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。     

啮齿动物对印度栲种子扩散的效率

从2007 年11 月到2009 年11 月,在西双版纳热带雨林中选取三棵母树,在每棵树下每年释放标记印度栲种子200 粒(三年共计1 800 粒),并追踪其命运。通过调查啮齿动物搬运和分散贮藏印度栲种子的比例,以及
查贮藏种子的微生境、贮藏点大小和扩散距离,分析贮藏种子的存活情况,进而评估啮齿动物对印度栲种子扩散的效率。结果表明,啮齿动物搬运了69.3% 的印度栲种子,被搬运的种子中18% 被分散贮藏。所有被分散贮藏的种子均被埋于落叶下或埋于土壤表层,并且大部分贮藏点仅含一粒种子。种子的扩散距离从0.5 m 到43.8 m,平均距离为7.1 m,扩散距离在年间没有显著差异。2007 年(种子密度低,啮齿动物密度高) 没有种能最终存活到实验结束,而2008 和2009 年(种子密度高,啮齿动物密度低)分别有0.3% 和1.5% 的种子存活。研究表明,啮齿动物是印度栲有效的种子扩散者,但其扩散效率很大程度上取决于森林中种子的密度和啮齿动物的丰富度。     

动物与植物种子更新的关系Ⅱ．动物对种子的捕食、扩散、贮藏及与幼苗建成的关系

植物的繁殖体总是面临来自各类生物（如昆虫、脊椎动物、真菌）的捕食风险。因动物捕食引起的种子死亡率影响植物的适合度、种群动态、群落结构和物种多样性的保持。种子被捕食的时间和强度成为植物生活史中发芽速度、地下种子库等特征的主要选择压力,而种子大小、生境类型等因素也影响动物对植物种子的捕食。捕食者饱和现象被认为是植物和种子捕食者之间的高度协同进化作用的结果,是限制动物破坏种子、提高被扩散种子存活率的一种选择压力。大部分群落中的大多数植物种子被动物扩散。种子扩散影响种子密度、种子被捕食率、病原体攻击率、种子与母树的距离、种子到达的生境类型以及建成的植株将与何种植物竞争,从而影响种子和幼苗的存活,最终影响母树及后代植物的适合度。种子被动物扩散后的分布一般遵循负指数分布曲线,大多数种子并没有扩散到离母树很远的地方。捕食风险、生境类型、植被盖度均影响动物对种子的扩散。植物结实的季节和果实损耗的过程也体现了其对扩散机会的适应。许多动物有贮藏植物种子的行为。动物贮藏植物繁殖体的行为,一方面调节食物的时空分布,提高了贮食动物在食物缺乏期的生存概率;另一方面也为种子萌发提供了适宜条件,促进了植物的扩散。于是,植物与贮食动物形成了一种协同进化关系,这种关系可能是自然界互惠关系（mutualism)的一种。影响幼苗存活和建成的因子包括种子贮蒇点的微生境、湿度、坡向、坡度、林冠盖度等。许多果食性动物吃掉果肉后,再将完好的种子反刍或排泄出来。种子经动物消化道处理后,发芽率常有所提高。     

神农架巴山冷杉种子雨的时空格局

巴山冷杉(Abies fargesii)是我国特有树种, 其分布区以秦巴山地为中心。在湖北神农架自然保护区选择不同的巴山冷杉群落设置样地, 通过布设种子收集器结合室内实验分析, 对巴山冷杉的种子雨进行研究。各样地巴山冷杉种子雨开始和持续时间大致相同, 始于10月初, 持续超过2个月, 但是不同的巴山冷杉种群表现出不同的种子雨时空格局。巴山冷杉－箭竹(Fargesia nitida) 群落种子雨的平均强度为167.93±111.14粒/m2, 有活力种子比例占22.31%, 落种高峰期集中于10月27日到11月2日间, 种子雨呈现聚集分布。巴山冷杉－茵芋(Skimmia reevesiana)群落的种子雨强度在3个样地中最小, 只有16.41±14.41粒/m2, 有活力种子仅占3.05%。对巴山冷杉－陕甘花楸(Sorbus koehneana)群落, 沿等高线方向和垂直等高线方向布置收集器带, 收集器自群落内中心母树延伸至林冠范围之外(距中心母树的距离>18 m)。收集结果表明种子雨落种高峰为10月15–21日, 种子雨主要集中在林冠范围内, 林冠内种子雨数量占到了收集总数的87.95%。方差分析结果显示各方向的种子雨数量不存在显著差异, 按照远离中心母树的距离进行分段比较, 0–6 m区段内的收集数量和其他各区段间(6–12 m, 12–18 m, >18 m)存在显著差异, 扩散的种子数量与离开中心母树的距离间的关系近似正态分布。研究结果说明巴山冷杉种子雨的强度大但种子质量不高, 而且种子雨的扩散距离有限。这些特征将影响到巴山冷杉种群的分布格局和种群更新。     

