外来入侵植物风传扩散过程模拟模型选择

种子扩散是外来入侵植物异地入侵的重要手段,对入侵种群扩散机制的清晰认识是有效进行入侵防治和生态管理的基础。本文在研究入侵植物风传扩散过程中的主要影响参数的基础上,分析了各参数对不同扩散过程造成的影响,总结出不同的扩散模拟过程中需要应用的影响参数,阐述了这些参数的参数化过程及相应的计算方法。对已有风传扩散模拟模型做了重点介绍,并结合入侵植物自身特征重点解析了其风传扩散过程模拟模型的选择依据。其中,对于风传扩散过程有直接、明显的影响参数是种子释放高度和水平风速。对于入侵草本、灌木植物而言,入侵区域植被高度是必须关注的影响参数。在长距离扩散研究中,以乔木和大灌木为研究对象时需要重点考虑植被叶片与水平风速垂直分布引起的冠层及冠层以上区域大气湍流所带来的影响;小灌木与草本植物则需注意由地表温度引起的温暖气团上升带来的影响。模拟模型中,2Dt模型和混合韦布尔模型可以有效描述扩散体扩散核近端与远端的扩散曲线。对更为详细的长距离扩散模拟模型选择时,需要考虑不同的大气湍流类型、扩散体大小、植被类型、扩散核维度以及模拟精度要求。     

风传草本植物种子空间传播新模型

种子阶段产生的时空格局对植物种群迁移十分关键,在植物种群动态、外来种入侵、保护生物学领域有重要意义。大多数种子从母株处传播距离较短,少数种子可以进行长距离传播。近30多年对种子传播的大量研究集中在近距离传播,这与目前的技术很难直接跟踪长距离传播的种子有关,尤其是草本植物种子传播数据极难获取。在现实条件约束下,利用模型模拟种子传播显示了重要作用,但目前尚未形成种子密度随距离变化的统一性函数或分布曲线,我国更是鲜见此方面研究成果。提出了一种全新的风传草本植物种子的空间传播模型,即准麦克斯韦空间分布模型,将种子长、中、短距离的传播归纳于统一的模型之中;实现无风或有风时,任意方向及区域的种子传播模拟,并且利用前人已有的实验或实测数据对模型进行了适应性分析,旨在推动此领域研究的进一步探索与发展。     

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。     

五种杂草种子沉降速度

在室内无风条件下,研究了加拿大一枝黄花(Solidago canadensis)、紫菀(Aster tataricus)、小飞蓬(Conyza canadensis)、山莴苣(Lagedium sibiricum)和芦苇(Phragmites communis)等5种具有风传扩散潜力的杂草种子的沉降速度,并对5种杂草种子的远距离扩散能力进行了推测.结果表明:在不受其他作用如气流等影响的情况下,加拿大一枝黄花、紫菀、小飞蓬、山莴苣和芦苇等5种杂草种子,分别在0.5、1.0、1.5和2.0 m等4个高度自由释放一定数量的种子,记录其种子在各个高度的沉降时间,该5种子沉降速度依次为14.1、20.4、20.8、23.3和33.3 cm·s-1;5种杂草种子中加拿大一枝黄花种子沉降速度较慢,更适于长距离风传扩散.     

石林地质公园不同群落类型植物果实组成与种子散布特征

喀斯特植物群落果实类型和种子散布特征的研究, 对于理解喀斯特植物群落更新与拓展特征, 进而揭示喀斯特石漠化地区恢复植被的自然扩展机制及制定植被恢复规划具有重要意义。该研究以石林地质公园不同群落类型为对象, 研究不同群落的果实类型谱与种子散布谱, 以揭示果实类型与种子散布方式对恢复植被的影响。采用石林地质公园不同群落各1 hm ²样地记名计数法调查的维管束物种名录, 依据Flora of China对物种果实的描述确定其果实类型; 根据文献、数据库以及果实、种子形态性状确定种子散布方式。结果表明: (1)石林地质公园282种维管束植物共16种果实类型(含孢子), 其中瘦果、蒴果、浆果和核果的物种比例均大于10%。木本植物、草本植物、藤本植物的优势果实类型分别为核果、瘦果和浆果。石林地质公园各群落的木本植物、草本植物和藤本植物的果实类型谱差异不显著。在原生林、次生林、灌丛、灌草丛和人工辅助云南松林不同群落中, 浆果和核果的物种比例减少, 瘦果、颖果和蒴果的物种比例增加。(2)石林地质公园植物种子以动物传播为主, 其次为风传播。在原生林、次生林、灌丛、灌草丛和人工辅助云南松林不同群落中, 风传播的物种比例增加了73%-87%, 动物传播(包括鸟类传播)减少了31%-36%。(3)物种的种子散布方式直接影响到植被恢复物种和恢复策略的选择。基于恢复目标, 选择自然传播能力强的乡土物种或地带性植被关键种, 辅以传播廊道, 有助于喀斯特的植被恢复。     

