Effects of seed aggregation on the removal rates of elaiosome-bearing Chelidonium majus and Viola odourata seeds carried by Formica polyctena ants

In a field dominated by Formica polyctena Foerst. ants, we examined the effect of seed aggregation on the seed-removal rates of two plant species: a large-seeded obligate myrmecochore Viola odorata L. and a small-seeded diplochore Chelidonium majus L., which was autochorous as well as myrmecochorous. The effect was statistically non-significant in V. odorata but significant in C. majus, with more closely aggregated seeds having higher removal rates. The large seeds of the obligate myrmecochore were more quickly discovered and repeatedly removed by ant workers than were the small seeds of the diplochore.     

川西亚高山65年人工云杉林种子雨、种子库和幼苗定居研究

川西亚高山人工针叶林已成为亚高山森林的重要组成部分, 它们是否具有持续的自然更新能力, 是决定川西亚高山针叶林群落演替方向和维持该区针叶林大面积存在的基础。以川西米亚罗亚高山人工云杉(Picea asperata)林(65 a)为研究对象, 对种子雨量年际变化、土壤种子库动态、种子萌发和幼苗定居等更新过程的关键环节进行了连续7年(2002-2008年)的野外观测, 以研究人工云杉林更新潜力及影响其更新的限制因素。结果表明: 该区云杉林种子雨一般从每年的10月初开始下落, 一直到翌年的1月底或2月初结束; 云杉种子散落存在明显的大小年现象, 种子散落周期为4年, 且大小年之间种子产量差异极大; 云杉种子从下落到土壤到种子完全失去活力不到1年时间, 属于Thompson和Grime定义的第II类土壤种子库类型。腐烂死亡和动物取食是土壤种子库损耗的主要因素, 而种子通过萌发真正转化为幼苗的比例非常低, 仅占2002年下落种子总量的3.6%。种子萌发后, 环境筛的作用导致云杉幼苗大量死亡, 尤其是在种子萌发后的一个生长季节内, 其死亡率高达78%。凋落物和苔藓是构成人工云杉林下地表的两种主要地被物类型, 二者占所有调查幼苗数量的93%左右; 两种地被物类型上0-2 cm层幼苗存活率最高, 分别占存活幼苗总数的76.07%和86.72%, 随地被物厚度增加, 幼苗存活率呈明显下降趋势, 而幼苗死亡率呈明显升高的趋势, 表明林下地被物厚度也是影响云杉幼苗定居的重要因素。两种地被物对幼苗生长的影响不同, 除株高之外, 分布在苔藓上的云杉幼苗生长参数(地径、分枝数、干重以及干重年增长率)明显高于分布于凋落物上的幼苗, 表明苔藓地被物更有利于云杉幼苗定居。尽管该区大量云杉种子下落, 但由于种子的高损耗率、幼苗的低输出率以及萌发幼苗的高死亡率, 使得人工云杉林下种子通过萌发转为实生幼苗的数量非常少, 最终真正能补充到云杉种群的个体数量非常有限。     

Spatial patterns in seed bank and vegetation of semi-natural mountain meadows

Soil seed bank composition and vegetative spatial patterns were studied in four mountain meadow communities in the Broto Valley (N. Spain), in order to analyse the differences that might exists between the two life forms. Soil and vegetation samples were taken at 1 m intervals from 10 m × 10 m quadrats in each meadow in one-year study. The spatial distribution of species was analysed along with the calculation of an autocorrelation coefficient which takes account of the relative position of samples: (Moran’s I). The results indicate that the abundance of the majority of the species in the seed bank and in the vegetation are randomly distributed, the percentage of species with a clumped distribution only exceeds 35% in the vegetation of one meadow and none of the taxa identified showed a uniform spatial organisation. The species that were distributed in the seed bank in a clumped pattern in more than one meadow were those of the pioneer species (Anagallis arvensis, Centaurium erythraea, Lamium purpureum and Stellaria media). All of these formed long-term persistent seed banks but were absent in the established vegetation in these meadows. According to the results, there exist not only differences between the spatial distribution of the species present in the same community, but also, that some species change their pattern of distribution according to the life form and to the grassland type in which they are found.     

