Structure, Pattern and Mechanisms of Formation of Seed Banks in Sand Dune Systems in Northeastern Inner Mongolia, China

A comparison of structure and pattern of the soil seed bank was made between active and stabilized sand dunes in northeastern Inner Mongolia, China. The objective of this paper was to determine the significance of seed bank in vegetation restoration of sand dunes. The results showed that (1) average seed density decreased from stabilized sand dune to interdune lowland of stabilized sand dune, to interdune lowland of active sand dune, and to active sand dune; (2) horizontally, along the transect from interdune lowland to ecotone and to sand dune top, a ‘V’ shaped pattern was presented in the active dune system, and a reverse ‘V’ shaped pattern in the stabilized sand dune system; (3) vertically, the proportion (accounting for the total seeds) of seeds found in 0–20 mm soil profile decreased from stabilized sand dune to interdune lowland of stabilized sand dune, to interdune lowland of active sand dune, and to active sand dune. The same order was also found in 20–50 mm and 50–100 mm soil profiles; (4) the Sokal and Sneath similarity indices in the species-composition between soil seed bank and above-ground vegetation were ranked as: the stabilized sand dune (24%) > the interdune lowland of active sand dune (21%) > the interdune lowland of stabilized sand dune (18%) > the active sand dune (5%); and (5) vegetation restoration of active sand dunes depends on the dispersal of seeds from nearby plant communities on the interdune lowlands. Much effort must be made to preserve the lowlands, as lowlands are the most important seed reservoir in the active sand dune field.     

Seed mucilage interacts with soil microbial community and physiochemical processes to affect seedling emergence on desert sand dunes

Seedling emergence is a critical stage in the establishment of desert plants. Soil microbes participate in plant growth and development, but information is lacking with regard to the role of microbes on seedling emergence. We applied the biocides (captan and streptomycin) to assess how seed mucilage interacts with soil microbial community and physiochemical processes to affect seedling emergence of Artemisia sphaerocephala on the desert sand dune. Fungal and bacterial community composition and diversity and fungal–bacterial interactions were changed by both captan and streptomycin. Mucilage increased soil enzyme activities and fungal–bacterial interactions. Highest seedling emergence occurred under streptomycin and mucilage treatment. Members of the phyla Firmicutes and Glomeromycota were the keystone species that improved A. sphaerocephala seedling emergence, by increasing resistance of young seedlings to drought and pathogen. Seed mucilage directly improved seedling emergence and indirectly interacted with the soil microbial community through strengthening fungal–bacterial interactions and providing favourable environment for soil enzymes to affect seedling emergence. Our study provides a comprehensive understanding of the regulatory mechanisms by which soil microbial community and seed mucilage interactively promote successful establishment of populations of desert plants on the barren and stressful sand dune.     

Habitat degradation,topography and rainfall variability interact to determine seed distribution and recruitment in a sand dune grassland

Question: Understanding the mechanisms underlying how habitat degradation, topography and rainfall variability interactively affect seed distribution and seedling recruitment is crucial for explaining plant community patterns and dynamics. Interactions between these major factors were studied together in a semiarid sand dune grassland. Location: Eastern Inner Mongolia, China. Methods: The study system used four sites of fixed, semifixed, semishifting and shifting sand dune grasslands, representing a gradient of habitat degradation. We investigated the density of germinable seeds deposited in the top 5 cm of soil and in situ seedling emergence (number of seedlings emerging early in the growing season) and establishment (number of plants recruited at the end of the growing season) at three topographic positions (dune top, windward and leeward sides) within each site over 2 years that differed in rainfall. Habitat characteristics (i.e. vegetation cover, plant species composition and diversity, soil moisture and nutrient availability and soil erodibility) of the four sites were also measured. Results: Habitat degradation (i.e. decreased vegetation cover and enhanced wind erosion rate) significantly reduced the size of the germinable soil seed bank. On average, germinable seed number from the high‐vegetation cover fixed dune was 36‐fold larger than the low‐vegetation cover shifting dune, and eight‐ and two‐fold larger, respectively, than the semishifting and semifixed dunes with intermediate vegetation cover. We observed within‐habitat variability in seed distribution, but among‐topographic position variation differed among habitats. Seedling recruitment showed large between‐year, and among‐ and within‐habitat variability, but these variations varied significantly depending on the response variables evaluated (i.e. initial seedling density, final plant density, emergence rate and recruitment rate). Path analysis revealed complex density‐dependent positive and negative, direct and indirect effects of germinable seed density and initial seedling density on recruitment, but the relative importance of these density‐dependent effects varied depending on habitat type and rainfall availability. Conclusion: Our results suggest that habitat degradation, microtopography and rainfall availability interact in shaping sand dune seed bank and plant community recruitment patterns and dynamics. Their effects were mainly mediated through changes in both the biotic and abiotic environment during the process of habitat deterioration.     

