Factors influencing the natural regeneration of the pioneering shrub Calligonum mongolicum in sand dune stabilization plantations in arid deserts of northwest China

Calligonum mongolicum is a successful pioneer shrub to combat desertification, which is widely used for vegetation restoration in the desert regions of northwest China. In order to reveal the limitations to natural regeneration of C. mongolicum by asexual and sexual reproduction, following the process of sand dune stabilization, we assessed clonal shoots, seedling emergence, soil seed bank density, and soil physical characteristics in mobile and stabilized sand dunes. Controlled field and pot experiments were also conducted to assess germination and seedling emergence in different dune soil types and seed burial depths. The population density of mature C. mongolicum was significantly different after sand dune stabilization. Juvenile density of C. mongolicm was much lower in stabilized sand dunes than mobile sand dune. There was no significant difference in soil seed bank density at three soil depths between mobile and stabilized sand dunes, while the emergence of seedlings in stabilized dunes was much lower than emergence in mobile dunes. There was no clonal propagation found in stabilized dunes, and very few C. mongolicum seedlings were established on stabilized sand dunes. Soil clay and silt content, air‐filled porosity, and soil surface compaction were significantly changed from mobile sand dune to stabilized dunes. Seedling emergence of C. mongolicm was highly dependent on soil physical condition. These results indicated that changes in soil physical condition limited clonal propagation and seedling emergence of C. mongolicum in stabilized sand dunes. Seed bank density was not a limiting factor; however, poor seedling establishment limited C. mongolicum's further natural regeneration in stabilized sand dunes. Therefore, clonal propagation may be the most important mode for population expansion in mobile sand dunes. As a pioneer species C. mongolicum is well adapted to propagate in mobile sand dune conditions, it appears unlikely to survive naturally in stabilized sand dune plantations.     

Ecophysiological behavior ofArtemisia ordosica on the process of sand dune fixation

The population structure and water relations ofArtemisia ordosica were studied at different stages of the sand dune fixation process. Vegetation coverage and biomass increased as the sand dune fixation process progressed. In contrast, individual growth rate decreased in the late fixation stage. On fixed sand dunes the modal age ofA. ordosica plants was higher and seedlings or saplings were rearely observed. On active sand dunes, settlement of seedlings was regulated by sand mobility; that is, seedlings were observed only when sand mobility was below 10 cm per year. Leaf transpiration was highest in active sand dunes. Stand transpiration in fixed sand dunes was lower than in semi-fixed sand dunes. These differences in transpiration were related to a decrease in soil water availability, which was affected by the increase in the aeolian fine soil component during the course of sand dune fixation. The reduction in soil water status and the cessation of sand movement were considered to be important factors in the decline of theArtemisia ordosica community.     

科尔沁沙地盐蒿种群的空间分布格局及关联性

以科尔沁沙地半固定沙丘和流动沙丘两种不同沙丘上的盐蒿(Artemisia halodendron)种群为研究对象,采用空间点格局方法研究了 0～20 m尺度上盐蒿种群的空间分布格局及其关联性.结果表明,半固定沙丘上盐蒿种群的数量远大于流动沙丘,不同生长发育阶段的种群结构呈偏正态分布,属于稳定型种群;流动沙丘上不同生长发...     

Effects of Sand Dune Stabilization on the Spatial Pattern of Artemisia ordosica Population in Mu Us Desert,Northwest China

