种子的长距离风传播模型研究进展

 植物种子的长距离传播在物种迁移、生物入侵、保护生物学等领域有重要的生态和进化意义。种子传播有很多方式,开阔草原等地区的草本植物和许多热带和温带的树木都是通过风传播种子的。风传播的方式最适合进行种子长距离传播现象的模拟研究。种子的风传播模型是传播生态研究的一个重要领域,尤其是种子的长距离风传播模型,对于外来入侵植物的扩散和破碎化景观中植物种群的基因交流等生态过程研究举足轻重,然而国内鲜见这方面的研究成果。本文综述了种子长距离风传播现象研究的背景和意义,分析了风传播种子模型的基本形式和构成原理,并分别就现象模型和机理模型的相关研究进展进行了总结,同时指出了未来发展的几个重要方向。种子的风传播模型可以分为现象模型和机理模型两类,现象模型按种子传播核心的形式包括短尾模型、偏峰长尾模型和混合传播核心模型,后两者对于长距离传播数据的模拟可以取得很好的效果。机理模型按照模拟机制可分为欧拉对流扩散模型和拉格郎日随机模型两类。本文重点介绍了种子的长距离风传播现象的形成机理和两类机理模型的参数构成和处理方式。适合种子脱落的天气和适合传播的天气的同步性可能是形成种子长距离风传播的一个重要前提,林缘和地表存在的上升气流及大风和暴风中形成的速度梯度都可能对于种子的长距离传播有重要的作用。机理模型的操作因子主要包括生物方面的因子、气象方面的因子和地形方面的因子。同时对目前几个应用比较成功的机理模型进行了简要的介绍和评价,包括倾斜羽毛模型、对流-扩散-下降模型、无掩蔽模型、背景模型、WINDISPER及其改进模型和PAPPUS模型。最后指出,目前在风传播种子的长距离模型研究中,对草本植物种子的传播模拟的投入明显不如树木种子的长距离传播模拟,对于破碎化景观中种子长距离的风传播的研究还存在很大的差距,而对提高机理模型预测能力的高分辨率物理环境数据输入技术的需求则为多学科交叉提供了很好的机会。     

植物种群更新限制——从种子生产到幼树建成

更新限制是指种子由于各种原因,不能够萌发并生长成幼树。它作为解释生物多样性的理论,一直受到国内外群落生态学家关注。从种源限制、传播限制和建成限制3个角度,对更新限制机制研究进展进行了综述。从种源限制而言,时空因素是影响植物种群更新限制的重要因素,因为植物结实量存在明显时空变化,造成植物更新个体出现明显的时空规律。从传播限制而言,传播数量、距离和食果动物行为均限制植物种群更新。数量上,缺乏有限传播者势必减少传播数量,但如果种子拥有较高质量,则能逃脱数量限制;距离上,植物更新个体显示出明显的Janzen-Connell格局,但传播距离趋向稳定,形成植物种群的进化稳定对策;食果动物行为上,不同传播者对更新贡献存在差异,捕食者直接降低更新,融入两类动物行为的模型更能反映食果动物对更新的限制。从建成限制而言,环境因子制约植物生长。小尺度下,微生境的好坏对于植物幼苗建成至关重要;大尺度下,植物提供较好的广告效应则能摆脱生境限制。将传播者行为、捕食者行为与幼苗的空间分布格局、种子传播机理模型等结合,建立植物更新限制机理模型应是更新限制未来的研究热点。选择稀有种和古老种为主题的长期更新限制研究,为种群恢复提供指导,也是未来重要研究方向。     

动物传播者对植物更新的促进与限制

在动物 植物的关系网络中,传播者对植物更新具有促进与限制两方面作用.本文从种群尺度总结了传播者取食、空间利用等行为对植物更新的影响;从群落尺度分析了多种传播者传播有效性对植物更新的促进与限制.传播者对食物的处理方式决定了种子的命运,且具有明显的种间差异;植物在传播者食谱中的地位亦决定其更新的成败,成为动物偏好的食物可助其摆脱森林破碎化等不利事件的影响.动物的空间行为可导致种子命运发生改变.传播者移动距离能否逃逸同种成树对种子的距离限制,影响种子的更新命运;动物偏好的适宜生境与适于植物萌发生境的空间一致性程度影响传播者传播的成效.有效传播者的非冗余性促使种子传播网络更稳定,利于植物更新;无效传播直接限制植物更新,但为其他植物定殖提供了可用空间.今后应将传播者行为融入植物种群更新研究,而从生态系统服务角度揭示传播者在植被恢复的作用应是未来恢复生态学研究的重点.     

