

Seed Rain and Seed Limitation in a Planted Gallery Forest in Brazil

With seeds collected monthly during one year from 53 1‐m² seed traps, we investigated the seed rain and seed limitation in a gallery forest planted in 1994 in SE Brazil. Contrasting animal‐ (zoochorous) and wind‐dispersed (anemochorous) plants we investigated (1) which aspects of the composition and structure of the vegetation influence the abundance and species richness of the seed rain; (2) if such influences differ between zoochorous and anemochorous seeds; (3) if the abundance and richness of the seed rain sampled under zoochorous and nonzoochorous plant species differ; and (4) if seed limitation (given by the proportion of sites to which seeds were not dispersed) differs between zoochorous and anemochorous plant species, and also between species that have been planted and those that further colonized the area (colonists). Seed rain was intense and dominated by anemochorous species. The overall seed rain was not influenced by the vegetation parameters we analyzed (canopy height and cover, plant size, abundance, and richness) or by the plant species above the seed trap. The abundance and richness of zoochorous seeds in a given spot was influenced by the abundance and richness of zoochorous plants in its immediate vicinity. Seed limitation was higher for anemochorous than zoochorous species and higher for planted than for colonist species. We concluded with recommendations for the initial establishment of a planted forest, including the homogeneous distribution of zoochorous plants to permit a spatially homogeneous zoochorous seedfall, which will likely enhance the chances of survival and successful establishment of seeds.     

Seed predation and dispersal in relict Scots pine forests in southern Spain

For two years, the seed rain and magnitude of seed losses due to predation were evaluated in Scots pine forests in southern Spain. The Crossbill was the most important pre-dispersal predator, consuming more than 80% of ripening seeds. In addition, other birds, mainly Tits and Siskin, also consumed seeds just before seed dispersal, reaching values of 16 and 51% losses in 1996 and 1997, respectively. Seed rain was monitored in different microhabitats (under pine canopies, under shrubs and in open areas), and was most intense under the canopy of mother plants both years. Post-dispersal seed predators (rodents and birds) consumed up to 96% of seeds reaching the ground. Both pre- and post-dispersal seed predators preferentially harvested filled seeds. Post-dispersal predation was similarly intense in all microhabitats, so predators did not change the spatial distribution of the seed rain. These high predation rates were constant between years, localities and habitats (woodland and treeline). We hypothesize that this high rate of seed predation is a major factor limiting the regeneration of these relict populations of Scots pine in its southernmost limit.     

Neighbourhood analyses of tree seed predation by introduced rodents in a New Zealand temperate rainforest

House mice Mus musculus and other introduced rodents represent a novel source of predation on tree seeds in New Zealand forests. In the northern temperate forests where these rodents are native, spatial and temporal variation in tree seed production can result in dramatic fluctuations in the distribution and abundance of seed predators, with subsequent feedbacks on the distribution and abundance of seedlings. We use neighbourhood models to examine variation in rodent predation on seeds of 4 tree species of the temperate rainforests of New Zealand as a function of 1) spatial variation in local canopy composition and 2) spatial and temporal variation in mouse activity. We placed seeds throughout mapped stands of mixed forests in alluvial valley bottoms and on elevated marine terraces in the Waitutu Forest, South Island. The risk of predation on seeds of 2 dominant canopy trees – rimu Dacrydium cupressinum and mountain beech Nothofagus solandri var . cliffortioides – peaked in neighbourhoods dominated by those species and by silver beech N. menziesii , particularly in a year of plentiful seed rain from these species. The risk of predation on rimu and beech seed was also related to measures of local mouse activity. These relationships suggest that the highest local abundance of mice was concentrated in rimu and beech neighbourhoods because of the food provided by seed rain from those trees. Predation on seed of miro Prumnopitys ferruginea , which is eaten by rats but not mice, was low in rimu neighbourhoods and where mouse activity was high. These patterns may reflect spatial segregation in the activity of rats versus mice within stands. Our results suggest that the spatial distribution of canopy trees translates into predictable patterns of variation in mouse activity and seed predation. Heterogeneity in rodent activity and seed predation within stands may have important implications for tree population dynamics.     

