Geographical variation in the early regeneration process of Siebold's Beech (Fagus crenata BLUME) in Japan

The causes and timing of seed death in early regeneration process of Siebold's beech (Fagus crenata Blume) was studied at 15 sites along a snowfall gradient in Japan, in order to clarify why the seedling density of the species has geographic difference remarkably. Seed production did not significantly differ along the snowfall gradient. Pre-dispersal seed mortality by insect damage was higher at sites with light snowfall than at sites with heavy snowfall, but this only seemed to be a minor factor influencing the population. A large proportion of the viable nuts that fall in autumn ware killed in winter before germination. Winter mortality was much higher at sites with thin snow cover than that at sites with thick snow cover, and this factor was strongly correlated with the geographic variation of seedling regeneration probability. There was little seed mortality by winter desiccation. The main factor contributing to the geographic difference seemed to be a seed predation by rodents in winter. Deep snow cover may reduce the success of rodents finding seeds in winter. Thus the observed relationship between snowpack depth and early mortality may be due to an indirect effect through the process of seed predation.p>     

Regeneration dynamics, causal factors, and characteristics of pacific ocean-type beech (Fagus crenata) forests in Japan: A review

This paper reviews the differences in the distribution and regeneration ofFagus crenata between two types of Japanese beech forests, the Japan Sea (JS)-type and the Pacific Ocean (PO)-type, and discusses the causal factors and characteristics of these forests, particularly the PO-type.F. crenata in PO-type forests regenerates sporadically rather than constantly, whereas regeneration in the JS-type forests is relatively constant with gap dynamics.F. crenata has dominated in snowy areas both in the past, after the last glacial age, when there was less human disturbance, and in the present. Snow accumulation facilitates beech regeneration in snowy JS-type forests, but not in the less snowy PO-type. Snow protects beechnuts from damage caused by rodents, desiccation, and freezing. In addition, snow suppresses dwarf bamboo in the spring, thus increasing the amount of sunlight available for beech seedlings on the forest floor. Snow also supplies melt water during the growing season and limits the distribution of herbivores. Moreover, snow reduces the number of forest fires during the dry winter and early spring seasons. The low densities ofF. crenata impede its regeneration, because disturbed wind pollination lowers seed fertility and predators are less effectively satiated. In snowy JS-type beech forests,F. crenata dominates both at the adult and the juvenile stages because it regenerates well, while other species are eliminated by heavy snow pressure. On the less snowy PO-side, deciduous broad-leaved forests with various species are a primary feature, althoughF. crenata dominates because of its large size and long lifespan.     

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.     

Differences in beech (Fagus crenata) regeneration between two types of Japanese beech forest and along a snow gradient

Differences in beech (Fagus crenata) regeneration were quantitatively investigated using power function analysis for the size–class (diameter at breast height, DBH) distribution and juvenile-to-canopy tree (J/C) ratio along a snow gradient throughout Japan. In snowy areas, all species combined, as well as F. crenata alone, showed constant regeneration, with parameter b≈−1.6 for the power function y=ax ^b (x=DBH, y=density), which is related to the DBH–class distribution. The good fit of the data to the function suggests that beech regenerates constantly with self-thinning patch dynamics. Parameter a, which indicates the abundance of small trunks, was high. Furthermore, the mean J/C ratio was ≈8, i.e., each parent beech tree produced eight juveniles. These results suggest that beech regenerates constantly with gap dynamics in snowy beech forests on the Japan Sea side of Japan (snowy). However, the fit of F. crenata was lower and nonsignificant in some forests in less snowy areas, despite the high fit of all species combined. In these areas, the mean of a was low, and b was often near zero for F. crenata regressions. These results suggest that the abundance of beech was low, and self-thinning was not evident because of the initial low abundance. Moreover, the mean J/C ratio was <1.0, suggesting that juvenile density was lower than that of canopy trees. Thus, the regeneration of F. crenata on the Pacific Ocean side of Japan (less snowy) is rather sporadic. Less snowy conditions may promote seed desiccation, predation of beechnuts and seedlings, and water stress. Lower F. crenata density may also reduce predator satiation and wind pollination.     