基于个体的空间显性模型和遥感技术模拟入侵植物扩张机制

基于个体的空间显性模型和遥感技术,以互花米草为例,模拟了自1997到2010年的种群扩张动态,揭示了土地利用变化与潮间带高程的影响;并通过全局敏感性分析揭示了种子扩散、成体存活率、有性和无性繁殖等种群统计学特征对互花米草种群扩张的相对重要性。研究结果发现:1)有性繁殖与无性繁殖共同决定互花米草种群快速扩张;2)潮间带高程和土地利用变化显著影响模型预测的精度,对互花米草种群扩张有非常重要的影响;3)成体存活率与种子长距离扩散是影响互花米草种群扩张速度最重要的因素;无性繁殖比有性繁殖对种群扩张的影响更大;种子长距离扩散比本地扩散更为重要,同时,小概率的种子长距离扩散事件对种群扩张有非常重要的影响。为了经济有效地控制外来入侵植物的扩张,应该抑制种子的长距离扩散和移除种子长距离扩散形成的位于入侵前沿的小斑块。     

胡杨种子散布的时空分布格局

以额济纳胡杨为研究对象,对种子雨的散布时间、强度、散布距离以及种子雨和空气湿度、风之间的关系进行了研究。胡杨种子雨可以分为初始期、高峰期和消退期3个阶段,大部分的种子集中在高峰期落下。种子的散布主要受湿度和风的影响。湿度对种子雨的强度起主要作用,在一天之中,种子在湿度较低的中午和下午集中散落。应用一元线性回归模型对种子雨强度和相对湿度进行分析后表明二者之间存在显著的负相关关系。对种子的散播距离进行研究后发现,大部分种子落在母树附近,少部分种子能够进行长距离传播。风对种子的传播的方向和距离起决定性的作用,不同方向上的种子传播距离和强度相差很大。在顺风方向上,种子的传播距离最远,所有的长距离传播现象几乎都发生在这一方向上;而在主风向的垂直方向和逆风方向上,种子的散布距离较小,很少有种子能够进行长距离传播。对风的观测表明中午后和下午初的风力较强,而此时种子雨强度又最大,有利风力条件和高种子雨强度出现的同步性可能是促进胡杨种子进行长距离传播最有效的生物控制机制。由于胡杨种子在自然条件下的存活时间非常短暂,所以研究中不同胡杨母树林间种子散播时间的差异可能是胡杨种群内部为适应不同洪水期所表现出的风险分摊机制所造成的。     

食果动物传播植物种子的有效性

传播有效性是评估食果动物对植物种群更新贡献的关键指标,它决定了植物种群的续存,影响着群落生物多样性的维系。本文从种子传播数量、质量角度分析了影响动物传播有效性的因素,并从有效传播动物组合角度分析了其对植物种群更新的影响。传播数量的多少与动物的取食行为有关。以整吞取食的食果动物常被认为是植物的有效传播动物,而植物的果实为动物偏好选择的食物时,势必导致该种植物拥有高的种子传播数量。传播质量与食果动物肠道种子滞留时间以及其后的种子传播距离和排放地相关。体型较大、生境适应能力和迁徙性较强的动物常为植物提供较远的传播距离,而被认为传播质量较高。取食后动物的生境选择决定了种子的排放地,进而影响它们的种子传播质量。如果取食后动物偏好利用植物适宜更新的生境时,动物则拥有较高的传播质量。有效传播动物通常在传播数量和传播质量以互补形成一些组合对植物种群更新起贡献。今后工作应将动物行为野外监测和种子萌发实验相结合、取食后多种行为相结合来分析动物的传播有效性;而以一些国内特有濒危植物为研究对象开展传播有效性研究应是今后恢复生态学研究的重点。     

