蒙古栎种子相对丰富度对小兴安岭5种木本植物种子扩散的影响

2009和2010年秋季,在黑龙江省伊春市带岭区东方红林场研究了蒙古栎结实量及啮齿动物密度变化对胡桃楸、红松、蒙古栎、毛榛和平榛5种木本植物种子扩散的影响.结果表明:在小兴安岭林区,啮齿动物种群数量具有年际变化,2009年小型啮齿动物的总捕获率(31.0％)显著高于2010年(16.7％);2009年蒙古栎种子雨密度(6.2±2.1粒·m-2)和种子相对丰富度(20.O)均显著低于2010年(26.7±10.2粒·m-2和160.0).2009年,除胡桃楸种子外,其他种子全部被扩散或者被原地取食,其中蒙古栎种子被分散埋藏的比例最高,且种子平均扩散距离最大;2010年,胡桃楸种子被分散埋藏的比例最高,且种子平均扩散距离最大.研究区蒙古栎种子相对丰富度是影响其他木本植物种子扩散的重要因素.     

不同类型城市森林对土壤肥力的影响

以东北林业大学城市林业示范研究基地中9种人工林(兴安落叶松林、樟子松林、黑皮油松林、黄波罗林、胡桃楸林、水曲柳林、白桦林、蒙古栎林、针阔混交林)及附近的农耕地和撂荒地土壤为研究对象,通过对林地土壤不同层次pH值、有机质及主要养分含量的测定分析,研究了不同林型对土壤肥力的影响.结果表明：阔叶林(除蒙古栎林)的土壤趋于中性,针阔混交林、兴安落叶松林、樟子松林和黑皮油松林的土壤呈微酸性,蒙古栎林的土壤呈酸性;随土壤深度增加,土壤有机质、水解氮、速效钾、有效磷、全氮、全磷含量均呈下降趋势.不同林地土壤同一层次化学指标整体差异显著(P＜0.05).土壤肥力优劣为：水曲柳林>黄波罗林>针阔混交林>胡桃楸林>白桦林>撂荒地>农耕地>樟子松林>兴安落叶松林>蒙古栎林>黑皮油松林,说明阔叶林(除蒙古栎林)和针阔混交林中的土壤肥力增加,而针叶林的土壤肥力趋于下降.     

啮齿动物对六盘山区辽东栎、野李和华山松种子的取食和搬运

为了阐明不同种子投放方式和覆盖处理对啮齿动物取食和搬运/贮藏种子行为的影响,2010年春季,我们在宁夏六盘山区的辽东栎幼林,用塑料标签标记法研究了不同种子投放方式(单独投放、两两组合投放和混合投放)和覆盖处理(对照、凋落物覆盖和土壤覆盖)下啮齿动物对辽东栎、野李和华山松种子的取食和搬运特点。结果表明:(1)不同物种、不同种子投放方式和覆盖处理对种子的就地取食率、搬运后的取食率和贮藏率均具有显著影响,种子较大或具有厚而坚硬的种皮(内果皮)、组合和混合投放及覆盖处理均不同程度地促进啮齿动物对种子的搬运;(2)3种植物的种子被搬运后取食和贮藏的平均距离范围均在5.5m以内,而且种子被搬运后贮藏的距离均大于搬运后取食的距离,其中华山松种子被搬运后取食的距离显著大于辽东栎和野李种子,但搬运后的贮藏距离显著小于后两者。(3)种子被搬运后单个种子的取食点和贮藏点占多数,其他大小的取食点和贮藏点较少,种子被搬运后以土壤埋藏为主要的贮藏方式。结果表明,种子的种皮特征以及种子在群落中的分布方式和存在状态,可能通过延长啮齿动物处理的时间而增大啮齿动物的被捕食风险,使动物改变对种子的取食和搬运行为,进而决定了种子的命运。     