Soil seed bank and species turnover in a limestone grassland

Abstract. The vegetation and soil seed bank in a limestone grassland in southern Sweden were studied in permanent 1-m² subplots which had been either grazed or not grazed for 17 yr. Of the 92 species recorded, 18 were present only in the seed bank, 28 were more frequent and 24 were less frequent in the seed bank than in the vegetation and 22 were not detected in the seed bank. Among the species present in the seed bank, therophytes were over-represented. Species turnover in the vegetation was estimated from presence/absence data collected in 1980, 1986 and 1990. Turnover was high, but there were no differences between grazed and ungrazed subplots. The turnover for individual species was also high in many cases. There was no clear relationship between the turnover of a species and its presence in the persistent seed bank.     

可获得种子量和种子出苗对草地裸沙斑植被恢复的相对制约

以科尔沁沙化草地的裸斑为研究对象,布设种子捕捉器监测生长季传播到裸斑的种子数量、物种组成,并人工播种监测裸斑中出苗情况,研究可获得种子数量和出苗对裸斑植被恢复的制约.结果表明:沙生植物3种,种子数为71 ～623粒·m-2,非沙生植物9种,种子数为73 ～410粒·m-2;裸斑中沙生植物的出苗数(9.3 ～22.3株·m-2)高于非沙生植物(0 ～3.4株·m-2);沙生植物在裸斑的出苗率最高为沙蓬(12.2％ ～34.1％),其次是乌丹蒿(4.5％ ～10.3％),小叶锦鸡儿(0～0.5％),非沙生植物狗尾草出苗率为0～6％,蒲公英为0 ～0.4％,其余植物种出苗率为0;沙生植物裸斑出苗数(9.3 ～22.3株·m-2)高于植被区(7.7株·m-2),而非沙生植物在裸斑的出苗数(0～3.4株·m-2)低于植被区(4.5株·m-2).可见,裸斑植被恢复中,沙生植物的可获得种子量制约高于出苗制约;非沙生植物的可获得种子量制约与出苗制约相当.     

Roles of species-preferential seed dispersal by ants and endozoochory in Melampyrum (Orobanchaceae)

Aims Melampyrum pratense and M. subalpinum are two myrmecochorous species, which possess similar habitat requirements and frequently occur together. Despite this, their population sizes differ markedly. Melampyrum pratense populations are usually very large, whereas M. subalpinum has rather small and isolated populations. We suggest that such an imbalance might be partially influenced by the difference in ant-mediated seed-removal rates. Genus Melampyrum is considered to be exclusively myrmecochorous, though to achieve the recent distribution of some Melampyrum species during the Holocene myrmecochory would be highly insufficient. We suggest that endozoochory takes place in the long-distance migration, whereas myrmecochory is important for the removal of seeds on a local scale.Methods For seed-preference analysis, M. pratense and M. subalpinum mixed seed samples were placed around Formica polyctena anthills. After a period of time, the remaining seeds of both species were counted for each sample. The results were analysed by analysis of variance and generalized linear mixed-effect model. To test myrmecochorous removal distances, M. pratense seeds were covered with fluorescent dactyloscopic powder and placed in the vicinity of a large ant trail. The area around the starting plot was searched in the dark using UV LED torchlight 7h after the beginning. The distance from the starting plot was measured for each seed found. Birds, rodents, leporine and a ruminant were fed with M. pratense seeds and fresh plants to test the possibility of endozoochorous dispersal of the species. Animal droppings were searched for intact seeds.Important findings Our field studies show that from mixed seed samples, containing both species, ants significantly preferred the seeds of M. pratense. This may be one of factors that has positive influence on M. pratense success in seed dispersal on mixed stands and consequently in the colonization of favourable sites. Experiments focusing on ant-mediated dispersal distance revealed that F. polyctena ants are able to move seeds over a distance of 36 m in only 7h. This distance is among the furthest known myrmecochorous removals of forest plant seeds. A new Melampyrum seed disperser Oligolophus tridens (Opiliones) was observed repeatedly. Our pilot study documented that Melampyrum seeds are able to pass through the digestive tract of a cow intact. This suggests that large ruminants such as deer, bison or forest-grazing livestock may function as important long-distance dispersers of Melampyrum species.     