Aerial and soil seed banks enable populations of an annual species to cope with an unpredictable dune ecosystem

Background and Aims

Simultaneous formation of aerial and soil seed banks by a species provides a mechanism for population maintenance in unpredictable environments. Eolian activity greatly affects growth and regeneration of plants in a sand dune system, but we know little about the difference in the contributions of these two seed banks to population dynamics in sand dunes.

Methods

Seed release, germination, seedling emergence and survival of a desert annual, Agriophyllum squarrosum (Chenopodiaceae), inhabiting the Ordos Sandland in China, were determined in order to explore the different functions of the aerial and soil seed banks.

Key Results

The size of the aerial seed bank was higher than that of the soil seed bank throughout the growing season. Seed release was positively related to wind velocity. Compared with the soil seed bank, seed germination from the aerial seed bank was lower at low temperature (5/15 °C night/day) but higher in the light. Seedling emergence from the soil seed bank was earlier than that from the aerial seed bank. Early-emerged (15 April–15 May) seedlings died due to frost, but seedlings that emerged during the following months survived to reproduce successfully.

Conclusions

The timing of seed release and different germination behaviour resulted in a temporal heterogeneity of seedling emergence and establishment between the two seed banks. The study suggests that a bet-hedging strategy for the two seed banks enables A. squarrosum populations to cope successfully with the unpredictable desert environment.     

地中海沿岸沙丘种子大小对植物及其种子多度的影响

分析了地中海沿岸沙丘3种微生境(灌丛下、灌丛之间的开阔地、路径)以及整个沙丘生态系统的种子大小与土壤种子库中休眠种子的数量、土壤种子库中总种子的数量、地上植被中各个植物种的个体数量和幼苗数量、每种植物在小样方中的出现频率等方面之间的关系,也分析了种子尺寸(长、宽、高之和的平均值)与土壤种子的长久性、土壤种子库中休眠种子数量之间的关系;还分析了具有不同种子大小的植物在沙丘生态系统和各微生境的分布比率,验证了生活周期短的植物的种子在土壤中更长久和被干扰的微生境具有更多的休眠种子这种假说。结果发现,在地中海沿岸沙丘生态系统中,具有特别大的种子和特别小种子的植物种类都很少,不同植物种子的大小呈现近正态分布,但绝大多数植物的种子重量都不超过10 mg;微生境影响种子尺寸与种子长久性的关系,在灌丛下、开阔地区域等两个微生境以及整个的沙丘生态系统都呈现显著的正向关系(p<0 .0 5 ) ,而在路径这种关系不很明显;种子多度与植被物种出现频率呈现强烈的正相关(p<0 .0 5 ) ,没有发现种子大小与土壤种子库中休眠种子数量、种子库中总的种子数量、植被物种出现频率、植被的物种多度、幼苗植物多度等方面有显著的关系;具有中度大小种子(0 .1～10 mg)的植物在总的土壤种子库、休     

The role of reproductive phenology, seedling emergence and establishment of perennial Salix gordejevii in active sand dune fields