香果树(Emmenopterys henryi)种群种子雨、种子库及 实生苗数量的海拔梯度变化

香果树为我国Ⅱ级野生保护植物,由于原生境中香果树种群的数量迅速减少,目前已濒临灭绝,种群亟待恢复。研究了武夷山不同海拔的香果树种群种子雨、种子库及其幼苗存活现状,目的是阐明不同海拔地区的香果树种群自然更新的动态过程,寻找其更新脆弱的环节及其影响因素,为香果树自然种群的恢复提供理论依据。以分布于不同海拔高度（819、980、1 140及1 301 m）的香果树母树为中心,在其周围（东、南、西、北四个方位）布设种子雨收集框和土壤种子库样方,连续观测种子雨和种子库的动态,并对不同地表覆盖物中种子萌发及幼苗数量进行观测和记录。结果表明,武夷山香果树种群种子雨持续时间近2个月,尤其是11月底至12月中旬为种子雨高峰期;随着海拔的上升,香果树种子总密度以及虫蛀种子密度均显著降低,但千粒重以及饱满种子密度则有所增加;香果树土壤种子库为瞬时种子库,其水平分布以母树南侧和西侧种子密度较大,母树北侧种子密度最小。香果树种子的垂直分布主要集中于枯落物和苔藓层;3月时香果树土壤种子库中存留的饱满种子仅占种子散布后1.80%,而1年实生苗存活率为6.18%,土壤种子库的损耗是其种群自然更新困难的主要原因;不同地表覆盖物对香果树幼苗的存活产生显著影响,枯落物及苔藓层的幼苗死亡率显著高于土壤表面。建议清理林内枯落物及苔藓,增加林内裸露土壤的面积,减小地上草本的盖度,以减少种子霉烂、虫蛀的几率,增加香果树实生苗胚根与土壤接触的几率以及改善光照条件,促进香果树自然更新。     

内生真菌侵染对黑麦草种子萌发、幼苗生长及渗透胁迫抗性的影响

 以含有内生真菌的黑麦草(Lolium perenne L.)种子为材料,采用4 ℃冰箱内和20 ℃培养箱内保存18个月的方式分别构建内生真菌侵染（EI）和内生真菌非侵染（EF）的黑麦草种群,通过比较EI和EF种群在正常条件下（对照）和渗透胁迫条件下种子发芽、幼苗生长等方面的差异,探讨内生真菌对其宿主植物的直接和间接影响。结果表明：在对照和胁迫条件下,EI种子的发芽势及发芽率均明显高于EF种子,而在重度胁迫下EI植株的叶延伸速率、根系总长度高于EF植株。内生真菌对宿主植物分蘖数和生物量的变化没有促进作用,但     

内生真菌侵染对黑麦草种子萌发、幼苗生长及渗透胁迫抗性的影响

以含有内生真菌的黑麦草(Lolium perenne L.)种子为材料,采用4 ℃冰箱内和20 ℃培养箱内保存18个月的方式分别构建内生真菌侵染（EI）和内生真菌非侵染（EF）的黑麦草种群,通过比较EI和EF种群在正常条件下（对照）和渗透胁迫条件下种子发芽、幼苗生长等方面的差异,探讨内生真菌对其宿主植物的直接和间接影响。结果表明：在对照和胁迫条件下,EI种子的发芽势及发芽率均明显高于EF种子,而在重度胁迫下EI植株的叶延伸速率、根系总长度高于EF植株。内生真菌对宿主植物分蘖数和生物量的变化没有促进作用,但     