啮齿动物捕食压力下生境类型和覆盖处理对辽东栎种子命运的影响

在六盘山区的退化辽东栎灌丛和辽东栎次生林 (以下分别称“灌丛样地“和“次生林样地”),研究了群落生境和清除凋落物、凋落物覆盖及土壤覆盖等处理对啮齿动物取食和搬运/贮藏辽东栎种子的影响。结果表明：(1) 凋落物和土壤覆盖处理在种子释放的前期阶段有利于种子留存,但释放40天后,种子在释放点的最终留存率在灌丛样地显著高于次生林样地 (Z = -2.333, P = 0.020)。灌丛样地的自然状态的种子最终留存率显著高于凋落物覆盖处理 (Z = -0.674,P = 0.05),但其它处理间无显著差异;次生林样地的凋落物覆盖和清除凋落物处理种子的最终留存率为0%,自然状态和土壤覆盖处理均不足1%,各处理间无显著差异。(2) 种子被啮齿动物的就地取食率在不同群落生境间差异显著 (Z = -2.333,P = 0.020);在灌丛样地,凋落物覆盖处理种子被啮齿动物就地取食率最高 (45.56%),而次生林样地的土壤覆盖处理最高 (64.81%),种子被啮齿动物就地取食率在灌丛样地和次生林样地均为清除凋落物处理最低 (分别为23.70%和40.00%);生境类型和覆盖处理对种子搬运后的取食率均无显著影响。(3) 次生林样地种子被啮齿动物搬运后的埋藏率显著高于灌丛样地 (Z = -2.88,P = 0.004);在灌丛样地,土壤覆盖处理种子搬运后的埋藏率最高仅5.56%,而在次生林样地,清除凋落物处理最高达9.26%。(4) 种子被啮齿动物搬运后取食的距离在灌丛样地和次生林样地分别为3.01 m和2.13 m,差异显著 (Z = -2.080,P = 0.038),而种子搬运后的埋藏距离前者小于后者 (分别为1.35 m和2.10 m),两者间差异不显著。     

啮齿动物对秦岭松栎混交林建群种种子扩散格局的影响

森林群落中的啮齿动物对林木种子存在着一定的取食偏好性,这种偏好性会使啮齿动物形成不同的捕食和贮藏策略,从而导致林木种子形成不同的扩散格局。以秦岭中段松栎混交林建群种锐齿槲栎、油松、华山松为研究对象,采用塑料标签标记种子方法,研究啮齿动物对种子传播和扩散格局的影响。结果表明:(1)种子特征是影响啮齿动物对其进行扩散的重要因子,油松种子以其质量小、种皮薄等特点吸引啮齿动物大量捕食,其原地取食率达到83.33%,显著高于华山松和锐齿槲栎种子。(2)锐齿槲栎和华山松种子大部分被搬运一定距离后再被取食和埋藏,其中,扩散距离在1 m以内、1—3 m和5—8 m的锐齿槲栎种子分别占59.12%、18.23%和13.26%,最远扩散距离达12 m处;华山松种子扩散距离主要介于在1—3 m(37.85%)和3—5 m(23.73%),在距离5—8 m(13.56%)、8—10 m(11.86%)和大于10 m(11.30%)区间也有一定的分布,最大扩散距离为12 m。(3)啮齿动物倾向于将锐齿槲栎种子搬运至松林内取食,而将华山松种子搬运至栎林内埋藏,说明不同林分类型及其环境因素对林木种子扩散后的分布格局具有重要影响。(4)3种类型种子被啮齿动物捕食的比例以油松最大(96.90%),锐齿槲栎次之(73.57%),华山松最小(50%);次年调查时,未被捕食的种子大部分已经被取食,仅有极少数锐齿槲栎种子萌发成幼苗(1.67%)。啮齿动物的捕食和贮藏行为对林木种子扩散及其成功更新都至关重要。     