Demographic response of tundra small mammals to a snow fencing experiment

Snow cover is a key environmental component for tundra wildlife that will be affected by climate change. Change to the snow cover may affect the population dynamics of high‐latitude small mammals, which are active throughout the winter and reproduce under the snow. We experimentally tested the hypotheses that a deeper snow cover would enhance the densities and winter reproductive rates of small mammals, but that predation by mustelids could be higher in areas of increased small mammal density. We enhanced snow cover by setting out snow fences at three sites in the Canadian Arctic (Bylot Island, Nunavut, and Herschel Island and Komakuk Beach, Yukon) over periods ranging from one to four years. Densities of winter nests were higher where snow depth was increased but spring lemming densities did not increase on the experimental areas. Lemmings probably moved from areas of deep snow, their preferred winter habitat, to summer habitat during snow melt once the advantages associated with deep snow were gone. Our treatment had no effect on signs of reproduction in winter nests, proportion of lactating females in spring, or the proportion of juveniles caught in spring, which suggests that deep snow did not enhance reproduction. Results on predation were inconsistent across sites as predation by weasels was higher on the experimental area at one site but lower at two others and was not higher in areas of winter nest aggregations. Although this experiment provided us with several new insights about the impact of snow cover on the population dynamics of tundra small mammals, it also illustrates the challenges and difficulties associated with large‐scale experiments aimed at manipulating a critical climatic factor.     

The effects of food availability and distance to protective cover on the winter foraging behaviour of tits (Aves: Parus)

To maximize fitness, many animals must trade off their need to forage efficiently against their need to avoid predators. We studied such a trade-off in four species of tits (Paridae) in a forest near Oxford, UK. During winter, tits form flocks which increase feeding efficiency and reduce predation risk. These flocks feed extensively on beech (Fagus sylvatica) seeds, the abundance of which may be critical for winter survival. Because these seeds drop to the ground, where birds are exposed to sparrowhawk (Accipiter nisus) attack, tits need to trade off their need to find seeds against the proximity to protective cover, provided by dense clusters of hawthorn (Crataegus spp.). The quality of the beech crop differs markedly between trees and years. During a year of abundant beechmast, most tits searched for seeds close to protective cover. This 'safety-first' strategy precluded visits to superabundant food patches if they were too far from protective cover. Among beech trees near to cover, tits tended to prefer those with high seed density. Tits benefited from foraging under trees with high seed density because this correlated significantly with seed mass per square metre and because mean search times decreased with increasing seed density. Finally, we show experimentally that great tits, Parus major, can discriminate between edible (viable) and inedible (empty) seeds.     

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.     

亚热带林区啮齿动物对樱桃种子捕食和搬运的作用格局

在都江堰林区,通过在原生林、次生林和灌丛3类生境释放野生和栽培樱桃(Cerasus pseudocerasus)的种子,研究了啮齿动物对樱桃种子的捕食及其对樱桃种群更新的作用。结果表明,啮齿动物对樱桃两类种子的搬运无显著差异,而啮齿动物对野生樱桃种子的收获则明显快于栽培樱桃种子,且在3类生境均有类似的趋势。这说明啮齿动物偏爱于收获具有较高收益(种仁重/种子重)的野生樱桃种子。啮齿动物在小于10 d的时间尺度收获了所有释放的樱桃种子,其中,70%以上为啮齿动物所搬运。春季食物的匮乏可能是导致啮齿动物对樱桃种子有较大捕食压力的主要原因,而生境类型间的差异较小。因此,啮齿动物是都江堰林区樱桃地表种子的主要捕食者,它们对野生樱桃种子的选择性捕食和搬运能影响樱桃种子/果实的进化及其种群更新。     