Vegetation patterns are strongly influenced by sand mobility in desert ecosystems. However, little is known about the spatial patterns of Artemisia ordosica, a dominant shrub in the Mu Us desert of Northwest China, in relation to sand fixation. The aim of this study was to investigate and contrast the effects of sand dune stabilization on the population and spatial distribution of this desert shrub. Spatial autocorrelation, semi-variance analysis, and point-pattern analysis were used jointly in this study to investigate the spatial patterns of A. ordosica populations on dunes in Yanchi County of Ningxia, China. The results showed that the spatial autocorrelation and spatial heterogeneity declined gradually, and the distance between the clustered individuals shortened following sand dune fixation. Seedlings were more aggregated than adults in all stage of dune stabilization, and both were more aggregated on shifting sand dunes separately. Spatial associations of the seedlings with the adults were mostly positive at distances of 0–5 m in shifting sand dunes, and the spatial association changed from positive to neutral in semi-fixed sand dunes. The seedlings were spaced in an almost random pattern around the adults, and their distances from the adults did not seem to affect their locations in semi-fixed sand dunes. Furthermore, spatial associations of the seedlings with the adults were negative in the fixed sand dune. These findings demonstrate that sand stabilization is an important factor affecting the spatial patterns of A. ordosica populations in the Mu Us desert. These findings suggest that, strong association between individuals may be the mechanism to explain the spatial pattern formation at preliminary stage of dune fixation. Sand dune stabilization can change the spatial pattern of shrub population by weakening the spatial association between native shrub individuals, which may affect the development direction of desert shrubs.     

VARIATION IN THE BIG BLUESTEM (ANDROPOGON GERARDII)-SAND BLUESTEM (ANDROPOGON HALLII) COMPLEX ALONG A LOCAL DUNE/MEADOW GRADIENT IN THE NEBRASKA SANDHILLS

Twenty four quantitative characteristics of 95 specimens of the big bluestem (Andropogon gerardii Vitman)—sand bluestem (Andropogon hallii Hack.) complex collected from six topographic positions along a local dune to meadow topographic gradient in the eastern Nebraska Sandhills were examined to determine the relationship between morphological variation and habitat variation in adjacent bluestem populations. All morphological characters showed intergradation between big bluestem and sand bluestem but based on awn length, ligule length, cilia length on pedicels and rachises, rhizome length, and hybrid indexes, as well as comparisons with isolated bluestem populations, dune and meadow bluestems were found to be morphologically distinct. Sand bluestem types occupied dry, upland sand dunes, big bluestem types were restricted to adjacent, subirrigated meadows and plants intermediate in morphology were found only in narrow dune/meadow transition zones. For all bluestems, four characters (awn, ligule, cilia and rhizome) were significantly correlated with each other but correlations were most non-significant when only intermediates were considered; pollen stainability, however, was highly variable and was not related to morphology. Variation in leaf thickness, adaxial stomatal frequency and leaf absorptance to photosynthetically active radiation was found to exist in bluestems, which previously has not been quantitatively described in this complex. Habitat assortment in bluestem types along this dune to meadow topographic gradient was closely related to soil moisture levels which, in these sandy soils, were controlled primarily by the depth of the underlying water table.     

Dune slack restoration in Dutch mainland coastal dunes

The 45,000 ha Dutch dunes are relatively well protected but they suffer from severe desiccation. During the last decade a lot of knowledge has been obtained about the steering processes of eco-hydrology in dunes resulting in well-founded schemes to restore wet dune slack communities. In two case studies a large-scale restoration of the landscape was necessary. Since 1987 in the North-Holland Dune Reserve the groundwater catchment has been reduced (from 8 million m³ to 2 million m³ per year) and the resulting development monitored. In another case, large-scale intervention on 35 ha of the Amsterdam Water Supply Dunes in 1995 resulted in the restoration of active sand drift with opportunities for the establishment of pioneer vegetation. In small parts of the area, the surface has stabilised as a result of vegetation growth, but elsewhere drifting sand has spread and covered previously stable surfaces. Many wet to moist dune slacks developed and results relating to geomorphology, hydrology and vegetation are presented. Finally, in 1996 yet another project of 50 ha was carried out in the Meijendel dunes, in which infiltration ponds were removed to allow vegetation succession to develop from scratch. Results of these three case studies will be presented. If only hydrological restoration is carried out, sod-cutting was found to be necessary to restore the whole series of vegetation types of dune slacks. In two cases an area was denuded. The seed bank and surrounding slacks seem to play an important role in the success of restoration, this is especially true for the pioneer vegetation types. Future monitoring will prove whether critical (Red List) plant species will return.     