鼠类对辽东栎橡子的搬运

在北京东灵山地区,在灌丛、森林2 种生境类型中共设计了5 个处理,以检验种子质量、种子可视度、生境类型如何影响鼠类对辽东栎（Quercus liaotungensis）橡子的搬运。每个处理包括200 枚种子。每日检查种子状态,实验持续16 d。鼠类在几天之内将大多数种子搬运走。完好橡子比虫蛀橡子消失得快。橡子在森林生境中明显比在灌丛生境中消失的快。鼠类对放置在灌丛下方和灌丛之间的橡子的搬运速率差别不大。放置在落叶下方的橡子消失速率明显慢于地表的橡子。本研究利用半存活时间测量种子的搬运率。各处理的半存活时间范围是0. 6 ～ 8. 6 d。各处理的橡子搬运率均较高,这说明鼠类有能力在种子下落到冬季来临的2 个月时间内,搬运绝大多数的橡子。实验说明,鼠类是辽东栎橡子重要的捕食者和扩散者。     

猴耳环天然更新特性

为研究药用植物猴耳环（Archidendron clypearia）的天然更新特性,2015年7月和2016年7月对广东省广州市、惠州市和深圳市主要森林公园、自然保护区和生态公益林内天然分布的42株母树进行了天然更新状况调查,并采用熵值法对更新效果进行评价。结果表明：（1）母树的上下和左右冠幅十分接近,但97.6%的母树存在偏冠现象,尤以向坡下偏冠更为明显,上坡方向树冠宽度约为下坡的1/2;偏冠是影响种子散布范围的主要原因之一;（2）与母树的距离与不同方向对种子雨量影响显著,但两者对苗木的数量和径高生长没有显著影响;（3）林冠下≤ 1 a的苗木占总数的87.8%,且林下无5年及以上的幼树,预示猴耳环依靠天然更新维持种群增长困难较大;（4）用信息熵理论导出的更新指数来评价与母树不同距离的更新效果显示,与母树距离越远更新效果越差;距母树0~3 m范围内更新效果最好;（5）猴耳环具有诸多"极小种群野生植物"的特征,亟需对其资源分布现状及生物生态学特性开展进一步的深入调查和研究。     

黄檗的更新特点及食果实鸟类对其种子的传播

在帽儿山实验林场和哈尔滨实验林场 ,黄檗母树下没有幼苗分布 ,不能进行天然更新 ,需要靠食果实鸟类等将果实和种子传播至远离母树的其他林下。捕食黄檗果实的食果实鸟类有 9种。其中 6种是食果肉鸟类 ,吞入果实后 ,消化果肉 ,而种子完整地随粪便排出而得以传播。其余 3种是食种子鸟类 ,没有传播作用。果实在鸟类消化道内的滞留时间达 2 0～ 30min ,具有很长的潜在传播距离。将鸟类消化后的种子与完整果实和人为去果肉种子进行萌发对比实验 ,消化后种子的累计萌发率与其余二者之间均没有显著性差异 ,说明食果实鸟类的消化 (道 )过程对种子萌发没有明显影响 ,同时证明果肉对种子萌发率没有显著影响 ,果肉中不含萌发抑制物质。黄檗提供多种鸟类以食物 ,而鸟类也同时以多种肉质果植物为食物。因此食果实鸟类和肉质果植物 (包括黄檗 )之间形成了松散的互利共生关系     

Effect of distance,aggregation, and habitat on levels of seed predation for two mammal — dispersed neotropical rain forest tree species

The effect of seed aggregation and distance from conspecific trees on seed predation was experimentally examined for two neotropical tree species, Macoubea guianensis (Apocynaceae) and Pouteria sp. (Sapotaceae) in a lowland tropical rain forest in northeastern Peru. Results of these experiments are discussed in the context of the Janzen-Connell model (Janzen 1970; Connell 1971), which predicts decreased seed survival near parent trees due to either density-or distance-responsive mortality, and Howe's model (Howe 1989) which predicts that trees with seeds dispersed in clumps (aggregated) will not suffer density-dependent predation, and will have higher survival of seeds near the parent tree than other trees. We also examined whether predation on seeds of these species was affected by seed placement in or near 30-m-wide strips regenerating after clear-cutting. Both species appeared to be mammal-dispersed but differed in how frugivores handled seeds, seed size, overall fruit crop size, and gemination time. Neither of the two species studied appeared to suffer seed predation in a manner predicted by the Janzen-Connell model, and patterns of seed predation for only one of the species was similar to predictions of Howe's model. For neither species did seed predation along the edge of, or in the center of, regenerating clear cuts differ from predation 15 m into the primary forest. For Pouteria, seed predation in and near regnerating strips was significantly greater than around forest trees, but the opposite pattern held for Macoubea. Overall, seed predation was much greater on Macoubea. The difference in seed predation for these two species was most likely a result of differences in the types of seed predators that attacked these two species.     