Influence of termites on the soil seed bank in an African savannah

In savannah ecosystems, termites drive key ecosystem processes, such as primary production through creation of patchiness in soil nutrients availability around their nests. In this study, we evaluated the role of termites in altering the soil seed bank size, an important ecosystem component that has often been overlooked in previous work. Data on above ground vegetation and soil seed bank samples were collected from four microhabitats, that is, the wooded mound, unwooded mound, tree sub‐canopy and the open grassland matrix in a protected game reserve in south‐central Zimbabwe. The seedling emergence method was then used to identify species present in the soil samples. One‐way analysis of variance followed by Tukey's multiple comparison tests was executed to test for significant differences in plant species richness among the four microhabitats. The results indicate that plant species richness was high on wooded termite mound but did not differ between the unwooded and the sub‐canopy microhabitats. The open grassland microhabitat had the lowest plant species richness. The influence of termites on the soil seed bank composition was also life form specific. The herb and woody life forms had significantly (α = 0.05) higher species richness in the soil seed bank at wooded and unwooded termite mounds when compared to the other two microhabitats. Overall, these results imply that termites alter the soil seed bank and the findings enhance our understanding of the significant role termites play in regulating processes in savannah ecosystem.     

Impact of seed abundance on seed processing and dispersal by the red ant Myrmica rubra

1. Myrmecochory sensu stricto is an ant–plant mutualism in which non‐granivorous ants disperse plant diaspores after feeding on their nutrient‐rich seed appendage, the elaiosome. Phenological traits associated with the diaspore can influence the behaviour of ants and thus their ultimate efficiency as seed dispersers. 2. This study investigated how a contrasting availability of seeds (20 vs. 200 seeds) from the diplochorous Chelidonium majus (Papaveraceae, Linnaeus) plant species influences the behaviour of Myrmica rubra (Formicidae, Linnaeus) ants, from the retrieval of seeds until their dispersal outside the ant nest. 3. Regardless of seed abundance, the ants collected the first diaspores at similar rates. Then, seed retrieval sped up over time for large seed sources until satiation took place with only one‐third of the tested colonies wholly depleting abundant seed sources. 4. No active recruitment by trail‐laying ants was triggered, even to an abundant seed source 5. In both conditions of seed abundance, the majority of the diaspores retrieved inside the nest were discarded with the elaiosome removed and were dispersed at similar distances from the nest. 6. The paper concludes with a discussion of how the quantity of seeds released by a plant with a dual mode of dispersal can potentially influence the behaviour of ant dispersers and hence the dispersal efficiency derived from myrmecochory.     

淡水湿地种子库的小尺度空间格局

以长江中下游淡水湿地湖里沼泽为对象,研究了种子库在小尺度空间范围的水平分布格局。在沼泽中选择地表植被分布比较均一的16m×4m大小的方形区域,按1m间隔获取64个内径7·8cm、深5cm的圆柱状土样,通过幼苗萌发法鉴定每个土样的种子库组成。采用2个格局指数(离散系数和Lloyd平均拥挤指数)以及Moran空间自相关系数分析了种子库中优势种的空间分布格局。结果显示种子库由17个物种组成,多年生的锐棱荸荠(Eleocharisacutangula)和龙师草(E·tetraqueter)是种子库中密度最大的物种。两个格局指数显示7个分布频率大于10%的物种的种子全部为聚集分布。MoransI统计分析显示其中只有3个物种为显著的正的空间自相关,表明这3个物种的种子斑块大小超过了目前的两个取样单位之间的距离(1m),而其它4个物种的种子斑块则小于这个尺度。     

Inflow of seeds through the forest edge: evidence from seed bank and vegetation patterns

To determine the influence of the proximity of a forest edge on seed bank composition and diversity, we performed a seed bank sampling at ancient deciduous forests bordering intensive arable fields. Also vegetation patterns were taken into account. We hypothised that forest edges may facilitate the entrance of diaspores of invasive species into the forest and the successive incorporation of these species in the forest seed bank. We noticed a substantial influence of the proximity of an edge on seed bank composition at as well the forested side of the edge as the field side. The forest edge zone was limited to 3 m into the forest and the field edge zone extended 3m into the field. The seed bank samples of field and forest edge are characterised by a higher species diversity and seed density and a higher similarity between seed bank and vegetation, compared to field or forest samples. The forest edges contains fewer pioneer species in comparison with the forest interior and more competitive species and species of edges and clearings compared with field and forest samples. The seed longevity index increases towards the forest interior. We can conclude from our data that the forest and edge seed bank are composed by both seeds from recent dispersal processes and local seed set and by seeds originating from past vegetation on the site. Near the edge, actual seed input seems of primal importance. Further towards the forest interior seed input decreases and long-living seeds of past vegetation become more important. Ancient forest edges thus act as a barrier for seeds of species of the surrounding arable field.     