Wintering space use and site fidelity in a nomadic species,the snowy owl

Migratory species can exploit many habitats over vast geographic areas and adopt various patterns of space and habitat use throughout their annual cycle. In nomadic species, determinants of habitat use during the non‐breeding season are poorly known due to the unpredictability of their movement patterns. Here, we analysed variability in wintering space and habitat use by a highly nomadic species, the snowy owl, in eastern North America. Using 21 females tracked by satellite telemetry between 2007 and 2016, we 1) assessed how space use patterns in winter varied according to the type of environment (marine vs terrestrial), latitudinal zone (Arctic vs temperate), local snow conditions and lemming densities and 2) investigated winter habitat and site fidelity. Our results confirmed a high inter‐individual variation in patterns of habitat use by wintering snowy owls. Highly‐used areas were concentrated in the Arctic and in the marine and coastal environments. Owls wintering in the marine environment travelled over longer distances during the winter, had larger home ranges and these were divided in more smaller zones than individuals in terrestrial environments. Wintering home range sizes decreased with high winter lemming densities, use of the marine environment increased following high summer lemming densities, and a thick snow cover in autumn led to later settlement on the wintering ground. Contrary to expectations, snowy owls tended to make greater use of the marine environment when snow cover was thin. Snowy owls were highly consistent in their use of a given wintering environment and a specific latitudinal zone between years, but demonstrated flexibility in their space use and a modest site fidelity. The snowy owls’ consistency in wintering habitat use may provide them with advantages in terms of experience but their mobility and flexibility may help them to cope with changing environmental conditions at fine spatial scale.     

啮齿动物及凋落物存留对天山野果林野生樱桃李种子命运的影响

2010年9月至2013年4月通过设置人工土壤种子库,研究了野生樱桃李土壤种子库的动态及啮齿动物和凋落物覆盖对种子库中种子命运的影响.结果表明: 在有取食动物扰动下,48.3%的种子萌发输出为幼苗,50%的种子被动物搬运或当场取食,4%的种子腐烂.在排除了取食动物干扰的条件下,樱桃李种子形成了短期持久的土壤种子库,3年后依然有部分种子萌发并输出为幼苗.凋落物覆盖和对照处理中,被搬运和当场取食种子的比例均显著低于清除凋落物的裸地.地表凋落物存留能减少动物搬运、取食,但不足以导致新生幼苗的大量出现,而啮齿类动物的搬运或取食是影响野生樱桃李种子命运和种子库动态的主要因素.
     

Post‐dispersal seed depletion by rodents in marginal populations of yew (Taxus baccata): consequences at geographical and local scales

Post‐dispersal seed predation is a crucial phenomenon for plant recruitment, and its incidence can be hypothesized to increase in ecologically and geographically marginal populations of threatened species, such as yew (Taxus baccata). Here we examine the among‐ and within‐population patterns of seed consumption by rodents and evaluate to what extent they are linked to marginality in Mediterranean low‐density yew stands. Among populations we tested: (i) whether the rates of seed predation found in our marginal sites were consistently higher than in populations from core regions; (ii) within populations we evaluated whether rodents preferred microhabitats with greater seed availability (beneath female yew trees) or with lower predation risk (shrubs) in two seeding seasons (fall–winter 2005 and 2006). Predation rates were extremely high (92.5%) and they were well above values reported for core populations (65.4%), to the extent that rodents almost completely depleted the experimental seeds in all populations and years. Our expectation of lower predation rates with decreasing vegetation cover was also confirmed for all years and populations, suggesting that rodent foraging was risk‐sensitive. This microhabitat effect outweighed the effect of seed availability under female yew trees, implying also that rodents selectively consumed the most valuable seeds in terms of their recruitment prospects. Overall, our results suggest that the mechanisms underlying seed depletion and its demographic consequences are linked to the effects of reduced yew performance in ecologically marginal habitats.     