Reproductive potential and seedling establishment of the invasive alien tree Schinus molle (Anacardiaceae) in South Africa

Schinus molle (Peruvian pepper tree) was introduced to South Africa more than 150 years ago and was widely planted, mainly along roads. Only in the last two decades has the species become naturalized and invasive in some parts of its new range, notably in semi‐arid savannas. Research is being undertaken to predict its potential for further invasion in South Africa. We studied production, dispersal and predation of seeds, seed banks, and seedling establishment in relation to land uses at three sites, namely ungrazed savanna once used as a military training ground; a savanna grazed by native game; and an ungrazed mine dump. We found that seed production and seed rain density of S. molle varied greatly between study sites, but was high at all sites (384 864–1 233 690 seeds per tree per year; 3877–9477 seeds per square metre per year). We found seeds dispersed to distances of up to 320 m from female trees, and most seeds were deposited within 50 m of putative source trees. Annual seed rain density below canopies of Acacia tortillis, the dominant native tree at all sites, was significantly lower in grazed savanna. The quality of seed rain was much reduced by endophagous predators. Seed survival in the soil was low, with no survival recorded beyond 1 year. Propagule pressure to drive the rate of recruitment: densities of seedlings and sapling densities were higher in ungrazed savanna and the ungrazed mine dump than in grazed savanna, as reflected by large numbers of young individuals, but adult : seedling ratios did not differ between savanna sites. Frequent and abundant seed production, together with effective dispersal of viable S. molle seed by birds to suitable establishment sites below trees of other species to overcome predation effects, facilitates invasion. Disturbance enhances invasion, probably by reducing competition from native plants.     

Seed rain of fleshy and dry propagules in different habitats in the temperate rainforests of Chiloé Island,Chile

In temperate rainforests on Chiloé Island in southern Chile (42°S), most canopy trees bear fleshy, avian‐dispersed propagules, whereas emergent tree species have dry, wind‐borne propagules. In the present study, the following hypothesis was tested: regardless of species, fleshy propagules are deposited in greater numbers in canopy gaps and in forest margins and hence have a more heterogeneous seed shadow than wind‐dispersed propagules. To test this hypothesis, the seed rains of these two types of propagules were compared in the following forest habitats: (i) tree‐fall gaps (edges and centre); (ii) forest margins with adjacent pastures; and (iii) under closed canopy (forest interior). Seed collectors (30‐cm diameter) were placed in two (15 and 100 ha) remnant forest patches (n = 60–100 seed collectors per patch) distributed in the four habitats. Seeds were retrieved monthly from each collector during two reproductive seasons (1996, 1997). In both years, the seed rain was numerically dominated by two species with dry propagules (Laureliopsis philippiana and Nothofagus nitida) and three species with fleshy fruits (Drimys winteri, Amomyrtus luma, and Amomyrtus meli). The seed shadows of the two species with dry, wind‐dispersed seeds differed markedly. Seeds of L. philippiana were deposited predominantly in canopy openings, whereas N. nitida seeds fell almost entirely in the forest interior. The fleshy‐fruited species, Drimys and Amomyrtus spp., had similar seed deposition patterns in the various habitats studied, but the between‐year differences in seed rain were greater in Drimys winteri than in Amomyrtus spp. Although no more than 10% of fleshy‐fruited propagules reached the margins of the patch, approximately 7% of these were carried there by birds. Every year, canopy gaps (pooling data from edges and centres) concentrated approximately 60% of the total seed rain of both propagule types in both forest patches. Forest margins received less than 20% of the total seed rain, which was largely dominated by fleshy‐fruited species. Seed shadows were a species‐specific attribute rather than a trait associated with propagule type and dispersal mode.     