Arbuscular Mycorrhizae of Leymus arenarius on Coastal Sands and Reclamation Sites in Iceland and Response to Inoculation

The occurrence of arbuscular mycorrhizae fungi (AMF) was examined in natural and reclaimed sand dunes in Iceland. On the coastal sand‐plain of Myrdalssandur no spores of AMF were found on barren sand, and very few were found in 1‐ and 5‐year‐old reclamation sites of Leymus arenarius (lymegrass). A significantly higher number of AMF spores and root colonization were found in a 10‐year‐old reclamation site and in a natural old dune system of L. arenarius. AMF spores showed seasonal variation with higher occurrence in the fall than in the spring. On the volcanic island Surtsey, no AMF spores or root colonization were found on L. arenarius in 6‐ and 10‐year‐old dunes. However, AMF spores and root colonization were found in the 22‐year‐old Leymus dune. On Surtsey, a colonization pattern was found where AMF non‐dependent plants were the first colonizers followed by AMF facultative plants, then by AMF‐dependent plants. Today AMF facultative plants have the highest number of species but AMF non‐dependent plants have the largest population size. Three different AMF inocula were tested on L. arenarius; growth of seedlings was improved significantly by an indigenous AMF inoculum compared with commercial inocula. Inoculation of nursery grown seedlings of L. arenarius followed by transplantation into barren sand seeded with L. arenarius is recommended because this method will ensure establishment of the plant and enhance succession.     

沙丘坡面异质性小生境中准噶尔无叶豆对水分条件变化的响应

以多年生克隆植物准噶尔无叶豆(Eremosparton songoricum(Litv.)Vass.)为材料,选择河边(A种群)和沙漠腹地(B种群)两个沙丘,研究从沙丘底部至顶部,沿着水分条件连续变化的梯度,准噶尔无叶豆在分株种群和克隆片段水平的形态变化特征,以期能揭示其在异质性小生境内利用水分资源的对策,并为准噶尔无叶豆的资源保护、培育和利用提供有意义的参考.研究发现:①在分株种群水平,A种群分株高度及地上部生物量显著高于B种群,而B种群地下部(根)的生物量则显著高于A种群;②在克隆片段水平,随着沙丘底部至顶部,A种群与B种群克隆片段高度和地上生物量都减小,而分株密度都增加,但升高或降低的强度不同;A种群根的生物量和长度增加,主要是水平的位于地下0～10 cm层面的直径10mm以下的根长度增加,而B种群根的生物量减小,但长度却在增加,主要是水平的位于地下0～10 cm层面的直径6mm以下的细根长度增加.水平细根的长度增加,更利于无性系进行广泛觅食,同时促进无性系尽快越过不利生境斑块和提高分株在有利生境中的生长概率.结果表明,准噶尔无叶豆对沙丘坡面水分条件连续变化的异质性小生境存在分株种群及克隆片段两个等级的可塑性响应,并通过可塑性变化适应了沙丘坡面水分条件的分异.     

科尔沁沙地农田-沙丘交错区白草无性系的生态适应性分析

对生长在不同沙地生境下的白草无性系进行了个体生长与形态以及种群生物量等指标的比较研究,结果表明,生长在不同生境下的白草无性系表现出显著的形态可塑性和不同的生物量分配模式,生长在农田-沙丘过滤带上的白草无性系与生长在半固定沙丘上的白草无性系相比,具有更大的生物量,更快的叶片延伸速率,根茎总长度和节间长度更长,不定芽更多,地下生物量所占比例更大且分布更深;这些特征的出现与农田-沙丘过渡带上土壤表层比较疏松而下层水分状况相对优越有着密切关系,因而生长在农田-沙丘过渡带上的白草无性系能更快地拓展其生存空间,在固沙和沙生植被演替中起重要作用。     