Fruit types and seed dispersal modes of plants in different communities in Shilin Geopark,Yunnan, China

喀斯特植物群落果实类型和种子散布特征的研究, 对于理解喀斯特植物群落更新与拓展特征, 进而揭示喀斯特石漠化地区恢复植被的自然扩展机制及制定植被恢复规划具有重要意义。该研究以石林地质公园不同群落类型为对象, 研究不同群落的果实类型谱与种子散布谱, 以揭示果实类型与种子散布方式对恢复植被的影响。采用石林地质公园不同群落各1 hm ²样地记名计数法调查的维管束物种名录, 依据Flora of China对物种果实的描述确定其果实类型; 根据文献、数据库以及果实、种子形态性状确定种子散布方式。结果表明: (1)石林地质公园282种维管束植物共16种果实类型(含孢子), 其中瘦果、蒴果、浆果和核果的物种比例均大于10%。木本植物、草本植物、藤本植物的优势果实类型分别为核果、瘦果和浆果。石林地质公园各群落的木本植物、草本植物和藤本植物的果实类型谱差异不显著。在原生林、次生林、灌丛、灌草丛和人工辅助云南松林不同群落中, 浆果和核果的物种比例减少, 瘦果、颖果和蒴果的物种比例增加。(2)石林地质公园植物种子以动物传播为主, 其次为风传播。在原生林、次生林、灌丛、灌草丛和人工辅助云南松林不同群落中, 风传播的物种比例增加了73%-87%, 动物传播(包括鸟类传播)减少了31%-36%。(3)物种的种子散布方式直接影响到植被恢复物种和恢复策略的选择。基于恢复目标, 选择自然传播能力强的乡土物种或地带性植被关键种, 辅以传播廊道, 有助于喀斯特的植被恢复。     

影响加拿大一枝黄花种子非随机脱落的因素

为揭示加拿大一枝黄花(Solidago canadensis)种群扩散机制, 明确种子的脱落及风传扩散在其种群蔓延中的作用, 在人工环境下测定了不同湍流强度、风速和湿度处理下种子脱落的差异, 并对脱落种子与未脱落种子进行形态学特征对比。结果表明: 加拿大一枝黄花的种子脱落受湍流、风速和湿度等因素的共同影响。水平气流下种子的脱落阈值为5.1 m·s^-1, 并随着风速增加, 种子的脱落率增加。与模拟水平气流相比, 模拟垂直气流下种子的脱落阈值显著偏小。相对于层流状态, 湍流的存在显著提高了种子的脱落率, 平均增幅超过300%; 但单纯提高湍流强度对种子脱落率的影响不显著。增加湿度则显著降低种子的脱落率。种子形态学特征对比结果表明, 脱落种子的冠毛数量和冠毛夹角显著高于未脱落种子。该研究结果为研究加拿大一枝黄花种子脱落规律和风传扩散机制提供了科学依据, 也为其他入侵性杂草种子的扩散机制及入侵过程提供了 借鉴。     

扩散对破碎化景观上宿主-寄生种群动态的影响

景观破碎化和扩散是空间种群模型的重要因素,对生物入侵存在着深远的影响。本章将基于偶对近似模型,探讨由局部和全局宿主-寄生相互作用共同决定的扩散模式对破坏性景观上疾病入侵与传播的影响。其中,生境破坏由生境丧失量与生境破碎化程度来描述。模拟结果显示,宿主和病毒的全局扩散对疾病的入侵与种群密度产生不对称效应：病毒的全局扩散对系统产生的影响较宿主的全局扩散更为显著。不同扩散模式下,生境丧失越高或破碎化程度越低,均将越有害于寄生病毒的入侵;同时,生境的破坏程度也显著地影响了入侵阈值对扩散模式的响应机制。本文研究结果暗示,景观破碎化的空间分布格局以及病毒扩散的限制均可作为物种保护与管理中有效的疾病控制策略。该研究结果在一定意义上丰富和发展了寄生感染理论,为物种保护提供了生态学理论依据。     