BACKGROUND AND AIMS: The function of sexual reproduction of perennials in restoration of vegetation of active dune fields frequently has been underestimated. The objective of this study was to evaluate the role of sexual reproduction of the perennial Salix gordejevii in the revegetation of active dunes. METHODS: Seedling emergence and establishment of S. gordejevii were examined both in controlled experiments (germination at different burial depths with different watering regimes) and in field observations in three dune slacks. The reproductive phenology and soil seed bank of S. gordejevii, the dynamics of soil moisture, the groundwater table and the landform level of three dune slacks were monitored. KEY RESULTS: Seeds of S. gordejevii began maturation on 1 May, and seed dispersal lasted from 8 May to 20 May. Seeds on the soil surface germinated significantly faster than those buried in soil (P<0.05). Seedling emergence was negatively correlated with landform level. When most seedlings emerged, there was a significantly positive correlation between soil moisture and seedling emergence (P<0.01). Rainfall was negatively correlated with seedling emergence. Seedling establishment was significantly and positively correlated with seedling emergence (P<0.05), and 72.3 % of the emergent seedlings were established at the end of the growing season. These results indicated that (a) seeds matured and dispersed before the rainy season; (b) seeds germinated as soon as they contacted a moist surface and relied more on soil moisture than on rainfall; and (c) more seedlings emerged at lower sampling points in dune slacks. CONCLUSIONS: In natural conditions, restoration of active sand dune fields generally commences with revegetation of dune slacks where sexual reproduction of perennials contributes greatly to species encroachment and colonization and hence plays an important role in restoration of active dune fields. Furthermore, aeolian erosion in dune slacks, leading to good soil moisture, facilitates seed germination, seedling emergence and establishment of S. gordejevii.     

石羊河中游沙漠化逆转过程土壤种子库的动态变化

在石羊河中游,应用空间代替时间的方法,选择流动沙丘以及封育恢复5a、15a和25a的沙漠化逆转过程序列,研究了沙漠化逆转过程土壤种子库的变化特征。结果表明:石羊河中游沙漠化土地土壤种子库由4科12种植物组成,种子库主要分布于表层0-5cm。在沙漠化逆转过程中,土壤种子库物种数趋于增加,物种组成以1年生草本植物占优势逐渐向多年生草本植物和半灌木植物转变;种子库密度、表层土壤种子比例、物种多样性指数、与地上植被的相似性呈现先增大后降低趋势;沙漠化土地生态恢复间隔的时间越长,土壤种子库间的相似性程度越低;沙漠化土地与地带性植被区种子库的相似性逐渐增大,但是恢复25a沙漠化土地也仅达到0.36。研究认为,沙漠化逆转过程也是沙漠化土地土壤种子库向地带性植被土壤种子库演变的过程,而且是一个十分缓慢的过程,该研究有助于丰富干旱区土壤种子库的理论和指导干旱内陆河流域沙漠化土地的生态恢复实践。     

Aerial seed bank in Artemisia species: how it responds to sand mobility

Aerial seed bank has been found in the arid dune ecosystem. However, the relationship of aerial seed bank to sand mobility intensity is unknown. The adaptation of aerial seed bank to sand mobility was analyzed in four Artemisia species dominating at different habitats in northeastern Inner Mongolia, China. The four species, Artemisia gmelinii, A. frigida, A. halodendron and A. wudanica, are an interdune lowland, stabilized dune, semi-stabilized dune and active dune species, respectively. Seed (achene and capitulum) persistence on the parent plants, the supplement for the soil seed bank, the maintenance of seed viability, and the effect of burial depth on seedling emergence of these four species were studied. Dispersal delay differed interspecifically responding to sand mobility intensity: maximal dispersal was in the following May after maturation for A. wudanica, in the following April after maturation for A. halodendron, and in November of the year of seed maturation for A. frigida and A. gmelinii. The maximal supplement to the soil seed bank for A. wudanica and A. halodendron occurred in the following May and March, respectively. The viability of achenes in aerial seed bank of A. wudanica and A. halodendron maintained at 79.5 and 79.6% until the following May after maturation. This study indicated that aerial seed bank in the sand dune ecosystem functions to postpone seed dispersal until the end of windy season and the start of growing season, and therefore is a mechanism for psammophyte to adapt to sand mobility.     

青海三江源地区风沙土养分及微生物区系

采用常规方法研究了青海三江源地区风沙土的养分状况及微生物区系.结果表明,从流动、半固定风沙土到固定风沙土的演化过程中,土壤有机质含量明显增加,固定风沙土的有机质含量分别为流动和半固定风沙土的5.9和3.8倍;土壤氮素和磷素含量的变化趋势与有机质基本一致,均呈递增趋势;土壤钾素含量和土壤pH无明显变化规律.随着植被发育、流沙固定及土壤养分状况改善,风沙土中的微生物数量和区系组成也发生了显著变化.固定和半固定风沙土中的细菌、真菌及放线菌数量均明显高于流动风沙土,其细菌数量分别约为流动风沙土的4.0和2.8倍,真菌数量分别约为19.6和6.3倍,放线菌数量分别约为12.4和2.6倍;真菌种类数明显增加,放线菌区系组成也变得复杂.即随着风沙土由流动变为固定,土壤微生物生态系统中微生物的生物多样性增强,微生物组成趋于多样化.风沙土中的微生物数量与土壤有机质、全氮、速效氮及速效磷含量显著或极显著相关,与土壤全磷、全钾、速效钾含量及土壤pH之间相关性未达显著水平.     