生境破碎化对食果动物及种子传播的影响

食果动物与依赖其传播种子的植物间在进化过程中形成互惠关系,生境破碎化往往干扰种子传播过程,继而破坏这种关系.生境破碎化通常降低食果动物的多样性,但亦有相反的情况出现.食果动物对生境破碎化的适应能力不同,泛性森林动物和广食性动物具有较强的适应性.生境破碎化对依赖动物传播的植物影响有差异,多数植物受到负面影响,但也有一些植物不受影响,甚至受益.动物在破碎生境中对种子传播的有效性是种子搬运量、传播距离、种子萌发及种群建立等环节的综合效果.破碎化生境中种子的搬运量与动物的觅食行为和食物选择有关;种子传播距离受食物资源可获得性的改变和生境斑块异质性的影响;种子萌发和更新种群建立成功与否决定于是否存在有效的种子传播者.生境破碎化如何影响种子传播以及动植物相互关系,尤其是异质斑块的空间分布如何影响食果动物的传播有效性、破碎化生境下动植物互惠共生关系如何建立,生境破碎化导致的植物入侵对本地植物种子传播的影响是未来需要深入研究的问题.     

生物土壤结皮对种子萌发和幼苗建植的影响研究进展

生物土壤结皮（BSC）在陆地生态系统中具有重要的生态地位,尤其是旱地生态系统中,BSC占据了种子植物之间的广阔地面。因此,BSC的发展必然影响种子植物更新过程与植被空间格局;但其作用方式、影响程度等因相关研究涉及多气候要素、土壤类型、BSC组成物种和种子植物物种的差异及其不同组合,导致目前的研究结论存在广泛争议。研究综合论述了BSC改变地表微形态对种子传播过程的影响;BSC改变土壤特性（物理、化学、生物学）对种子萌发和幼苗存活与建植等关键环节的影响;并结合种子形态特征及种子萌发、幼苗建植的性状等,综合分析了BSC对种子传播、种子萌发与幼苗建植等更新过程的潜在影响机理;探讨了目前研究矛盾性结论产生的原因。总体来说,深入研究并全面揭示BSC对种子植物更新过程的影响,应加强学科交叉,将分子生物学、植物生理学、生物化学等微观研究,与遥感、野外生态因子过程监测、控制实验等宏观、中观研究结合,从机理到过程方面动态研究BSC对种子植物更新过程的影响,并引入水文模型、气候模型、种群动态模型等模型预测方法,研究气候变化、各类干扰频发的情景下,BSC对种子传播、萌发及幼苗建植过程的潜在影响,以期促进对BSC与种子植物间相互关系的研究,加深对干旱脆弱生态系统植被发展规律的认识。     

内蒙古退化草原狼毒种子的种群分布格局与散布机制

狼毒 (Stellera chamaejasme)为瑞香科多年生草本植物 ,是主要的草地有毒植物种类之一。采用邻接格子样方法和分布指数 (DI)以及平均拥挤度 (m* )指标 ,研究了内蒙古西辽河平原北部典型草原区内的狼毒 (Stellera chamaejasme)种子种群分布格局强度、类型与规模 ;根据顺序远离母株的取样调查 ,定量地分析了狼毒种子在 8个方向上的散布格局。结果表明 ,狼毒种子在落种期之前为随机分布 ,而在落种期结束后为聚集分布 ,种子种群斑块大小平均为 0 .0 8m2 。狼毒种子在 8个方向上的散布表现为不同的散布格局模式。在不同方向随着累积面积增加狼毒种子散布数量变化符合 L ogistic曲线增长。各方向种子散布数量与顺风风向频率显著正相关。狼毒种群具有“近母株散布”的种子散布机制 ,基本散布半径为 0～ 5 0 cm。狼毒种群通过种子散布扩展其分布空间的能力是很有限的     

李园桃蚜和草间小黑蛛种群空间格局的地学统计学研究

利用地学统计学的原理和方法分析了李园桃蚜种群和草间小黑蛛种群的空间结构和时间相关性.结果表明,桃蚜种群6月27日和11月22日此两次的半变异函数拟合模型为直线,表明此两次两种空间分布为均匀分布或随机分布,其余9次半变异函数拟合模型均为球形,表明此9次李园桃蚜种群的空间分布是聚集分布,变程RSD(m)为4.173903～34.7837.草间小黑蛛空间格局5月21日、5月31日及10月19日、11月22日4次半变异函数拟合模型为直线,表明此4次种群的空间分布为均匀分布或随机分布,其余8次半变异函数拟合模型均为球形,表明此8次李园草间小黑蛛种群的空间分布是聚集分布,变程RSD(m)为2.3169～42.00711,表明草间小黑蛛种群对桃蚜种群在空间位置上具有追随关系,对其种群数量的控制有较大作用.     