Seed consumption and dispersal of ant-dispersed plants by slugs

In beech-dominated forests in Central Europe, many spring geophytes show adaptations to seed dispersal by ants (myrmecochory). Ants, however, can be rare in such moist forests. Motivated by observations of slug feeding on seeds we investigated the seed consumption of two plant species, Anemone nemorosa and Asarum europaeum, by slugs, in a series of experiments. In a seed predation experiment in a beech forest, we found that seed removal was strongly reduced when gastropods were excluded from the seed depots. The contribution of insects, including ants, and rodents to seed removal was relatively less but differed between May and July. In the laboratory, slug species, in particular Arion sp., consumed seeds of both plant species. Slugs either consumed the elaiosomes of seeds or swallowed seeds intact. Swallowed seeds were defecated undamaged and germinated as well as control seeds when buried overwinter, indicating the potential for seed dispersal by slugs. We also recovered seeds of myrmecochores in the faeces of several slugs caught in forests. In a slug release experiment in the forest, slugs moved up to 14.6 m (mean 4.4 m) in 15 h, which is the median gut passage time of seeds based on measurements made in the laboratory. We also found that when slug-defecated seeds were offered to rodents, these were less attractive than control seeds, suggesting that passage through the slug gut reduces seed predation risk. Our results demonstrate that slugs are significant consumers of elaiosomes or entire seeds of ant-dispersed plants and that they can function as seed dispersers of these plants.     

Early reduction of post-fire recruitment of Pinus nigra by post-dispersal seed predation in different time-since-fire habitats

This study analyses the effects of post-dispersal predation of Pinus nigra seeds on the initial recruitment of this species in areas burned by large wildfires, where P. nigra shows very low regeneration. In three different habitats obtained in a gradient of time since fire in Catalonia (NE Spain), we have evaluated the effects of seed predators (ants, rodents and birds) on post-dispersal seed removal and early seedling establishment of P. nigra by using selective exclosures limiting their access to seeds. Ants were the most efficient seed predator group, followed by rodents and birds. The contribution of each group to overall predation showed large seasonal variations. The first seeds dispersed in winter were mainly predated by rodents, which also registered their highest abundance in this season of the year. In spring, at the end of the natural dissemination period of P. nigra seeds, ants became the major predators, this fact coinciding with their increased abundance. Birds showed the lowest predation values. In the seedling establishment experiment, only in the exclusion treatment of the three predator groups was there initial establishment in all habitats, especially in the recently burned area, where there was seedling establishment in all exclusion treatments. The post-dispersal seed predation by different animal groups and low seedling emergence in the different habitats obtained in this study, together with the low seed availability of P. nigra seeds in burned areas, do not predict a favourable outlook for the natural post-fire recolonization of this species, which might even affect its overall distribution area in the region.     

Factors affecting seed germination and seedling establishment of a long-lived desert shrub (<Emphasis Type="Italic">Coleogyne ramosissima</Emphasis>: Rosaceae)

Long-lived desert shrubs exhibit infrequent, episodic recruitment from seed. In spite of this long time scale, selection on life history attributes that affect seedling recruitment should be strong. We studied factors affecting germination phenology and seedling establishment for Coleogyne ramosissima, a dominant shrub species in the ecotone between warm and cold deserts in western North America. We also examined ecotypic differentiation in establishment strategy in response to selection regimes in two contrasting habitats. We followed patterns of dormancy loss, germination, emergence, and survival in reciprocal field experiments at warm winter Mojave Desert and cold winter Colorado Plateau study sites. Seed germination took place in late winter, under winter rain conditions at the warm desert site and under snow at the cold desert site. Distinctive germination phenologies for the two seed populations at contrasting field sites followed predictions based on laboratory germination experiments. There was no seed bank carryover across years. Seedling survival at the end of three growing seasons was remarkably high (mean survival 54%). Most seedling mortality was due to sprout predation by rodents early the first spring in unprotected caches. Emergence and establishment at each site were significantly higher for seeds from the local population, supporting the idea of ecotypic differentiation in establishment strategy. Establishment success was an order of magnitude greater overall at the Colorado Plateau site, which represents the leading edge of an upward elevational shift in distribution for this species under the current climatic regime. The Mojave Desert site is on the trailing edge of this shift, and recruitment there is apparently a much less frequent occurrence.     