The potential role of seed banks in the recovery of dune ecosystems after removal of invasive plant species

Question: How resilient is the seed bank of an invaded dune system? Is that resilience dependent on duration of invasion? How does the accumulated litter layer contribute to the soil seed bank? Location: Coastal sand dunes invaded by Acacia longifolia, Portugal. Methods: Seedling emergence was used to quantify and compare soil seed banks in long‐invaded, recently invaded and non‐invaded areas. Changes in seed banks were also compared with areas where A. longifolia and the litter layer were removed. Results: Species richness, seedling density and diversity were higher in non‐invaded and recently‐invaded areas than in long‐invaded areas. Although there was an apparent similarity between non‐invaded and recently‐invaded areas, analyses of species traits revealed differences. Non‐invaded areas had a wider array of traits. Exotic/invasive species dominated invaded seed banks while native species dominated non‐invaded seed banks. Life forms, growth forms, longevity and dispersal mode showed differences between areas, with cleared plots of long‐invaded areas being apparently the most similar to non‐invaded plots. Acacia longifolia seeds were most abundant in long‐invaded areas, particularly where the litter layer remained. Removal of A. longifolia plus the litter had little effect on the seed bank composition of recently‐invaded areas but resulted in noticeable changes in seed banks of long‐invaded areas. Conclusions: Long‐invaded areas are less resilient and show a higher reinvasion potential, despite severe alteration of the seed banks of both areas. Seed bank studies can be a useful tool to guide management, but can give misleading results when invasion periods are protracted.     

地中海沿岸沙丘微生境对幼苗出现时空格局的影响

实验样地设在地中海沿岸沙丘,选择了3个不同的微生境代表：(1)稳定沙丘上的开阔地片段,(2)稳定沙丘上的灌丛下区域,(3)不稳定沙丘的路径区域;从2001年11月至2002年4月的整个生长季节,每一个微生境出现的幼苗在4个日期被监测,并在每一个取样日把每一株幼苗鉴定、计数后,用剪刀把地上部分移去;研究调查了3种微生境幼苗出现的时空分布格局,并分析了雨量与幼苗出现数量的关系。结果发现：在地中海沿岸沙丘生态系统,幼苗出现在时间上具有明显的分布特征,大多数幼苗出现在第一观测期,整个生长季幼苗都不断出现,但幼苗出现的数目却逐渐下降;各功能群的幼苗占幼苗总数的比例分别为：1年生阔叶草47．4％,多年生阔叶草2．5％,豆科植物17,0％,菊科植物14．5％,1年生禾草为11．7％,多年生禾草为1．9％,十字花科植物3．7％,伞形科植物1,4％。在空间上,总幼苗密度、物种丰富度和物种多样性等显示出重要的微生境差异,开阔地区域具有最大的幼苗密度、物种丰富度和物种多样性;3个微生境的幼苗出现不是同步的,微生境影响种子萌发的时间分布格局,灌丛下种子萌发具有滞后现象;大多数功能群的幼苗密度分布基本上具有显著的微生境差异,主要物种的幼苗分布也具有显著的微生境差异。雨量和萌发的幼苗数量间未发现显著的关系。     