Comparing short and long‐distance dispersal: modelling and field case studies

Dispersal is a factor of great importance in determining a species spatial distribution. Short distance dispersal (SDD) and long distance dispersal (LDD) strategies yield very different spatial distributions. In this paper we compare spatial spread patterns from SDD and LDD simulations, contrast them with patterns from field data, and assess the significance of biological and population traits. Simulated SDD spread using an exponential function generates a single circular patch with a well‐defined invasion front showing a travelling‐wave structure. The invasive spread is relatively slow as it is restricted to reproductive individuals occupying the outer zone of the circular patch. As a consequence of this dispersal dynamics, spread is slower than spread generated by LDD. In contrast, the early and fast invasion of the entire habitat mediated by power law LDD not only involves a significantly greater invasion velocity, but also an entirely different habitat occupation. As newly dispersed individuals soon reach very distant portions of the habitat as well as the vicinity of the original dispersal focus, new growing patches are generated while the main patch increases its own growth absorbing the closest patches. As a consequence of both dispersal and lower density dependence, growth of the occupied area is much faster than with SDD. SDD and LDD also differ regarding pattern generation. With SDD, fractal patterns appear only in the border of the invasion front in SDD when competitive interaction with residents is included. In contrast, LDD patterns show fractality both in the spatial arrangements of patches as well as in patch borders. Moreover, values of border fractal dimension inform on the dispersal process in relation with habitat heterogeneity. The distribution of patch size is also scale‐free, showing two power laws characteristic of small and large patch sizes directly arising from the dispersal and reproductive dynamics. Ecological factors like habitat heterogeneity are relevant for dispersal, although its importance is greater for SDD, lowering the invasion velocity. Among the life history traits considered, adult mortality, the juvenile bank and mean dispersal distance are the most relevant for SDD. For LDD, habitat heterogeneity and changes in life history traits are not so relevant, causing minor changes in the values of the scale‐free parameters. Our work on short and long distance dispersal shows novel theoretical differences between SDD and LDD in invasive systems (mechanisms of pattern formation, fractal and scaling properties, relevance of different life history traits and habitat variables) that correspond closely with field examples and were not analyzed, at least in this degree of detail, by the previously existing models.     

Retention time variability as a mechanism for animal mediated long-distance dispersal

Long-distance dispersal (LDD) events, although rare for most plant species, can strongly influence population and community dynamics. Animals function as a key biotic vector of seeds and thus, a mechanistic and quantitative understanding of how individual animal behaviors scale to dispersal patterns at different spatial scales is a question of critical importance from both basic and applied perspectives. Using a diffusion-theory based analytical approach for a wide range of animal movement and seed transportation patterns, we show that the scale (a measure of local dispersal) of the seed dispersal kernel increases with the organisms' rate of movement and mean seed retention time. We reveal that variations in seed retention time is a key determinant of various measures of LDD such as kurtosis (or shape) of the kernel, thinkness of tails and the absolute number of seeds falling beyond a threshold distance. Using empirical data sets of frugivores, we illustrate the importance of variability in retention times for predicting the key disperser species that influence LDD. Our study makes testable predictions linking animal movement behaviors and gut retention times to dispersal patterns and, more generally, highlights the potential importance of animal behavioral variability for the LDD of seeds.     

Dandelion Seed Dispersal: The Horizontal Wind Speed Does Not Matter for Long-Distance Dispersal - it is Updraft!