Seed dynamics of resprouting shrubs in grassy woodlands: Seed rain,predators and seed loss constrain recruitment potential

Abstract Measuring the fate of seeds between seed production and seedling establishment is critical in understanding mechanisms of recruitment limitation of plants. We examined seed fates to better understand the recruitment dynamics of four resprouting shrubs from two families (Fabaceae and Epacridaceae) in temperate grassy woodlands. We tested whether: (i) pre‐dispersal seed predation affected seed rain; (ii) post‐dispersal seed predation limited seed bank accumulation; (iii) the size of the seed bank was related to seed size; and (iv) viable seeds accumulated in the soil after seed rain. There was a distinct difference in seed production per plant between plant families with the legumes producing significantly more seeds per individual than the epacrids. Seed viability ranged from 43% to 81% and all viable had seed or fruit coat dormancy broken by heat or scarification. Pre‐dispersal predation by Lepidopteran larvae removed a large proportion of seed from the legume seed rain but not the epacrids. Four species of ants (Notoncus ectatomoides, Pheidole sp., Rhytidoponera tasmaniensis and Iridomyrmex purpureus) were major post‐dispersal seed removers. Overall, a greater percentage of Hardenbergia (38%) and Pultenaea (59%) seeds were removed than the fleshy fruits of Lissanthe (14%) or Melichrus (0%). Seed bank sizes were small (<15 seeds m^?2) relative to the seed rain and no significant accumulation of seed in the soil was detected. Lack of accumulation was attributed to seed predation as seed decay was considered unlikely and no seed germination was observed in our study sites. Our study suggests that seed predation is a key factor contributing to seed‐limited recruitment in grassy woodland shrubs by reducing the number of seeds stored in the soil.     

西双版纳望天树林种子雨9年动态

种子雨是森林生态系统过程的重要组成部分。虽然国内外已有部分关于种子雨的研究, 但多数集中在温带和亚热带地区, 热带地区种子雨的长期监测研究仍显不足。为全面了解热带森林种子雨输入的数量动态和物种组成的长期规律, 本文对我国西双版纳望天树(Parashorea chinensis)林群落水平上9年(2008年5月至2017年4月)的种子雨输入进行了研究。9年共收集到木本植物的种子226种8,993,224粒, 其中大型种子(I-1型) 77种9,779粒; 中型种子(I-2型) 61种24,237粒; 小型种子(I-3型) 72种83,399粒; 粒状微小种子(II-1型) 13种1,215,235粒; 粒状极小种子(II-2型) 1种7,244,667粒; 细丝状种子(II-3型) 2种415,907粒。种子雨输入量排名前10位的优势种贡献了70%以上的种子数量。各类型种子的种子雨输入量均具有年际波动, 呈“大小年”的交替变化, 间隔期为1-2年, 物种组成也呈现一定的年际差异。持续9年的监测中, 每类型种子至少观察到了1次大年结实事件。     

Postdispersal seed predation and seed viability in forest soils: implications for the regeneration of tree species in Ethiopian church forests

Almost all dry Afromontane forests of Northern Ethiopia have been converted to agricultural, grazing or scrub lands except for small fragments left around churches (‘Church forests’). Species regeneration in these forests is limited. We investigated (i) how intense postdispersal seed predation was in church forest, and if this seed predation varied with species and/or habitat, and (ii) for how long tree seeds maintained their viability while buried in forest soil. In the seed predation experiment, we monitored seeds of six tree species in four habitats for a period of 14 weeks (the peak seeding season). In the seed viability experiment, we assessed seed viability of five species in four habitats after being buried 6, 12, or 18 months. Ninety‐two percent of the tree seeds were predated within 3.5 months. Predation was mainly dependent on species whereas habitat had a weaker effect. Seed viability decreased sharply with burial time in soil for all species except for Juniperus. To minimize seed availability limitation for regeneration of such species in the forest, the standing vegetation needs to be persistently managed and conserved for a continuous seed rain supply. Additional seed sowing, and seed and seedling protection (by e.g. animal exclosures) may increase successful regeneration of important species in these forests.     

Temporal and spatial patterns of seed dispersal in two Cistus species (Cistaceae)