沙生植物倒披针叶虫实在古尔班通古特沙漠的分布

倒披针叶虫实(Corispermum lehmannianum)在古尔班通古特沙漠广泛分布,重要值仅次于该沙漠建群种白梭梭(Haloxylon persicum),对沙面的稳定起着非常重要的作用。采用固定样方法对自然条件下该植物的分布特征和种群数量动态进行了研究。结果表明:1)倒披针叶虫实在沙丘各坡位均占有较高重要值,尤其活化程度较高的中部和上部优势度更明显;2)生长季节的不同阶段,除坡顶外,其他各坡位种子主要分3批萌发出苗,不同时期种子萌发出苗数量存在差异,3月份有大量种子萌发出苗,4月份种子萌发出苗数最多,5月份萌发出苗量逐渐减少。而坡顶3月份种子萌发出苗量最多,4月至5月均无种子萌发出苗。因此,除坡顶外,种子萌发出苗数量和种群密度在其他各坡位呈先增后减的趋势。生长季结束时各坡位种群密度表现为:中部和上部底部和丘间顶部;3)成熟植株单位面积种子产量为:中部和上部丘间和底部顶部,且中部和上部单位面积种子数量极显著多于丘间、坡底和坡顶(P0.01);4)通过对0—5 cm和6—10 cm土壤种子库密度研究发现,沙丘中部和上部种子库密度显著高于其他坡位。综合以上研究表明,倒披针叶虫实在沙面较活化的中部和上部呈优势分布,对沙漠环境具有良好适应性。因此,该植物是一种极具开发潜力的沙生植物。     

雅鲁藏布江高寒河谷流动沙地适生植物种筛选和恢复效果

通过4a的人工模拟飞播试验研究(2008—2011年),观测了北方优良沙生植物种和西藏乡土沙生植物种在雅鲁藏布江中游高寒河谷流动沙地上的出苗、保存、生长和繁殖情况,筛选和确定了西藏高寒流动沙地人工模拟飞播的适生植物种。结果表明:籽蒿、花棒、沙拐枣、杨柴和砂生槐在高寒河谷流动沙地的适应性较好。籽蒿在播后第2年便有花序和种子出现,花棒和沙拐枣在第3年开花结实,籽蒿、花棒和沙拐枣均能完成生活史。籽蒿的再繁殖能力较弱,花棒和沙拐枣的再繁殖能力较强。不同类型沙丘的形态特征对人工模拟飞播效果影响较大,最适宜型(第Ⅰ类)流动沙地的人工模拟飞播效果最好,植被盖度达35%;较适宜型(第Ⅱ类)流动沙地的效果较好,植被盖度达30%;不适宜型(第Ⅲ类)流动沙地的效果最差,植被盖度尚不足10%;基本适宜型(第Ⅳ类)流动沙地的效果较差,但采用沿等高线人工脚踩回头撒播法,植被盖度可达20%。不同沙丘部位的人工模拟飞播效果差异明显。     

Spatiotemporal pattern of seed bank in the annual psammophyte Agriophyllum squarrosum Moq. (Chenopodiaceae) on the active sand dunes of northeastern Inner Mongolia, China

The seed banks of active sand dunes are critical for the adaptation of psammophytes to unstable habitats. However, existing reports have failed to effectively elucidate the spatiotemporal structure of these seed banks and the factors regulating them. The seed bank pattern of the annual psammophyte Agriophyllum squarrosum Moq. in active sand dunes was studied by taking into consideration the time, the depth of the soil profile, dune positions, and seed bank type (released seed bank, belowground and aboveground canopy seed banks). Most canopy-stored seeds were unburied before the following March of the maturation year; however, they were buried from March to May. The seed bank mainly comprised the aboveground canopy-stored seeds from September of the year of seed maturation to the following March. The seed bank was concentrated in the surface soil layers (0–10 cm) from March of the year following seed maturation to June and was concentrated in the deep soil layers (10–50 cm) after June. The seed bank in the lower leeward slope tended to be larger than those in the other dune positions. The canopy-stored seed:released seed ratio differed between dune positions. The findings of this study are: (1) sand burial and wind erosion play key roles in shaping the spatiotemporal pattern of the seed banks of active sand dunes, and (2) delayed seed release may be a major factor regulating the spatiotemporal pattern of the seed banks of certain psammophytes.     