啮齿动物对不同大小和种皮特征种子的取食和搬运

在宁夏六盘山区的辽东栎灌丛,以不同大小的辽东栎种子、野李和华山松种子为材料,研究了种子大小和种皮特征对啮齿动物取食和搬运行为的影响.结果表明:辽东栎小种子和华山松种子的就地取食率均显著高于辽东栎大种子和野李种子;具有厚而坚硬种皮(内果皮)的野李种子被啮齿动物搬运后的取食率和贮藏率均最高.辽东栎大种子被啮齿动物搬运后取食的距离最大(3.10 m),被搬运后贮藏的距离(6.48 m)显著大于其他类型种子.除野李种子外,其他3种类型种子的单个种子取食点都在80％以上,所有种子的单个种子贮藏点都在90％以上,含2个以上种子的取食点和贮藏点较少.较高比例的华山松种子在灌丛基部和洞穴以外的其他环境被取食,其他3种类型种子被啮齿动物搬运后主要集中在灌丛基部和洞穴中取食;种皮(内果皮)厚而坚硬的种子以土壤埋藏方式贮藏的比例偏高.     

哈尔滨市不同类型人工林土壤重金属含量

以东北林业大学城市林业示范研究基地9种人工林(兴安落叶松林、樟子松林、黑皮油松林、黄波罗林、胡桃楸林、水曲柳林、白桦林、蒙古栎林和针阔混交人工林)为对象,分析林地土壤不同层次(0～10 cm和10-30 cm)A8、Cd、Cu、Ni、Pb、Zn 6种重金属含量,并以哈尔滨市土壤背景值为标准,采用综合污染指数法评价各林型土壤重金属污染状况.结果表明:不同类型人工林同一土层重金属含量差异显著;除了As和Ni,同一人工林土壤重金属含量均随土壤深度增加呈下降趋势.各类型人工林同一土层重金属含量以Zn最高(62.29～126.35mg·kg-1),Cd最低(0.06～0.47 mg·kg-1).6种重金属含量由高到低顺序为:Zn＞Pb＞Cu(Ni)＞Ni(Cu)＞As＞Cd,累积程度为:Cd＞Pb＞Zn＞Cu＞Ni＞As.林地土壤Pb、Cd、Cu、Zn间(除水曲柳林)及其与土壤有机质、N、P、K(除水曲柳林和蒙古栎林)显著相关,且以上4种重金属含量高于背景值,为人为输入;Ni和As含量与本地背景值相当,为自然因素,不同林型土壤内梅罗综合污染指数依次为:水曲柳林＞黄波罗林＞针阔混交林＞胡桃楸林＞樟子松林＞黑皮油松林＞兴安落叶松林＞白桦林＞蒙古栎林.     

胡桃楸种子大小对鼠类分散贮藏行为的影响——基于无线电标记技术

种子大小是影响鼠类对种子扩散和分散贮藏的一个重要因素,但有关种子大小对分散贮藏影响的研究结果尚存在一定争议。2012年9月,在小兴安岭南麓带岭区东方红林场利用微型无线电标记技术研究鼠类对大、小胡桃楸种子的多次分散贮藏,进一步探讨种子大小对种子扩散的影响。结果表明:1)胡桃楸种子大小并不影响鼠类对坚果的选择及分散贮藏偏好。胡桃楸大、小种子被扩散的比例分别为100%和95.6%,鼠类对两类种子的埋藏率也十分接近(χ2=0.045,df=1,P=0.831);2)鼠类对胡桃楸大、小种子的扩散次数无显著差异(F=2.710,df=1,P=0.103),大种子被鼠类扩散的平均次数为1.67±0.10,小种子为1.91±0.10。3)胡桃楸大种子的初次、次级扩散距离均显著高于小种子,种子大小也显著影响扩散的净距离和总距离,大种子比小种子扩散的较远。基于无线电标记技术,研究结果准确评价了种子大小对胡桃楸种子扩散距离的影响,进一步支持了大种子扩散较远的假说。     