Cistus species are obligate seeding, early colonizers that follow disturbance, particularly fire, in Mediterranean ecosystems. We studied seed release, seed dispersal and soil seed populations in stands of Cistus ladanifer and C. libanotis. Seed release started in mid- to late summer (C. ladanifer) or in early autumn (C libanotis), and continued for a very extended period: 8-10 months in C. ladanifer, and for a mean of 16 months in C. libanotis. The xerochastic capsules of both species released seeds by successive dehiscence of the locules. All capsules begin to dehisce simultaneously at the start of the seed release period, but in C. libanotis capsule fragmentation replaced dehiscence early in the seed release period. In plants of both species, seed shadows were characterized by a peak of density beneath the plant canopy and a very short tail of much lower densities, indicating that seeds are concentrated beneath mother plants when dispersed. Nevertheless, in late May, at the onset of the fire season, soil seed densities beneath plant canopies were low compared with densities expected from seed shadows, but were apparently high enough to allow recovery of the stands if a disturbance, such as fire, had taken place. Seed-eating Bruchidae in summer and granivorous ants during the seed release period were apparently the main causes of seed losses. Results suggest that in both Cistus species, the staggered seed release could constitute an efficient risk-reducing trait. The plant pool of seeds existing throughout most of the year could be a relevant component of Cistus seed banks.     

Differences in seed rain composition in small and large fragments in the northeast Brazilian Atlantic Forest

Tropical forests are seriously threatened by fragmentation and habitat loss. The impact of fragment size and forest configuration on the composition of seed rain is insufficiently studied. For the present study, seed rain composition of small and large forest fragments (8–388 ha) was assessed in order to identify variations in seed abundance, species richness, seed size and dispersal mode. Seed rain was documented during a 1‐year period in three large and four small Atlantic Forest fragments that are isolated by a sugarcane matrix. Total seed rain included 20,518 seeds of 149 species of trees, shrubs, palms, lianas and herbs. Most species and seeds were animal‐dispersed. A significant difference in the proportion of seeds and species within different categories of seed size was found between small and large fragments. Small fragments received significantly more very small‐sized seeds (<0.3 cm) and less large‐seeded species (>1.5 cm) that were generally very rare, with only one species in small and eight in large fragments. We found a negative correlation between the inflow of small‐sized seeds and the percentage of forest cover. Species richness was lower in small than in large fragments, but the difference was not very pronounced. Given our results, we propose changing plant species pools through logging, tree mortality and a high inflow of pioneer species and lianas, especially in small forest fragments and areas with low forest cover. Connecting forest fragments through corridors and reforestation with local large‐seeded tree species may facilitate the maintenance of species diversity.     

The effect of seed mass and gap size on seed fate of tropical rain forest tree species in guyana

Abstract: For eleven tree species, differing in seed mass, germination success (emergence success for two small-seeded species) and the causes of failure to germinate were studied in the forest understorey and in logging gaps in the tropical rain forests of Guyana. In the forest understorey, germination success increased with seed mass. However, as gap size increased the difference between smaller and larger seeded species diminished because germination success of smaller-seeded species increased slightly, while that of larger-seeded species decreased dramatically. The negative effect of gap size on germination success of larger-seeded species was caused by an increased risk of desiccation with gap size, which was a far more important seed mortality agent for larger than for smaller-seeded species. Generally, seeds of smaller-seeded species suffered more from insect predation and were removed at higher rates than larger-seeded species. On the other hand, larger-seeded species were eaten more by mammals than smaller-seeded species. It is concluded that logging can result in shifts in the species composition in the tropical rain forests of Guyana which are dominated by species with large seeds, since germination success of larger-seeded species is dramatically reduced in large logging gaps.     

大兴安岭中部天然次生林种子雨动态

本研究于2018—2019年对大兴安岭中部白桦林、针叶林和针阔混交林3种林型的种子雨落种量进行了动态监测,并对3种林型主要树种的种子雨季节动态、落叶动态、种子雨千粒重、种子雨年际变化和种子雨空间格局进行了分析。结果表明: 各林型中兴安落叶松种子雨和白桦种子雨呈现明显的单峰型分布。针阔树种(落叶松、樟子松、云杉、白桦、山杨)的落叶量呈现出明显的季节动态,各林型落叶量大多在9月中上旬达到高峰。在针阔混交林和针叶林中,处于高峰期的兴安落叶松种子雨千粒重明显大于初始期和末尾期的兴安落叶松种子雨千粒重,3种林型下白桦千粒重在季节上未表现出明显的差异。兴安落叶松和白桦的种子雨均呈现出明显的年际变化,2018年为种子散种量的丰年,2019年为歉年。两年时间内,所有种子雨的空间格局在总体上均表现为聚集分布,种子雨和幼苗幼树在空间分布格局上存在一致性。     