Tracking Seed Fates of Tropical Tree Species: Evidence for Seed Caching in a Tropical Forest in North-East India

Rodents affect the post-dispersal fate of seeds by acting either as on-site seed predators or as secondary dispersers when they scatter-hoard seeds. The tropical forests of north-east India harbour a high diversity of little-studied terrestrial murid and hystricid rodents. We examined the role played by these rodents in determining the seed fates of tropical evergreen tree species in a forest site in north-east India. We selected ten tree species (3 mammal-dispersed and 7 bird-dispersed) that varied in seed size and followed the fates of 10,777 tagged seeds. We used camera traps to determine the identity of rodent visitors, visitation rates and their seed-handling behavior. Seeds of all tree species were handled by at least one rodent taxon. Overall rates of seed removal (44.5%) were much higher than direct on-site seed predation (9.9%), but seed-handling behavior differed between the terrestrial rodent groups: two species of murid rodents removed and cached seeds, and two species of porcupines were on-site seed predators. In addition, a true cricket, Brachytrupes sp., cached seeds of three species underground. We found 309 caches formed by the rodents and the cricket; most were single-seeded (79%) and seeds were moved up to 19 m. Over 40% of seeds were re-cached from primary cache locations, while about 12% germinated in the primary caches. Seed removal rates varied widely amongst tree species, from 3% in Beilschmiedia assamica to 97% in Actinodaphne obovata. Seed predation was observed in nine species. Chisocheton cumingianus (57%) and Prunus ceylanica (25%) had moderate levels of seed predation while the remaining species had less than 10% seed predation. We hypothesized that seed traits that provide information on resource quantity would influence rodent choice of a seed, while traits that determine resource accessibility would influence whether seeds are removed or eaten. Removal rates significantly decreased (p < 0.001) while predation rates increased (p = 0.06) with seed size. Removal rates were significantly lower for soft seeds (p = 0.002), whereas predation rates were significantly higher on soft seeds (p = 0.01). Our results show that murid rodents play a very important role in affecting the seed fates of tropical trees in the Eastern Himalayas. We also found that the different rodent groups differed in their seed handling behavior and responses to changes in seed characteristics.     

Consequences of habitat disturbance on seed fate of a Brazilian tropical dry forest tree Cavanillesia arborea (Malvaceae)

We compared seed fate (survival, mortality by rodent predators, desiccation and other causes) of the tree Cavanillesia arborea (Malvaceae) in preserved old‐growth tropical dry forests and in nearby abandoned pasturelands in Brazil. For this purpose, we performed an experiment where 15 seeds were placed in the surroundings of 15 parental individuals in each habitat. These 450 seeds were monitored over four months. At the end of the experiment, seed predation by rodents was higher in forest (56.5%) than in pasture (8.9%) areas, but seed desiccation showed the opposite pattern (8.9% vs. 80.4%). Mortality by desiccation was also faster in pasture than in forest areas, probably reducing their attractiveness to predators in these areas. None of the seeds placed in the pasture survived, whereas 26.2% of seeds became seedlings in forest areas. The absence of seedling recruitment of Cavanillesia arborea in pastures is likely a consequence of their incapacity to tolerate the harsh abiotic conditions in this habitat. Thus, forest conversion imposes a strong limitation to the long‐term population viability of this species. However, restoration and natural regeneration of abandoned pastures can recreate the forest structure and microclimatic conditions favourable to seed germination and seedling establishment. Remnant individuals of Cavanillesia arborea in agricultural landscapes may have a negligible contribution to current recruitment, but they can attract potential dispersers of pioneer species, with positive feedbacks to future recruitment during secondary succession.     