围栏条件下捕食风险对岩松鼠贮藏核桃种子行为的影响

面对捕食风险,贮藏植物种子的啮齿动物如何通过调整其贮藏行为,权衡捕食风险和贮食效益,需要深入研究。2006 年9 月,在北京东灵山地区,以黄鼬作为捕食风险源,在半自然围栏内比较了有无捕食风险源时,岩松鼠贮藏的核桃种子的状态,以及埋藏种子在实验围栏内的分布、微生境和搬运距离,以探讨捕食风险对岩松鼠贮藏种子行为的影响。结果表明,捕食风险源存在时,搬运和埋藏种子的比例都显著减少,弃置地表种子的比例无显著差异;埋藏于高风险区的种子比例减少,中风险区和低风险区的埋藏种子比例增加,但差异都不显著;位于裸地和草下的埋藏种子比例减少,位于墙基的埋藏种子比例增加,但差异都不显著;此外,埋藏种子的搬运距离也没有显著差异。可见,捕食风险源存在时,岩松鼠显著减少了搬运和埋藏种子的比例,但对埋藏种子时的埋藏区域和微生境选择,以及搬运距离无显著影响。差异不显著可能与围栏效应有关。     

橡胶树在西双版纳引种区的种子捕食与散布

在纯橡胶林、胶-竹复合林、沟谷季节雨林、次生林及毗邻次生林的橡胶林共5种不同生境中释放标记的橡胶树种子,观测种子捕食者以及种子被捕食、埋藏和扩散情况。结果表明:纯橡胶林和沟谷季节雨林内的橡胶树种子大多数都原地存留(80%和98%),而胶-竹复合林内几乎100%的种子均被捕食;次生林内释放的橡胶树种子捕食率也几为100%,而毗邻次生林的橡胶林种子捕食率约50%;卡方检验表明,胶-竹复合林的种子捕食率显著高于沟谷季节雨林和纯橡胶林,次生林内显著高于毗邻的橡胶林(P0.001)。绝大多数橡胶树种子被捕食者原地或搬运后取食,仅有少量种子被埋藏。大多数种子(80%)被搬运距离在10 m以内。红外感应相机拍摄和饲喂实验发现,黄胸鼠(Rattus tanezumi)和红刺鼠(Rattus surifer)为主要的种子捕食者。可见,在西双版纳引种区橡胶树种子主要是被小型鼠类取食,但捕食强度较大,而被埋藏种子比例很低,散布距离较短(10 m)。     

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。     

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.     

Survival and Scatterhoarding of Frugivores-Dispersed Seeds as a Function of Forest Disturbance

The effect of forest disturbance on survival and secondary dispersal of an artificial seed shadow (N= 800) was studied at Brownsberg Natural Park, Suriname, South America. We scattered single seeds of the frugivore‐dispersed tree Virola kwatae (Myristicaceae), simulating loose dispersal by frugivores, in undisturbed and disturbed secondary forest habitats. Seed survival rate aboveground was high (69%) within 2 wk and was negatively correlated with scatterhoarding rate by rodents, the latter being significantly lower in the undisturbed forest (9%) than in the disturbed forest (20%). Postdispersal seed predation by vertebrates was low (3%) and infestation of seeds by invertebrates was almost zero in all instances. Therefore, secondary seed dispersal by rodents in forest is not as critical for recruitment as observed among other bruchid‐infested large‐seeded species. Secondary seed dispersal by rodents may, however, facilitate seedling recruitment whether cached seeds experience greater survival than seeds remaining above ground surface.     