Impact of management on vegetation dynamics and seed bank formation of inland dune grassland in Hungary

Seed bank formation and 7 years of vegetation dynamics were studied on permanent plots of a dry sandy pasture, Cynodonti-Festucetum pseudovinae. A stand overgrazed by domestic geese and a reference stand void of overgrazing were compared. Apart from this both stands were accidentally grazed by cattle. Vegetation of the overgrazed stand was significantly more species-rich, especially in summer annuals. The reference was dominated with perennials and winter annuals. Composition and dominance changed considerably at both stands but only composition became more alike. Declining species richness and increasing dominance of perennials was found in the overgrazed stand. The reference stand became dominated with the dwarf-shrub Thymus degenianus. Species richness of the overgrazed stand showed greater seasonal and year-to-year variation than that of the reference. No temporal change of nutrient availability was found and neither was a difference detected between the stands.Greenhouse germination revealed more dense and more species-rich seed bank in the overgrazed stand. Its established vegetation and soil seed bank were also more alike. Higher species richness of the overgrazed stand can be associated with intensive propagule dispersal of geese, as indicated by dense seed banks of zoochorous hygrophytes delivered from neighbouring wetlands. Relatively high representation of persistent seed bank records suggests that, except for some sensitive perennials, the studied community is adapted to recurrent disturbances. Community regeneration seems to be limited by slow spread of perennial graminoids.     

Seed rain and soil seed bank of Picea asperata Mast. in subalpine coniferous forest of western Sichuan,China

Abstract

Dragon spruce (Picea asperata Mast.) is widely planted on clear-cuts from natural subalpine coniferous forests in western Sichuan. To assess the natural regeneration potential of this species in spruce plantations of different ages, field studies on the seed rain, seed bank, and seedling recruitment were conducted in 20-year, 30-year, and 60-year plantations, and in a retained natural forest ca. 150 years old for comparison. Moreover, a series of temperature and light regimes were also designed in March 2003 to test germination/dormancy responses of the P. asperata seeds to different conditions. In the plantations considered, both the densities of seed rain and soil seed bank increased with increasing stand age, whereas they were both low beneath the natural forest partially due to low adult density of P. asperata adult trees. P. asperata has a transient seed bank, and ca. 60% of seeds were found in the litter layer. Seed decay and seed predation were the two most important factors affecting soil seed bank dynamics, which caused a number of seed losses from the soil. Only a small fraction germinated and produced seedlings, and the indices of the losses from the seed bank via germination in the 30-year plantation, 60-year plantation and natural forest were 4.75, 5.10, and 2.80%, respectively. High seedling mortality was observed after seedling emergence, and most of the germinated seedlings died out within one growing season. The P. asperata seeds showed a high percent germination and no significant differences under most light and temperature regimes except for full sunlight or temperature below 5°C. In conclusion, despite a substantial number of seed produced, the high depletion of soil seed, the low seedling output, and high seedling mortality may obscure the natural regeneration potential of this tree species.     

Similarity between seed bank and vegetation in Mediterranean grassland: a predictive model

Abstract. The similarity in species composition between seed bank and vegetation was analysed in Mediterranean grasslands in relation to altitude, topography and grazing. Soil samples were collected in permanent plots in autumn at the end of the summer drought period and in spring, before the new seed fall and after the natural winter seed stratification. The seed bank composition was determined by greenhouse germination over a nine-month period. Presence/absence of species in the standing vegetation throughout the complete annual cycle, and the percentage area of bare ground in October, were recorded in the same plots. The species composition of the standing vegetation is clearly determined by altitude, topography and grazing, while the floristic composition of the seed banks is only related to altitude and topography in the case of autumn seed bank and with any of the three factors in the spring seed bank. Relative abundances of grasses, legumes and forbs also show different patterns in vegetation and seed bank data. Sørensen similarity between the autumn seed bank and the vegetation declines as altitude rises, but there are no significant differences for topography and grazing. This similarity decreases in the case of the spring seed bank and does not show any significant relationship with any of the factors. The perennial/ annual ratio and the proportion of bare soil in October are proposed as explanatory variables in a predictive model of similarity between the seed composition of the seed bank and vegetation.     