Abstract: Long-distance dispersal of seeds (LDD) surely affects most ecological and evolutionary processes related to plant species. Hence, numerous attempts to quantify LDD have been made and, especially for wind dispersal, several simulation models have been developed. However, the mechanisms promoting LDD by wind still remain ambiguous and the effects of different weather conditions on LDD, although recognized as important, have only rarely been investigated. Here we examine the influence of wind speed and updrafts on dispersal of dandelion ( Taraxacum officinale agg.), a typical wind-dispersed herb of open habitats. We used PAPPUS, a weather-sensitive mechanistic simulation model of wind dispersal, which considers frequency distribution of weather conditions during the period the simulation refers to. A simulation for the 4-month shedding period of dandelion shows that high wind speed does not promote LDD. In contrast, vertical turbulence, especially convective updrafts, are of overwhelming importance. Mainly caused by updrafts, in the simulations more than 0.05 % of dandelion seeds were dispersed beyond 100 m, a distance commonly used to define LDD. We conclude that long-distance dispersal of seeds of herbaceous species with falling velocities < 0.5 - 1.0 ms^-1 is mainly caused by convective updrafts.     

The role of migratory ducks in the long‐distance dispersal of native plants and the spread of exotic plants in Europe

Little is known about the role of migratory waterfowl in the long‐distance dispersal (LDD) of seeds. We studied the gut contents of 42 teals Anas crecca collected in the Camargue, southern France, and found intact seeds of 16 species. There was no relationship between the probability that a given seed species was found intact in the lower gut, and the seed hardness or size. The number of seeds found in the oesophagus and gizzard (a measure of ingestion rate) was the only significant predictor of the occurrence of intact seeds in the lower gut, so studies of waterfowl diet can be used as surrogates of dispersal potential. In a literature review, we identified 223 seed species recorded in 25 diet studies of teal, pintail Anas acuta, wigeon A. penelope or mallard A. platyrhynchos in Europe. We considered whether limited species distribution reduces the chances that a seed can be carried to suitable habitat following LDD. Overall, 72% of plant species recorded in duck diets in southern Europe (36 of 50) were also recorded in the north, whereas 97% of species recorded in duck diets in the north (137 of 141) were also recorded in the south. This suggests a great potential for LDD, since most dispersed plants species occur throughout the migratory range of ducks. Migratory ducks are important vectors for both terrestrial and aquatic plant species, even those lacking the fleshy fruits or hooks typically used to identify seeds dispersed by birds. Finally, we show ducks are important vectors of exotic plant species. We identified 14 alien to Europe and 44 native to Europe but introduced to some European countries whose seeds have been recorded in duck diet.     

Mechanisms of long-distance seed dispersal

Growing recognition of the importance of long-distance dispersal (LDD) of plant seeds for various ecological and evolutionary processes has led to an upsurge of research into the mechanisms underlying LDD. We summarize these findings by formulating six generalizations stating that LDD is generally more common in open terrestrial landscapes, and is typically driven by large and migratory animals, extreme meteorological phenomena, ocean currents and human transportation, each transporting a variety of seed morphologies. LDD is often associated with unusual behavior of the standard vector inferred from plant dispersal morphology, or mediated by nonstandard vectors. To advance our understanding of LDD, we advocate a vector-based research approach that identifies the significant LDD vectors and quantifies how environmental conditions modify their actions.     

Does seed retention potential affect the distribution of plant species in highly fragmented calcareous grasslands?

Long-distance dispersal is a crucial factor in the life-cycle of plants, especially in our modern, highly fragmented landscapes. Because natural herds of large animals have disappeared and grazing practices have been abandoned, important potential vectors for seed dispersal over large distances may have been lost. In the context of the re-establishment of grazing management for nature conservation purposes, it is therefore important to gain insight in the ability of grazing animals to act as seed dispersal vectors. Whereas local dispersal mainly occurs through standard vectors typically described based on morphological adaptations of the diaspore, large herbivores act as non-standard seed dispersers. Therefore, traditional dispersal classes are loosing scientific relevance and continuous predictors of dispersal potential have been proposed. Here, we explored whether dispersal related plant traits, including the "seed retention potential", could explain the distribution patterns of 180 plant species over 64 fragmented semi-natural calcareous grasslands in Belgium. The distribution of habitat specialist plant species was strongly determined by the degree of isolation of the grasslands. Interestingly, species distribution patterns were clearly linked with a species' potential to migrate through large grazers, as quantified by its retention potential: species producing seeds with high retention capacity were less affected by habitat isolation. Categorical dispersal classes based on seed morphology did not explain a species' response to fragment isolation. Although seed retention potential outperformed simple seed dimensional traits, plant height, which is an indicator of epizoochorous attachment potential, was even more important. Therefore we suggest further extension of the epizoochorous retention potential model by incorporating basic ecological mechanisms that effectively contribute to successful dispersal events.     