Factors influencing the nonrandom abscission of Solidago canadensis seeds

 为揭示加拿大一枝黄花(Solidago canadensis)种群扩散机制, 明确种子的脱落及风传扩散在其种群蔓延中的作用, 在人工环境下测定了不同湍流强度、风速和湿度处理下种子脱落的差异, 并对脱落种子与未脱落种子进行形态学特征对比。结果表明: 加拿大一枝黄花的种子脱落受湍流、风速和湿度等因素的共同影响。水平气流下种子的脱落阈值为5.1 m·s^–1, 并随着风速增加, 种子的脱落率增加。与模拟水平气流相比, 模拟垂直气流下种子的脱落阈值显著偏小。相对于层流状态, 湍流的存在显著提高了种子的脱落率, 平均增幅超过300%; 但单纯提高湍流强度对种子脱落率的影响不显著。增加湿度则显著降低种子的脱落率。种子形态学特征对比结果表明, 脱落种子的冠毛数量和冠毛夹角显著高于未脱落种子。该研究结果为研究加拿大一枝黄花种子脱落规律和风传扩散机制提供了科学依据, 也为其他入侵性杂草种子的扩散机制及入侵过程提供了借鉴。     

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.     

Human-mediated dispersal of seeds over long distances

Human activities have fundamental impacts on the distribution of species through altered land use, but also directly by dispersal of propagules. Rare long-distance dispersal events have a disproportionate importance for the spread of species including invasions. While it is widely accepted that humans may act as vectors of long-distance dispersal, there are few studies that quantify this process. We studied in detail a mechanism of human-mediated dispersal (HMD). For two plant species we measured, over a wide range of distances, how many seeds are carried by humans on shoes. While over half of the seeds fell off within 5m, seeds were regularly still attached to shoes after 5 km. Semi-mechanistic models were fitted, and these suggested that long-distance dispersal on shoes is facilitated by decreasing seed detachment probability with distance. Mechanistic modelling showed that the primary vector, wind, was less important as an agent of long-distance dispersal, dispersing seeds less than 250 m. Full dispersal kernels were derived by combining the models for primary dispersal by wind and secondary dispersal by humans. These suggest that walking humans can disperse seeds to very long distances, up to at least 10 km, and provide some of the first quantified dispersal kernels for HMD.     

Mechanistic analytical models for long-distance seed dispersal by wind

We introduce an analytical model, the Wald analytical long-distance dispersal (WALD) model, for estimating dispersal kernels of wind-dispersed seeds and their escape probability from the canopy. The model is based on simplifications to well-established three-dimensional Lagrangian stochastic approaches for turbulent scalar transport resulting in a two-parameter Wald (or inverse Gaussian) distribution. Unlike commonly used phenomenological models, WALD's parameters can be estimated from the key factors affecting wind dispersal--wind statistics, seed release height, and seed terminal velocity--determined independently of dispersal data. WALD's asymptotic power-law tail has an exponent of -3/2, a limiting value verified by a meta-analysis for a wide variety of measured dispersal kernels and larger than the exponent of the bivariate Student t-test (2Dt). We tested WALD using three dispersal data sets on forest trees, heathland shrubs, and grassland forbs and compared WALD's performance with that of other analytical mechanistic models (revised versions of the tilted Gaussian Plume model and the advection-diffusion equation), revealing fairest agreement between WALD predictions and measurements. Analytical mechanistic models, such as WALD, combine the advantages of simplicity and mechanistic understanding and are valuable tools for modeling large-scale, long-term plant population dynamics.     

Seed Dispersal Distances and Plant Migration Potential in Tropical East Asia

Most predictions of vegetation responses to anthropogenic climate change over the next 100 yr are based on plant physiological tolerances and do not account for the ability of plant species to migrate over the distances required in the time available, or the impact of habitat fragmentation on this ability. This review assesses the maximum routine dispersal distances achievable in tropical East Asia and their vulnerability to human impacts. Estimates for various plant–vector combinations range from < 10 m, for species dispersed by ants or mechanical means, to > 10 km for some species dispersed by wind (tiny seeds), water, fruit pigeons, large fruit bats (tiny seeds), elephants, rhinoceroses, and people. Most plant species probably have maximum dispersal distances in the 100–1000 m range, but the widespread, canopy-dominant Dipterocarpaceae and Fagaceae are normally dispersed < 100 m. Large fruit bats and fruit pigeons are particularly important for long-distance dispersal in fragmented landscapes and should be protected from hunting. The maximum seed dispersal distances estimated in this study are potentially sufficient for many plant species to track temperature changes in steep topography, but are far too small for a significant role in mitigating climate change impacts in the lowlands, where temperature and rainfall gradients are much more shallow.     