Offspring spatial patterns in Picconia excelsa (Oleaceae) in the Canarian laurel forest

We studied the spatial patterns of seedlings and seeds in isolated Picconia excelsa (Oleaceae) trees in the laurel forest of Anaga, Tenerife (Canary Islands). By finding isolated trees we assessed the correlation of seed and seedling bank traits and parent trees by removing the confounding effects of proximity (<100 m radius) of conspecific fruiting trees. We counted all the seedlings per age (height) class within its parental range, and sampled the seed number along transects departing from beneath the parent canopy at regular intervals. We mapped all seedlings per age class and plotted seed and seedling profiles in relation to distance to parent trees. Older Picconia seedlings tended to clump significantly further from parent trees than younger seedlings, which clumped just beneath the parents. We found significant differences among distances to parent tree in numbers of seedlings per age class. The seedling bank area was significantly correlated with maximum distance of seedlings to parent trees. The majority of seeds were deposited within the first 4 m below the parent crown. Seedlings amount at further distances from the trees is larger than seeds/fruits as counted on the ground. Our results suggest that disseminated, older seedlings have occupied germination sites far from the parent tree because there is probably lower seedling–seedling and parent–seedling competition for resources, and perhaps no intraspecific allelopathy and predation/disease.     

Effect of Bat Exclusion on Patterns of Seed Rain in Tropical Rain Forest in French Guiana

To assess the impact of bats on seed dispersal in a tropical mature forest (Nouragues, French Guiana), we conducted a bat exclusion experiment and tested the hypotheses that an artificial reduction in the abundance of bats would result in: (1) a decrease in seed species diversity, at the community level; and (2) an increase in seed limitation (a failure of seeds to reach all suitable sites for germination) at the species level. Seed rain was sampled in two contiguous forest plots for a total of 120 d, using 49 seed traps (1 m²) arranged in 7 × 7 grids and spaced at 5-m intervals. Using mist nets, bat activity was reduced in one forest plot for a total of 60 nights. Thirty-nine plant species, or species groups, likely to be consumed and dispersed by bats, were collected within a total sample of 50,063 seeds. The overall seed rain was dominated by epiphytic Araceae and Cyclanthaceae (83.3%) and tree species within the genera Cecropia and Ficus (16.0%). Seeds from bat-dispersed shrubs and treelets ( Piper , Solanum , and Vismia ) were relatively rare (0.7%). The bat exclusion resulted in a 30.5 percent reduction in seed species richness and increased seed limitation for most of the species sampled. Seed limitation was caused mainly by a reduced seed rain (seed source limitation) rather than a decrease in seed dispersal uniformity (seed dispersal limitation). Therefore, bat-dispersed plants with low seed production are likely to be particularly affected by a decline in bat abundance, as a result of anthropogenic change.     

六盘山区辽东栎的实生苗更新及其影响因子

研究了六盘山区辽东栎(Quercus liaotungensis)灌丛的种子产量、土壤种子库组成、湿沙和风干贮藏对种子寿命的影响以及动物取食子叶对种子萌发和幼苗建立的影响。结果表明, 辽东栎灌丛种子的完好率为27.51%, 被动物取食或搬运种子的比例(41.51%)显著高于其他类型种子(p < 0.01); 辽东栎次生林土壤种子库中萌发和虫蛀种子分别占35.16%和38.29%, 完好种子仅占13.65%, 捕食动物主要通过贮藏或搬运而影响土壤种子库中的种子密度。湿沙贮藏60天的辽东栎种子自动萌发率高达96.67%, 短期贮藏可加快种子的萌发进程, 提高萌发率和萌发指数, 但随着贮藏时间的延长, 种子萌发进程延迟, 萌发率、萌发指数和活力指数均不同程度地降低。排除动物取食处理的幼苗在林窗和林下生境的存活率分别为80%和83%, 而不排除动物取食幼苗在2种生境中分别仅有25%和31%能够存活, 表明子叶在幼苗建立中具有重要作用。林窗中幼苗子叶的动物取食率(85.00%)高于林下(71.00%), 子叶留存的幼苗在林窗中的存活率(6.00%)低于林下(15.50%), 而子叶被取食幼苗的存活率在两种环境中基本相等(分别为18.50%和18.00%)。     