小果野芭蕉种子散布和不同时空尺度上种子被捕食格局

 以分布在云南西双版纳地区的大型先锋草本植物小果野芭蕉(Musa acuminata)为研究材料,研究其种子初次散布过程和不同时空尺度上种子被 捕食格局。小果野芭蕉的成熟果实有75%在夜间被取食和传播,在白天消失的果实则占25%。蝙蝠是其最主要的种子传播者,鸟类在其种子传播 过程中也起到一定的作用。人工摆放种子试验结果显示小果野芭蕉种子的主要转移者是小型啮齿类(鼠类)和蚁类：在开放处理下3 d后转移率为 86%,排除蚁类(鼠类可进入)处理下种子转移率为69%以及排除鼠类(蚂蚁可进入)处理下种子被转移率为56%。季节、地点和生境均显著影响人工 摆放种子被转移强度：雨季显著高于旱季(p<0.001), 野芭蕉生境显著高于与其相连的自然森林和荒地(p<0.001),在人为干扰较少的补蚌自然 保护区显著低于西双版纳热带植物园和新山,而后两者之间并无显著差异(p>0.05)。同时,地点和生境以及季节、地点和生境都有显著的交互 作用。与相邻的森林和荒地相比,野芭蕉群落中种子被鼠类捕食的强度最大且受蚁类二次转移的比例最少,森林和荒地中种子被鼠类捕食的强 度相对较小且蚁类对种子的二次转移比例较高,从而更好地帮助种子逃避鼠类捕食。因此,依赖于食果动物(主要是蝙蝠, 也包括鸟类)的初次 散布是小果野芭蕉种子逃避捕食的关键。     

Spatial and temporal patterns of seed predation on three tree species in an oak-pine forest

We experimentally examined spatial and temporal patterns of seed predation on three tree species in an oak-pine forest in southern Maine. USA, Rodents were the principal seed predators. Rates of seed loss varied with tree species and exposure to three different suites of potential seed predators (all vertebrate seed predators, medium-sized rodents [primarily squirrels] and small rodents [primarily mice and voles]), Acer rubrum seeds were removed more slowly (mean of 17.7% within 2 d across all habitats and treatments in 1991) than seeds of either Quereus rubra or Pinus strobus (e,g., > 99% removed within 2 d across all habitats and treatments for each species in 1991), Levels of final removal (cumulative removal at final census) varied with year but not with microhabitat; i.e., seed predation in four types of forest gap was not significantly different than in intact forest. Both field data and experimental feeding trials with captive Peromyscus leucopus and Clethrionomys gapperi suggest that a significant proportion of while pine seeds is eaten at time of detection, while red oak acorns are more likely to be cached. Captive Peromyscus leueopus cached and ate significantly more red oak acorns than Clethrionomys gapperi. These results suggest that red oak mast may be more important in overwintering success in Feromyscus than in Clethrionomys. Nevertheless, both species did consume red oak acorns, suggesting that in southern Maine these species are not substantially inhibited by high tannin levels in red oak acorns, as has been suggested by researchers elsewhere.     

Temporal consistency in the spatial pattern of seed predation across a forest – old field edge

Seed predation is an important factor in determining the rate of tree establishment in abandoned agricultural land. Edges, through altered habitat use by small mammals, may influence the spatial pattern of forest regeneration in these successional sites. To determine the spatial pattern of seed predation across a forest-old field edge, we used a grid that began 30 m inside the forest and extended 60 m into the old field. Seed stations were placed at regular 10-m intervals and were monitored for removal of Acer rubrum seed for 50 d. This design was repeated over four years (1995–1998). Small mammal trapping was conducted in the final year of the study to determine the spatial pattern of seed predators within the site. Removal rates varied among the four years of the study with years of high and low removal rates. However, the spatial pattern of seed removal rate was similar in all years. Final survival ranged from 0.7–15.5% of seeds, with lower final survival in years with faster rates of seed removal (1996 and 1998). Seed removal rates and rates of discovery were greatest at the forest edge and decreased with distance into the old field. The number of seeds surviving to the end of the experiment also varied across the edge gradient, with highest survival at greater distances into the old field in low predation years. Seed removal rate covaried with spatial pattern of Peromyscus leucopus captures within the site. Seed removal and discovery was also higher under the exotic shrub Rosa multiflora, which may have provided cover for foraging seed predators. These indirect effects of edges on plant communities can potentially alter the rate and spatial pattern of tree invasion into disturbed lands and illustrate the importance of understanding plant-animal interactions in the context of habitat fragmentation.     