The chemical nature of P accumulation in agricultural soils��implications for fertiliser management and design: an Australian perspective

Sand burial, persistent seed bank and soil water content (SWC) are three factors that potentially can affect regeneration in sand dune plant populations. To evaluate the effects of these three factors on population regeneration of Eremosparton songoricum, a rare and endangered legume, we investigated seed germination, seedling emergence and seedling survival in greenhouse and controlled field experiments in different sand dunes microsites. Freshly matured seeds are physically dormant, and the highest germination was only 9.3?±?5.8% at 25/10°C. Seed germination occurred at burial depths from 0 to 10 cm, but the maximum depth from which seedlings emerged was 6 cm; from 1 to 6 cm, the deeper the burial, the lower the percentage of seedling emergence. Only 36.2% of the total soil seed banks occurred at depths of 0?C6 cm. For artificially sown seeds at different dune microsites, mean seedling emergence percentage was 6.8%. Of 150 seedlings that emerged in the field investigation at the study site, only those germinating in flat sandy areas survived, and mean survival percentage was only 2.0%. Thus, the proportion of non-dormant seeds in soil seed banks that developed into seedlings and survived to the end of the growing season was only 0.2%. Minimum SWC for seed germination, seedling emergence and seedling survival was 2.0%. During monitoring of emergent seedlings in the field, low seedling recruitment was at least partly due to the rate of root extension (1.6?±?0.3 cm day^?1) into the sandy soil, which was slower than that of the downward movement of plant-available moisture (2.8?±?0.6 cm day^?1). Thus, population regeneration under natural conditions rarely occurred via sexual reproduction, owing to the limited water resources available for seedling establishment. Rational field seeding practices, including manually scarified or dry stored seeds before sowing, sowing the seeds at right time and suitable place, are suggested for ecological restoration of endangered E. songoricum populations.     

Soil Moisture Rather than Soil Nutrient Regulates the Belowground Bud Bank of Rhizomatous Species <i>Psammochloa villosa</i> in Arid Sand Dunes

In arid and semi-arid sand dune ecosystems, belowground bud bank plays an important role in population regeneration and vegetation restoration. However, the responses of belowground bud bank size and composition to sand burial and its induced changes in soil environmental factors have been rarely studied. In arid sand dunes of Northwestern China, we investigated belowground bud bank size and composition of the typical rhizomatous psammophyte Psammochloa villosa as well as three key soil environmental factors (soil moisture, total carbon and total nitrogen) under different depths of sand burial. Total buds and rhizome buds increased significantly with increasing burial depth, whereas tiller buds first increased and then decreased, with a peak value at the depth of 20–30 cm. Soil moisture increased significantly with sand burial depth, and was positively correlated with the number of all buds and rhizome buds. Soil total carbon concentration first increased and then decreased with sand burial depth, and total nitrogen concentration was significantly lower under deep sand burial than those at shallow depths, and only the number of tiller buds was positively correlated with soil total nitrogen concentration. These results indicate that soil moisture rather than soil nutrient might regulate the belowground bud bank of P. villosa, and that clonal psammophytes could regulate their belowground bud bank in response to sand burial and the most important environmental stress (i.e., soil moisture). These responses, as the key adaptive strategy, may ensure clonal plant population regeneration and vegetation restoration in arid sand dunes.     

Ecological longevity of Polaskia chende (Cactaceae) seeds in the soil seed bank,seedling emergence and survival

• Soil seed banks are essential elements of plant population dynamics, enabling species to maintain genetic variability, withstand periods of adversity and persist over time, including for cactus species. However knowledge of the soil seed bank in cacti is scanty. In this study, over a 5‐year period we studied the seed bank dynamics, seedling emergence and nurse plant facilitation of Polaskia chende, an endemic columnar cactus of central Mexico.
• P. chende seeds were collected for a wild population in Puebla, Mexico. Freshly collected seeds were sown at 25 °C and 12‐h photoperiod under white light, far‐red light and darkness. The collected seeds were divided in two lots, the first was stored in the laboratory and the second was use to bury seeds in open areas and beneath a shrub canopy. Seeds were exhumed periodically over 5 years. At the same time seeds were sown in open areas and beneath shrub canopies; seedling emergence and survival were recorded over different periods of time for 5 years.
• The species forms long‐term persistent soil seed banks. The timing of seedling emergence via germination in the field was regulated by interaction between light, temperature and soil moisture. Seeds entered secondary dormancy at specific times according to the expression of environmental factors, demonstrating irregular dormancy cycling.
• Seedling survival of P. chende was improved under Acacia constricta nurse plants. Finally, plant facilitation affected the soil seed bank dynamics as it promoted the formation of a soil seed bank, but not its persistence.