Arbuscular mycorrhizae in a tropical sand dune ecosystem on the Gulf of Mexico

 Root samples of 37 species distributed on the beach and along a successional gradient (from mobile to stabilized areas) in a tropical sand dune system on the Gulf of Mexico showed that 97% of the species were mycorrhizal. The mycorrhizal inoculum potential of the sand from several dune areas was compared using two different bioassays. Firstly, the field rate of colonization by arbuscular mycorrhizal fungi of Chamaecrista chamaecristoides seedlings transplanted to random plots in the foredunes and in the mobile area was measured. The seedlings were harvested at intervals during 3 weeks to record mycorrhizal structures. In the mobile area, no mycorrhizal colonization was observed during the experiment. In the foredunes, hyphae and external mycelium were present in 40% of the seedlings as early as 8 days after transplanting. After 15 days, arbuscules and vesicles were observed in 60 and 20% of the seedlings, respectively, and after 21 days, 100, 46 and 20% of the seedlings showed hyphae, arbuscules and vesicles, respectively. Secondly, maize seedlings were transplanted to pots previously filled with sand from the foredunes, mobile dunes, grassland and a Dyphisa robinoides shrub area. After 1 month, the lowest mycorrhizal inoculum potential was recorded for the mobile dunes and the highest for the shrub area. As expected, mycorrhizal inoculum potential increased with dune stabilization. Accepted: 17 July 1996     

Diversity of AMF associated with Ammophila arenaria ssp. arundinacea in Portuguese sand dunes

Dune vegetation is essential for the formation and preservation of sand dunes and the protection of the coast line. Coastal sand dunes are harsh environments where arbuscular mycorrhizal fungi (AMF) play an important role in promoting plant establishment and growth. We present a study of the diversity of AMF associated with A. arenaria ssp. arundinacea in two locations of the Portuguese coast under a Mediterranean climate. These two locations were selected to compare a well-preserved dune system from a protected area with a degraded dune system from a public beach. AMF diversity was assessed mainly by cloning and sequencing of a fragment of the ribosomal SSU using the primer NS31 and AM1. Most of the 89 AMF clones obtained from the rhizosphere and roots of A. arenaria belonged to the genus Glomus, the largest clade within the Glomeromycota. Higher AMF diversity was found in the least disturbed site, in which spores of Scutellospora persica, Glomus constrictum and Glomus globiferum were found in the rhizosphere of A. arenaria.     

Temporal morph frequency changes in sand‐dune populations of Cepaea nemoralis (L.)

Changes in morph frequency over time in sand‐dune populations of Cepaea nemoralis in the British Isles have been examined using the Evolution Megalab database. Frequencies in colonies on a sand dune at Berrow, Somerset were estimated in 2008–9. This extends a survey started in 1926 by A. E. Boycott and C. Diver and continued in the mid‐20th Century by B. C. Clarke and J. J. Murray. An increase in the frequency of the mid‐banded morph, noted in earlier work, has continued. The apparent decrease in brown was not confirmed but the yellow frequency has increased. At a range of dunes in the British Isles, comparisons between data from the mid‐20th Century and the early 21st Century indicate an increase in yellow and mid‐banded morphs and a decrease in banding and brown morphs. These results differ from the overall trends derived from Europe‐wide comparison of early data with Evolution Megalab data, in which banded and mid‐banded increased in frequency, whereas there was no overall change in yellow. The general pattern of regional variation has been retained, although there is also high heterogeneity between samples, suggesting that a variety of factors are involved in explaining the changes. The mean shifts in frequency are consistent with climatic change over the period. Dunes are probably the most likely habitat in which to detect such a change. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 108 , 315–322.     

STIGMATIC POLLEN LOADS IN POPULATIONS OF PONTEDERIA CORDATA FROM THE SOUTHERN U.S.