SEED DENSITY, COVER, PREDATION AND THE DISTRIBUTION OF BIRDS IN A BEECH WOOD IN SOUTHERN SWEDEN

The distribution of six species of birds within a beech wood was analysed in two winters when beech seeds were abundant. All species were concentrated in patches where seed density was high in one winter, but only two showed a similar distribution in the second winter. The distribution of the other four species, three tits Parus spp. and the Nuthatch Sitta europaea, was positively correlated with low cover. Cover was inversely correlated with seed density. Bird predators were recorded many times during the second winter but not at all during the first winter. It is suggested that predator avoidance was more important than food density for some species in the second winter. The relationships between bird density and both food and cover were sigmoid in shape. The proportion of the seed crop consumed by the birds was estimated at about 4.5%, which was probably more than the amount consumed by mammals. The Woodpigeon Columba palumbus (which took 44% of the total seeds consumed by birds) and the Brambling Fringilla montifringilla (33%) were the most important seed consumers among the birds.     

啮齿动物捕食和搬运蒙古栎种子对种群更新的影响

2010-2011年,在东北林业大学城市林业示范基地研究了啮齿动物对蒙古栎种子的捕食和搬运。结果表明,蒙古栎种子捕食率在年际间无显著差异,但2010年的蒙古栎种子搬运率显著高于2011年,存留率显著低于2011年。啮齿动物在胡桃楸林、樟子松林、水曲柳林和白桦林内对蒙古栎种子的总捕食率和总搬运率分别达到(5.7±13.5)%和(27.1±37.1)%。蒙古栎种子在樟子松林内的捕食率和搬运率均最高,胡桃楸林内蒙古栎种子存留率最高;2010年蒙古栎种子在与蒙古栎林边缘距离0,10,20,30,40,50m组间的捕食率无显著差异,搬运率和存留率有显著差异,但2011年蒙古栎种子在不同距离组间的捕食率呈显著差异,搬运率和存留率无显著差异。与蒙古栎林边缘距离20m处蒙古栎种子的捕食率最高。2010年和2011年,分别有(37.8±49.7)%和(27.7±49.8)%的蒙古栎种子被啮齿动物利用,这表明啮齿动物是林业示范基地内蒙古栎地表种子的主要捕食者。冬季食物匮乏秋季贮藏种子是造成啮齿动物对蒙古栎种子有较大捕食、扩散压力的主要原因。因此,啮齿动物对蒙古栎种子的捕食和搬运影响了蒙古栎林的种群更新。     

啮齿类取食的物种偏好与时空格局

通过强烈消耗土壤种子库,动物取食种子对植物种群更新和群落动态产生深远的影响。一般认为种子被食概率的空间格局取决于种子密度和离母树的距离,而环境（如地形）异质性的影响则一直没有得到足够的关注,与此相关的机制及其影响程度亦不清楚。研究设计在野外埋放种子以模拟种子扩散后的情形,监测啮齿类对种子的取食,以检验种子取食受埋藏生境、时间及动物对种子种类的偏好等因素的影响。结果表明,经过1a实验,8种落叶阔叶树种子的累计取食概率为0～48．25％,平均值为20％;山顶部位的取食概率大致是其它部位概率的3倍;埋放在凋落物层中的种子被食概率大约为埋放在土壤层中概率的2倍。利用logistic回归模型进行统计分析表明,种子被食概率变化的45％可以被上述因素解释。其中,物种偏好是影响种子被取食概率的首要因素,其后依次是地形、埋藏时间和深度。啮齿类明显喜好较大的种子;其取食行为在山脊部位明显较其它部位更频繁和剧烈;对埋藏种子的取食从3月份开始加剧,到7月份以后平息下来。种子埋放深度对啮齿类的取食概率有显著影响。     

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.