Human-Mediated Dispersal of Seeds by the Airflow of Vehicles

Human-mediated dispersal is known as an important driver of long-distance dispersal for plants but underlying mechanisms have rarely been assessed. Road corridors function as routes of secondary dispersal for many plant species but the extent to which vehicles support this process remains unclear. In this paper we quantify dispersal distances and seed deposition of plant species moved over the ground by the slipstream of passing cars. We exposed marked seeds of four species on a section of road and drove a car along the road at a speed of 48 km/h. By tracking seeds we quantified movement parallel as well as lateral to the road, resulting dispersal kernels, and the effect of repeated vehicle passes. Median distances travelled by seeds along the road were about eight meters for species with wind dispersal morphologies and one meter for species without such adaptations. Airflow created by the car lifted seeds and resulted in longitudinal dispersal. Single seeds reached our maximum measuring distance of 45 m and for some species exceeded distances under primary dispersal. Mathematical models were fit to dispersal kernels. The incremental effect of passing vehicles on longitudinal dispersal decreased with increasing number of passes as seeds accumulated at road verges. We conclude that dispersal by vehicle airflow facilitates seed movement along roads and accumulation of seeds in roadside habitats. Dispersal by vehicle airflow can aid the spread of plant species and thus has wide implications for roadside ecology, invasion biology and nature conservation.     

Seed-dispersal distributions by trumpeter hornbills in fragmented landscapes

Frugivorous birds provide important ecosystem services by transporting seeds of fleshy fruited plants. It has been assumed that seed-dispersal kernels generated by these animals are generally leptokurtic, resulting in little dispersal among habitat fragments. However, little is known about the seed-dispersal distribution generated by large frugivorous birds in fragmented landscapes. We investigated movement and seed-dispersal patterns of trumpeter hornbills (Bycanistes bucinator) in a fragmented landscape in South Africa. Novel GPS loggers provide high-quality location data without bias against recording long-distance movements. We found a very weakly bimodal seed-dispersal distribution with potential dispersal distances up to 14.5 km. Within forest, the seed-dispersal distribution was unimodal with an expected dispersal distance of 86 m. In the fragmented agricultural landscape, the distribution was strongly bimodal with peaks at 18 and 512 m. Our results demonstrate that seed-dispersal distributions differed when birds moved in different habitat types. Seed-dispersal distances in fragmented landscapes show that transport among habitat patches is more frequent than previously assumed, allowing plants to disperse among habitat patches and to track the changing climatic conditions.     

Linking landscape history and dispersal traits in grassland plant communities

Dispersal limitation and long-term persistence are known to delay plant species’ responses to habitat fragmentation, but it is still unclear to what extent landscape history may explain the distribution of dispersal traits in present-day plant communities. We used quantitative data on long-distance seed dispersal potential by wind and grazing cattle (epi- and endozoochory), and on persistence (adult plant longevity and seed bank persistence) to quantify the linkages between dispersal and persistence traits in grassland plant communities and current and past landscape configurations. The long-distance dispersal potential of present-day communities was positively associated with the amounts of grassland in the historical (1835, 1938) landscape, and with a long continuity of grazing management—but was not associated with the properties of the current landscape. The study emphasises the role of history as a determinant of the dispersal potential of present-day grassland plant communities. The importance of long-distance dispersal processes has declined in the increasingly fragmented modern landscape, and long-term persistent species are expected to play a more dominant role in grassland communities in the future. However, even within highly fragmented landscapes, long-distance dispersed species may persist locally—delaying the repayment of the extinction debt.     

Seasonal and Diurnal Patterns of Spore Release Can Significantly Affect the Proportion of Spores Expected to Undergo Long-Distance Dispersal

Many of the fungal pathogens that threaten agricultural and natural systems undergo wind-assisted dispersal. During turbulent wind conditions, long-distance dispersal can occur, and airborne spores are carried over distances greater than the mean. The occurrence of long-distance dispersal is an important ecological process, as it can drastically increase the extent to which pathogen epidemics spread across a landscape, result in rapid transmission of disease to previously uninfected areas, and influence the spatial structure of pathogen populations in fragmented landscapes. Since the timing of spore release determines the wind conditions that prevail over a dispersal event, this timing is likely to affect the probability of long-distance dispersal occurring. Using a Lagrangian stochastic model, we test the effect of seasonal and diurnal variation in the release of spores on wind-assisted dispersal. Spores released during the hottest part of the day are shown to be more likely to undergo long-distance dispersal than those released at other times. Furthermore, interactions are shown to occur between seasonal and diurnal patterns of release. These results have important consequences for further modelling of wind-assisted dispersal and the use of models to predict the spread of fungal pathogens and resulting population and epidemic dynamics.