亚高山草地土壤原生生物群落结构和多样性海拔分布格局

生物多样性分布格局与维持机制即群落构建机制,是群落生态学研究的热点领域。微生物生态学中的一个关键问题是量化确定性过程和随机过程对微生物群落构建的相对贡献。尽管原生生物是土壤微生物群落中重要的组成部分,在微生物食物链中扮演着关键角色,但与细菌和真菌相比,目前对原生生物群落构建机制知之甚少。运用Illumina Miseq高通量测序技术,分析了五台山亚高山草地生态系统（海拔2000-3061 m范围内）土壤原生生物群落组成和多样性维持机制。结果表明,四个海拔梯度的土壤共获得有效序列520673条,分属于8个超群、24个门、65个纲、125个目、222个科和350个属。门水平上,丝足虫门（Cercozoa）、褐藻门（Ochrophyta）、纤毛门（Ciliophora）和顶复门（Apicomplexa）为主要优势类群。LEfSe分析显示,17个生物标志物对海拔梯度非常敏感,不同海拔梯度富集了不同的原生生物种群。尽管海拔对土壤原生生物群落的α多样性没有显著影响,但非度量多维尺度分析（NMDS）和相似性分析（ANOSIM）结果表明,原生生物群落组成和结构在海拔梯度上存在显著的差异（P<0.05）。冗余分析（RDA）显示海拔、土壤含水量、总氮和植物丰富度指数与原生生物群落结构存在显著相关性（P<0.05）。方差分解分析（VPA）和偏Mantel分析表明环境因子和空间变量对原生生物群落海拔分布格局均有显著影响,但是环境因子的相对作用（7.9%）明显大于空间变量（1.8%）。土壤原生生物群落之间的Bray-Curtis距离与海拔距离呈显著正相关关系（P<0.05）,说明选择过程可能是亚高山草地土壤原生生物群落分布格局的主要驱动因素。零模型分析进一步证实了确定性过程在原生生物群落构建中的相对作用大于随机过程。总之,五台山亚高山草地土壤原生生物群落组成和结构沿海拔梯度具有显著的变化格局,群落构建主要由确定性过程和随机过程驱动,但确定性过程占主导地位。     

Aerial seed bank dynamics and seedling emergence patterns in two annual Mediterranean Asteraceae

Question: We explored the functional significance of seasonal aerial seed banks in two coexisting, heterocarpic annual Asteraceae with dormant (Chrysanthemum coronarium) and non‐dormant (Anacyclus radiatus) achenes. We hypothesised that the plant achene pool is a significant component of total seed reserves, and that within‐season seedling emergence timing is shaped by achene release patterns. Location: SW Spain. Methods: In an observational study, we established temporal achene release patterns. We also quantified the aerial and soil achene pools throughout the release season, and assessed seedling emergence timing. Sowing experiments were used to explore the influence of achene release dynamics on emergence timing, and to establish achene morph‐specific patterns of between‐year distribution of germination. Results: Achene release extended from late spring to late autumn (Chrysanthemum), or from early autumn to early winter (Anacyclus). Within species, achene morphs differed in release timing. Only in Chrysanthemum, a small achene fraction seemed to persist in the soil, and between‐year germination distribution differed among morphs. In coexisting populations, the Anacyclus plant achene pool was an order of magnitude higher than the soil pool throughout the release season, whereas in Chrysanthemum both pools were of the same magnitude during autumn. Within‐year seedling emergence was significantly staggered beneath parent plants compared with the pattern resulting solely from the germination response in soil, with the exception of Chrysanthemum in one of the two study years. Conclusions: Results suggest that seasonal aerial seed banks are effective within‐season, risk‐reducing traits in ruderal Mediterranean habitats characteristic of the study species.     