More than just drought: complexity of recruitment patterns in Mediterranean forests

Understanding community dynamics during early life stages of trees is critical for the prediction of future species composition. In Mediterranean forests drought is a major constraint for regeneration, but likely not the only factor determining the observed spatial patterns. We carried out a sowing experiment aimed at identifying main filters during seed-seedling transition. Specifically, we studied seed fate (predation, fungi infection, emergence) and subsequent seedling performance (mortality during the first summer and overall recruitment after 2 years) of four co-occurring Mediterranean tree species (Quercus ilex, Quercus faginea, Juniperus thurifera, Pinus nigra). We related these processes to the dominant species composition, microhabitat heterogeneity, herb cover and seed mass. The identity of the dominant species in the forest canopy was more important for recruitment than the forest canopy being dominated by conspecific vs. heterospecific species. The patterns we found suggest that biotic interactions such as facilitation (lower mortality under the canopies) and herb competition (during emergence of J. thurifera) are relevant during recruitment. Moreover, our results pointed to ontogenetic conflicts regarding the seed mass of Q. faginea and to density-dependent seed mortality for Q. ilex, rarely described in Mediterranean ecosystems. We propose that our study species experience population growth in forests dominated by heterospecifics where the recruitment success depends on habitat heterogeneity and on moderated biotic and abiotic stresses created by each species. Our results reveal patterns and mechanisms involved in recruitment constraints that add complexity to the well-known drought-related processes in Mediterranean ecosystems.     

北京东灵山落叶阔叶林中辽东栎种子雨

在北京东灵山地区的一个落叶阔叶林中调查了辽东栎（Ｑｕｅｒｃｕｓ ｌｉａｏｔｕｎｇｅｎｓｉｓ Ｋｏｉｄｚ．）的种子雨。对于选定的４棵辽东栎中的３棵,树冠下的种子雨分布格局符合二次分布,具有很高的决定系数。由设置在树冠下的种子捕捉器收集的坚果数量来估计整棵树的种子雨。４棵树的种子雨中有活力的种子很少,变化范围从２６到２５９个。每棵树的树冠下的种子雨密度变化范围从０．７６到７．２６个／ｍ＾２。林中平均种     

Trophic and non-trophic pathways mediate apparent competition through post-dispersal seed predation in a Patagonian mixed forest

Post‐dispersal predation can be a major source of seed loss in temperate forests. Little is known, however, about how predator‐mediated indirect interactions such as apparent competition alter survival patterns of canopy tree seeds. Understorey plants may enhance tree seed predation by providing sheltered habitat to granivores (non‐trophic pathway). In addition, occurrence of different tree seeds in mixed patches may lead to short‐term apparent competition between seed types, because of the granivores’ foraging response to changes in food patch quality (trophic pathway). We hypothesised that understorey bamboo cover and mixing of seed species in food patches would both increase tree seed predation in a Nothofagus dombeyi?Austrocedrus chilensis forest in northern Patagonia, Argentina. Seed removal experiments were conducted for three consecutive years (2000–2002) differing in overall granivory rates. Seed patch encounter and seed removal rates were consistently higher for the larger and more nutritious Austrocedrus seeds than for the smaller Nothofagus seeds. Seed removal was greater beneath bamboo than in open areas. This apparent competition pathway was stronger in a low‐predation year (2000) than in high‐predation years (2001–2002), suggesting a shift in microhabitat use by rodents. Patch composition had a significant, though weaker, impact on seed survival across study years, whereas seed density per patch enhanced encounter rates but did not influence seed removal. Removal of the less‐preferred Nothofagus seeds increased in the presence of Austrocedrus seeds, but the reciprocal indirect effect was not observed. However, this non‐reciprocal apparent competition between seed species was only significant in the high‐predation years. Our study shows that granivore‐mediated indirect effects can arise through different interaction pathways, affecting seed survival patterns according to the predator's preference for alternative seed types. Moreover, results indicate that the occurrence and relative strength of trophic vs non‐trophic pathways of apparent competition may change under contrasting predation scenarios.