Why are small seeds dispersed through animal guts: large numbers or seed size per se?

Long-distance dispersal of seeds is an important process in metapopulation dynamics and in plant migrations, but at the same time extremely difficult to observe or quantify directly. If seed dispersal ability were related to attributes of seeds or motherplants, long-distance seed dispersal would be predictable by indirect approximation using easy-to-measure traits. Seed size has been suggested to be such a key trait in seed dispersal ability. However, having smaller seeds also implies having more numerous seeds per plant individual (given equal reproductive effort), and consequently increases the probability of seeds being ingested accidentally. The question is whether small-seeded species are more abundant in herbivore dung because smaller seed size increases survival rate during gut passage or because they are produced (and ingested) in greater numbers than larger seeds. We investigated endozoochorous seed dispersal via cattle grazing a meadow, and related seed abundance in dung samples to seed attributes. We found that seeds were ingested and passed through the bovine intestinal tract in proportion to the numbers produced per unit area in the grazed vegetation. In contrast, no relationship could be found between endozoochorous dispersal potential (measured as abundance of seeds in dung samples corrected for seed output in the grazed vegetation) and seed attributes such as seed mass, seed shape (roundness), and thickness of the seed coat. This finding underlines the importance of seed number in plant dispersal ability. In addition, it shows that grazing mammals may constitute an important dispersal vector for many plant species conventionally classified as 'unspecialised'.     

Long‐distance dispersal syndromes matter: diaspore–trait effect on shaping plant distribution across the Canary Islands

Oceanic islands emerge lifeless from the seafloor and are separated from continents by long stretches of sea. Consequently, all their species had to overcome this stringent dispersal filter, making these islands ideal systems to study the biogeographic implications of long‐distance dispersal (LDD). It has long been established that the capacity of plants to reach new islands is determined by specific traits of their diaspores, historically called dispersal syndromes. However, recent work has questioned to what extent such dispersal‐related traits effectively influence plant distribution between islands. Here we evaluated whether plants bearing dispersal syndromes related to LDD – i.e. anemochorous (structures that favour wind dispersal), thalassochorous (sea dispersal), endozoochorous (internal animal dispersal) and epizoochorous (external animal dispersal) syndromes – occupy a greater number of islands than those with unspecialized diaspores by virtue of their increased dispersal ability. We focused on the native flora of the lowland xeric communities of the Canary Islands (531 species) and on the archipelago distribution of the species. We controlled for several key factors likely to affect the role of LDD syndromes in inter‐island colonization, namely: island geodynamic history, colonization time and phylogenetic relationships among species. Our results clearly show that species bearing LDD syndromes have a wider distribution than species with unspecialized diaspores. In particular, species with endozoochorous, epizoochorous and thalassochorous diaspore traits have significantly wider distributions across the Canary archipelago than species with unspecialized and anemochorous diaspores. All these findings offer strong support for a greater importance of LDD syndromes on shaping inter‐island plant distribution in the Canary Islands than in some other archipelagos, such as Galápagos and Azores.     

Increases in air temperature can promote wind-driven dispersal and spread of plants

Long-distance dispersal (LDD) of seeds and pollen shapes the spatial dynamics of plant genotypes, populations and communities. Quantifying LDD is thus important for predicting the future dynamics of plants exposed to environmental changes. However, environmental changes can also alter the behaviour of LDD vectors: for instance, increasing air temperature may enhance atmospheric instability, thereby altering the turbulent airflow that transports seed and pollen. Here, we investigate temperature effects on wind dispersal in a boreal forest using a 10-year time series of micrometeorological measurements and a Lagrangian stochastic model for particle transport. For a wide range of dispersal and life history types, we found positive relations between air temperature and LDD. This translates into a largely consistent positive effect of +3°C warming on predicted LDD frequencies and spread rates of plants. Relative increases in LDD frequency tend to be higher for heavy-seeded plants, whereas absolute increases in LDD and spread rates are higher for light-seeded plants for which wind is often an important dispersal vector. While these predicted increases are not sufficient to compensate forecasted range losses and environmental changes can alter plant spread in various ways, our results generally suggest that warming can promote wind-driven movements of plant genotypes and populations in boreal forests.