Contemporary pollen and seed dispersal in a Prunus mahaleb population: patterns in distance and direction

Pollination and seed dispersal determine the spatial pattern of gene flow in plant populations and, for those species relying on pollinators and frugivores as dispersal vectors, animal activity plays a key role in determining this spatial pattern. For these plant species, reported dispersal patterns are dominated by short-distance movements with a significant amount of immigration. However, the contribution of seed and pollen to the overall contemporary gene immigration is still poorly documented for most plant populations. In this study we investigated pollination and seed dispersal at two spatial scales in a local population of Prunus mahaleb (L.), a species pollinated by insects and dispersed by frugivorous vertebrates. First, we dissected the relative contribution of pollen and seed dispersal to gene immigration from other parts of the metapopulation. We found high levels of gene immigration (18.50%), due to frequent long distance seed dispersal events. Second, we assessed the distance and directionality for pollen and seed dispersal events within the local population. Pollen and seed movement patterns were non-random, with skewed distance distributions: pollen tended moved up to 548 m along an axis approaching the N-S direction, and seeds were dispersed up to 990 m, frequently along the SW and SE axes. Animal-mediated dispersal contributed significantly towards gene immigration into the local population and had a markedly nonrandom pattern within the local population. Our data suggest that animals can impose distinct spatial signatures in contemporary gene flow, with the potential to induce significant genetic structure at a local level.     

荒漠草原土壤种子库对草地管理方式的响应

以宁夏中部不同封育年限(封育1a、3a、5a和7a)、不同放牧方式(中等强度自由放牧、中等强度4区轮牧)、补播改良(2006年补播沙打旺Astragalus adsurgens和紫花苜蓿Alfalfa stiva)及未封育等管理方式下的荒漠草原为研究对象,采用空间梯度代替时间梯度法,于2012年3月下旬用样线法采集土壤样品,采用温室萌发法,统计土壤种子库物种组成和种子数,研究了不同草地管理方式下的种子库与地上植被相似性、多样性等特征。结果表明:1)宁夏荒漠草原土壤种子库中共出现14种植物,分属6科14属,除牛枝子(Lespedeza potanimii)为半灌木外其余均为草本植物。多年生植物占土壤种子库植物总数的65.3%,一年生植物占土壤种子库植物总数的35.7%;多年生植物种子数量随封育年限增加而表现为下降趋势,封育1a和未封育草地最多;2)不同管理方式下土壤种子库和地上植被的共有物种数为2—6种,封育时间延长导致土壤种子库物种与地上植被相似性下降,土壤种子库和地上植被共有物种数有减少趋势,自由放牧降低了土壤种子库和地上植被的共有物种数,补播恢复的草地土壤种子库中未出现补播物种;3)种子库物种多样性与管理方式无明显的相关性;种子库物种多样性指数4区轮牧草地最高,封育7a草地最低,物种丰富度指数封育1a草地最高,均匀度指数自由放牧草地最高,补播草地最低。若荒漠草原仅依靠土壤种子库恢复自然植被,则不建议封育时间过久,需间以适当的干扰(放牧);补播外来种未对荒漠草原土壤种子库产生贡献,反而对乡土种产生了一定限制。     

Aerial seed bank in a cold desert annual-ephemeral species: Role of anatomical structure of stem and delayed fruit dehiscence in timing of seed dispersal

Why the fruits are retained on dead upright herbaceous plants and how this relates to seed dispersal and timing of germination remain unclear. Stems of the annual Euclidium syriacum (Brassicaceae) with infructescences bearing indehiscent silicles remain upright after plants die in the spring. We investigated the effect of anatomical structures of stem and pedicle and delayed silicle dehiscence on seed dispersal phenology of this species. For comparison, sections were made of the stem of the annual Goldbachia laevigata (Brassicaceae), which has stems that fall over when plants die. Compared to G. laevigata, the stem of E. syriacum has vascular bundles that are closer together, a thicker xylem and phloem, more fibers, a thicker perimedullary zone and a smaller pith diameter:stem diameter ratio. The thickened pedicle did not form an abcission layer. By late October, 5–20% of seeds were dispersed, depending on the position of infructescences on the plant. Snow covered the plants in late autumn and when it melted in mid-April many of the plants had fallen over, with a high number of seeds germinating in attached silicles; seedlings became rooted in soil. After snowmelt, 14–15% of the silicles on the remaining upright plants contained seeds; all seeds were dispersed by early July. The anatomical structures of the stem and pedicle plus the delayed dehiscence of silicles explain the presence of an aerial seed bank in E. syriacum and delay of germination of many of seeds until spring. Further, pieces of upright plants are broken off and dispersed by wind, which helps to explain the wide distribution of E. syriacum in the cold desert.