啮齿动物对不同林木种子的搬运和取食微生境选择机制

不同植物种子依靠不同的方式实现扩散,啮齿动物对林木种子搬运后在取食点微生境和贮藏方式的选择存在偏好,研究其贮藏行为与微生境的关系是探究幼苗建成的关键。在秦岭中段火地塘林区,采用标签标记法,以锐齿槲栎、华山松和油松种子为材料,探究了小型啮齿动物对松栎混交林建群种种子扩散过程的影响。结果表明:1)油松种子原地取食率显著高于锐齿槲栎和华山松种子,且啮齿动物更倾向于搬运后取食(60%)和埋藏(4.33%)华山松种子,搬运后取食距离也为华山松最大(2.49 m);锐齿槲栎小种子被搬运后埋藏的距离最大(4.92 m)。2)除华山松种子外,其他类型种子被搬运后单个取食的比例均在85%以上;油松种子不存在埋藏点,而其他类型种子90%以上均以单个形式被埋藏。3)大部分种子被啮齿动物搬运后选择在裸地丢弃;锐齿槲栎大种子(87.5%)、小种子(78.57%)和华山松种子(53.33%)较大比例被啮齿动物埋藏在灌丛下方,埋藏在裸地的种子较少。4)大部分种子在灌丛下方被取食,仅华山松种子被啮齿动物搬运到洞穴取食;除油松种子被大量原地取食外,其他类型种子被搬运到取食点的种子比例基本呈现由微生境植被复杂到简单(灌丛—草丛—灌丛边缘—裸地)而逐渐减小的趋势。种子的营养价值及取食和搬运过程中啮齿动物付出的成本是影响种子命运的关键性因子,且啮齿动物对种子埋藏和取食地点的微生境存在较明显的选择性。     

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.     

Tree seed rain and seed removal,but not the seed bank,impede forest recovery in bracken (Pteridium aquilinum (L.) Kuhn)‐dominated clearings in the African highlands

Considerable areas dominated by bracken Pteridium aquilinum (L.) Kuhn occur worldwide and are associated with arrested forest recovery. How forest recovery is impeded in these areas remains poorly understood, especially in the African highlands. The component processes that can lead to recruitment limitation—including low seed arrival, availability and persistence—are important determinants of plant communities and offer a potential explanation for bracken persistence. We investigated key processes that can contribute to recruitment limitation in bracken‐dominated clearings in the Bwindi Impenetrable National Park, Uganda. We examined if differences in seed rain (dispersal limitation), soil seed bank, or seed removal (seed viability and persistence) can, individually or in combination, explain the differences in tree regeneration found between bracken‐dominated areas and the neighboring forest. These processes were assessed along ten 50‐m transects crossing the forest–bracken boundary. When compared to the neighboring forest, bracken clearings had fewer seedlings (bracken 11,557 ± 5482 vs. forest 34,515 ± 6066 seedlings/ha), lower seed rain (949 ± 582 vs. 1605 ± 335 tree seeds m^?2 year^?1), comparable but sparse soil seed bank (304 ± 236 vs. 264 ± 99 viable tree seeds/m²), higher seed removal (70.1% ± 2.4% vs. 40.6% ± 2.4% over a 3‐day interval), and markedly higher rodent densities (25.7 ± 5.4 vs. 5.0 ± 1.6 rodents per 100 trapping sessions). Camera traps revealed that rodents were the dominant animals visiting the seeds in our seed removal study. Synthesis: Recruitment limitation contributes to both the slow recovery of forest in bracken‐dominated areas, and to the composition of the tree species that occur. Low seed arrival and low persistence of unburied seeds can both explain the reduced density of seedlings found in bracken versus neighboring forest. Seed removal, likely due to rodents, in particular appears sufficient to constrain forest recovery and impacts some species more severely than others.