     

Limiting factors for seedling emergence and establishment of the invasive non-native Rosa rugosa in a coastal dune system

The relative importance of seed, habitat and microhabitat limitation has rarely been investigated for invasive non-native species, although this is critical for their effective management and for predicting future range expansion. Rosa rugosa is an abundant non-native shrub in coastal habitats of NW Europe; it is common along the Baltic coast but more scarce in exposed dunes of the North Sea. To investigate whether invasion of exposed dunes is limited by seed, habitat and microhabitat limitation, seedling emergence and establishment were examined in a factorial sowing, transplant and disturbance experiment. Twenty plots were randomly placed in each of five dune habitats (white dune, Empetrum dune, grey dune, outer dune heath, inner dune heath), and studied over 2 years. Seedling emergence in control subplots was zero in all habitats, whereas 96% and 98% of the undisturbed and disturbed seeded subplots produced seedlings. Disturbance had a positive effect on emergence and subsequent survival in white dune, outer and inner dune heath. Seedling survival and growth, and growth of transplanted seedlings, were markedly lower in grey dune. These findings indicate that establishment of R. rugosa is seed-limited in coastal dune habitats, and that the species is able to establish in both active and fixed dunes once seeds have arrived. Although differential seedling emergence and growth indicate that habitats differ in their degree of invasion susceptibility by R. rugosa, the positive influence of small-scale disturbance suggests microhabitat limitation in some dune habitats as well. Dune management should aim to reduce seed production and dispersal of R. rugosa in near-natural sites, and anthropogenic changes of habitat dynamics should be prevented.     

Exotic Ecosystem Engineers Change the Emergence of Plants from the Seed Bank of a Deciduous Forest

The anthropogenic spread of exotic ecosystem engineers profoundly impacts native ecosystems. Exotic earthworms were shown to alter plant community composition of the understory of deciduous forests previously devoid of earthworms. We investigated the effect of two exotic earthworm species (Lumbricus terrestris L. and Octolasion tyrtaeum Savigny) belonging to different ecological groups (anecic and endogeic) on the emergence of plants from the seed bank of a northern North American deciduous forest using the seedling emergence method. We hypothesized that (1) exotic earthworms change the seedling emergence from the plant seed bank, (2) L. terrestris increases the emergence of plant seedlings of the deeper soil layer but decreases that of the upper soil layer due to plant seed burial, and (3) O. tyrtaeum decreases plant seedling emergence due the damage of plant seeds. Indeed, exotic earthworms altered the emergence of plant seedlings from the seed bank and the functional composition of the established plant seedlings. Surprisingly, although L. terrestris only marginally affected seedling emergence, O. tyrtaeum changed the emergence of native plant species from the seed bank considerably. In particular, the number of emerging grass and herb seedlings were increased in the presence of O. tyrtaeum in both soil layers. Moreover, the impacts of earthworms depended on the identity of plant functional groups; herb species benefited, whereas legumes suffered from the presence of exotic earthworms. The results highlight the strong effect of invasive belowground ecosystem engineers on aboveground ecosystem characteristics and suggest fundamental changes of ecosystems by human-spread earthworm species.     

Smoke and heat: influence on seedling emergence from the germinable soil seed bank of mesic grassland in South Africa

We examined if germination and seedling emergence of species from the soil seed bank of mesic grassland in South Africa differed in their response to smoke or heat treatments alone or combined. Soil seed bank samples taken from 0 to 5 cm depth of the topsoil were treated with smoke-water solution, heat and heat + smoke-water combined and subsequent emergence of seedlings monitored over 90 days. In total, 790 individuals from 11 different plant families representing 37 graminoid (Poaceae) and non-graminoid species of flowering plants were identified. The most abundant plant families that emerged were Poaceae followed by Asteraceae. Grasses contributed 18% of the germinable seed bank, while non-grass (forbs and trees) species contributed >80%. The most abundant grass species that emerged was Themeda triandra and the most common forb species was Centella asiatica. Compared to the control, smoke-water (SW) and/or smoke-water + heat (H + SW) treatment resulted in a 1.6-fold increase in mean number of seedling emergence. SW and H + SW treatment of soil seed bank also resulted in fourfold to fivefold greater biomass production. These results suggest that emergence of some smoke-responsive perennial grasses (e.g. Themeda triandra) from the soil seed bank can be enhanced using smoke techniques. Moreover, plant-derived smoke could potentially have a significant role in the restoration of degraded grasslands in South Africa and elsewhere.