Tropical tree species diversity: a test of the Janzen-Connell model

To test the premises and predictions of the Janzen-Connell model (Janzen's spacing mechanism), seeds of the rainforest canopy tree, Brosimum alicastrum, were placed at different distances from the parent tree and their removal observed over 3 weeks. The number and density of naturally occurring seeds at different distances from the parent tree were also estimated. Predation was not greater near the parent tree, except on the very small spatial scale: the proportion of experimental seeds removed was greater 1 m from the trunk than it was 5–25 m from the trunk. Predation was negatively correlated with seed density, not positively as the Janzen-Connell model assumes-presumably due to predator satiation. The density of seeds after predation peaked 5 m from the tree trunk, but this is well within the crown radius of the parent tree. There is a peak in the number of potential recruits at a distance of 10 m from the parent tree, due to the peaked initial distribution of seeds. This peak is caused by the interaction between the seed density curve and the increasing area of an annulus around the parent tree at increasing distances, not by the product of the density curve and the predation curve. However, it is important to realize that it is not the presence of a peak in recruitment away from the parent that is essential to maintaining tropical tree species diversity, but frequency-dependent recruitment induced by poor recruitment near conspecifics. Predator satiation seems to be an important factor in the survival of B. alicastrum seeds, possibly at several spatial scales. The number of seeds produced by the tree is negatively correlated with the loss to predators, and trees that have a fruiting conspecific nearby also suffer lower levels of predation. Seed predation increases as one moves from the forest edge into the interior, creating an edge effect that may have long-term effects on the forest composition and tree species diversity. More studies are needed, for other species, other localities, and larger spatial and temporal scales, on both the Janzen-Connell mechanism and this edge effect.     

Seed Dispersal of a High Quality Fruit by Specialized Frugivores: High Quality Dispersal?1

Dispersal quality, as estimated by the cumulative effects of dispersal, germination, seed predation, and seedling survival, was examined for Beilschmiedia pendula (Lauraceae) in Monteverde, Costa Rica. I determined the pattern of dispersal by finding seeds deposited by birds, protected the seeds from seed predators with cages to assess germination and seedling survival, and examined seed predation rates with marked seeds. Seed predation, germination, and seedling survival were compared between seeds naturally dispersed by birds and seeds placed at randomly located sites. Approximately 70 percent of seeds dispersed by birds (N= 244) were deposited <10 m from crown edges of fruiting B. pendula trees, although some seeds were dispersed at least 70 m away. Larger seeds were more likely to be dispersed under or close to the parent trees, and larger seeds produced larger seedlings. Seed size was not correlated directly with seedling survival, but larger seedlings at three months were most likely to survive one year. Seed predation by mammals and insects and seedling mortality due to fungal pathogens were concentrated beneath the crowns of parent trees. Seedlings and saplings were more abundant beneath fruiting B. pendula trees, but individuals farther away were taller on average. Thus, dispersal is beneficial for B. pendula, but such benefits appear most pronounced at a small spatial scale; seeds dispersed >30 m from the crown edges actually had a lower probability of survival than those dispersed 10–20 m. Only 10 percent of B. pendula. seeds received high‐quality dispersal in terms of landing in the zone with the highest per seed probability of seedling survival 10–20 m from parental crowns.     

Impacts of logging on density-dependent predation of dipterocarp seeds in a South East Asian rainforest

Much of the forest remaining in South East Asia has been selectively logged. The processes promoting species coexistence may be the key to the recovery and maintenance of diversity in these forests. One such process is the Janzen-Connell mechanism, where specialized natural enemies such as seed predators maintain diversity by inhibiting regeneration near conspecifics. In Neotropical forests, anthropogenic disturbance can disrupt the Janzen-Connell mechanism, but similar data are unavailable for South East Asia. We investigated the effects of conspecific density (two spatial scales) and distance from fruiting trees on seed and seedling survival of the canopy tree Parashorea malaanonan in unlogged and logged forests in Sabah, Malaysia. The production of mature seeds was higher in unlogged forest, perhaps because high adult densities facilitate pollination or satiate pre-dispersal predators. In both forest types, post-dispersal survival was reduced by small-scale (1 m(2)) conspecific density, but not by proximity to the nearest fruiting tree. Large-scale conspecific density (seeds per fruiting tree) reduced predation, probably by satiating predators. Higher seed production in unlogged forest, in combination with slightly higher survival, meant that recruitment was almost entirely limited to unlogged forest. Thus, while logging might not affect the Janzen-Connell mechanism at this site, it may influence the recruitment of particular species.     

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     

Postdispersal Seed Fate of Tropical Montane Trees in an Agricultural Landscape, Southern Costa Rica

Variation in postdispersal seed fate is an important factor driving patterns of forest regeneration. Because most previous studies have not tracked final seed fate and have commonly equated seed removal with predation without considering the possibility of secondary dispersal, little is known about individual seed mortality factors in successional and degraded habitats. This study tracked the postdispersal fate of large-seeded tropical montane trees in abandoned pasture, young secondary forests, and small forest fragments during two consecutive years in an agricultural landscape in southern Costa Rica. The incidence of secondary dispersal by animals, scatterhoarding in particular, and the effects of seed burial on germination were measured. Overall, seeds survived through germination more often in secondary forests with high levels of mortality occurring in abandoned pastures and forest fragments. The majority of seed mortality was caused by rodent predation in forest fragments, insects and fungal pathogens in secondary forests, and a combination of desiccation, insects, and fungal pathogens in pastures. Seeds were frequently secondarily dispersed in larger forest fragments, whereas they were only rarely moved in pastures and secondary forests. Burial tended to improve germination in pastures and was important for an often scatterhoarded species, Otoba novogranatensis, in all habitats. The results of this study suggest that: (1) seed mortality factors differ in response to the type of habitat degradation; (2) large-seeded species have high potential for survival when dispersed to young secondary forests; and (3) seed removal is not a reliable proxy for seed predation, particularly in less degraded forest fragments.     

Frugivory and Seed Fate in Bursera inversa (Burseraceae) at Tinigua Park,Colombia: Implications for Primate Conservation1

The large ateline primates are efficient seed dispersers in Neotropical forests and hunting is driving their populations to extinction, but we do not know whether other frugivores could substitute primates in their ecological role as seed dispersers. In this study we test this possibility using a potential keystone species (Bursera inversa) at Tinigua Park, Colombia. This plant species allows us to compare seed removal rates between emergent, isolated trees, without primate visitors and trees with connected crowns. We used traps to estimate fruit production and seed removal rates in six different trees, and fruiting trees were observed during 2 yr to quantify the number of seeds manipulated by different animal species. We carried out seed predation experiments to test if seed removal by predators was affected by distance or density effects. We found that the most productive trees attracted more visiting species and seed removal rates differed among trees, the lowest corresponding to trees without primate access. Seed removal rates from the ground by predators were not higher below parental trees than away from them, but the distribution of saplings in the forest suggests that seed dispersal is advantageous. Although it is likely that the effect of primate extinctions will vary depending on tree species traits, conserving the populations of primate seed dispersers is critical to maintain the ecological processes in this forest.     

Contagious seed dispersal beneath heterospecific fruiting trees and its consequences

An hypothesized advantage of seed dispersal is avoidance of high per capita mortality (i.e. density-dependent mortality) associated with dense populations of seeds and seedlings beneath parent trees. This hypothesis, inherent in nearly all seed dispersal studies, assumes that density effects are species-specific. Yet because many tree species exhibit overlapping fruiting phenologies and share dispersers, seeds may be deposited preferentially under synchronously fruiting heterospecific trees, another location where they may be particularly vulnerable to mortality, in this case by generalist seed predators. We demonstrate that frugivores disperse higher densities of Cornus florida seeds under fruiting (female) Ilex opaca trees than under non-fruiting (male) Ilex trees in temperate hardwood forest settings in South Carolina, USA. To determine if density of Cornus and/or Ilex seeds influences survivorship of dispersed Cornus seeds, we followed the fates of experimentally dispersed Cornus seeds in neighborhoods of differing, manipulated background densities of Cornus and Ilex seeds. We found that the probability of predation on dispersed Cornus seeds was a function of both Cornus and Ilex background seed densities. Higher densities of Ilex seeds negatively affected Cornus seed survivorship, and this was particularly evident as background densities of dispersed Cornus seeds increased. These results illustrate the importance of viewing seed dispersal and predation in a community context, as the pattern and intensity of density-dependent mortality may not be solely a function of conspecific densities.     

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.