Stigmatic pollen loads were examined in four tristylous populations of Pontederia cordata from the southern U.S. to evaluate Darwin's hypothesis that floral trimorphism promotes legitimate pollination. In each population, morph frequency, pollen production, the composition of stigmatic pollen loads and seed production were estimated. Goodness-of-fit tests and a measure of the efficiency of legitimate pollination were employed to evaluate pollen load patterns. Results from the four populations demonstrate that the long-styled (L) morph usually experiences legitimate pollination, whereas the mid-styled (M) and short-styled (S) morphs often display random pollination. However, at Taylor Road (Louisiana) all three floral morphs exhibited statistically significant levels of legitimate pollination. The size of legitimate pollen loads suggests that in populations of P. cordata tristyly may have only a minor influence on the fecundity of morphs. Two hypotheses are proposed to explain the maintenance of floral trimorphism in contemporary populations of Pontederia spp. The first assumes the polymorphism is selectively neutral; the second suggests that it contributes towards the male component of fitness by increasing the distance, and hence the number of genets, that pollen can be transported to by pollinators. In clonal species, such as Pontederia cordata, where geitonogamous pollinations are promoted by large floral displays, floral mechanisms that increase pollen carryover may have selective value.     

浑善达克沙地榆树疏林幼苗更新空间格局

榆树疏林广泛分布于浑善达克沙地,是适应半干旱、半湿润气候的沙地植被类型.采用Ripley的K函数统计方法,分析了浑善达克沙地固定沙丘与丘间低地封育状态下的榆树幼苗空间分布及更新格局.结果表明:两种生境下榆树幼苗密度分别为88株/hm2和77株/hm2,丘间低地更适合幼苗的生长.固定沙丘和丘间低地榆树幼苗均呈聚集分布,但丘间低地的幼苗聚集强度更高.固定沙丘上榆树幼苗与成熟树种群在小尺度上呈负关联,而丘间低地幼苗与成熟树种群呈现较显著的正关联,但二者在大尺度上均无关联性.研究对揭示浑善达克沙地榆树幼苗分布、更新空间格局及其恢复保护具有重要意义.     

入侵植物凤眼蓝繁育系统在中国境内的地理变异

 对中国境内的凤眼蓝（Eichhornia crassipes）进行了大规模地理范围的调查,分析了24个地点40个种群的克隆繁殖与有性繁殖的相对水平、三型花柱的花型变化以及花部综合特征的变异。40个种群都以克隆繁殖为主要更新方式。其中36个种群只有三型花柱中的M花型;南宁、重庆的种群中出现了频率极低的L花型;南宁、重庆、福州和海口种群内,出现了具有促进花内自交潜能的半同长花柱花型（柱头与花药相靠拢的M花型变异体或L花型变异体）。与其它地方种群相比,西南和海南岛的种群表现出较多样化的繁育系统。M和L花型个体的结实个体率都约为4％,没有显著差异,种子产量在两花型个体之间也没有差异。由于种群内花型的残缺,作为促进花型之间异交传粉适应机制的三型花柱繁育系统已经解体,部分种群中出现了可以促进花内自交的半同长花柱变异体。克隆繁殖是凤眼蓝适应入侵生境的一种形式,花内自交是凤眼蓝繁育系统在入侵过程中可能的进化方向,这些繁育系统上的变化对该植物的入侵机制和途径产生了深远影响。     

Clonal integration in Fragaria chiloensis differs between populations: ramets from grassland are selfish

In plants, only species with clonal growth are able to directly transfer resources between otherwise independent units of the same genetic individual. A simple conceptual model of plant performance as a function of internal resource supply and environmental resource availability suggests that resource sharing between ramets within clones is likely to be disadvantageous in uniform habitats and advantageous in patchy ones. It was therefore hypothesized that clones in populations from relatively uniform habitats will have been selected for low rates of resource sharing between ramets compared to clones in populations from patchier habitats. In coastal northern California, the clonal herb Fragaria chiloensis is common both in grasslands, where resources are relatively uniform, and on sand dunes, where resources are more patchy. It was predicted that clones from a grassland population of Fragaria would have “selfish” ramets with low rates of resource sharing compared to clones from an adjacent dune population. Ramets were subjected to contrasting light levels with and without connection between ramets. Patterns of biomass accumulation were consistent with the prediction. This appears to be the first report of genetically based variation in patterns of resource sharing in clonal plants. It supports the idea that these patterns are locally selected to increase plant performance in habitats with different patterns of resource availability. Received: 19 August 1998 / Accepted: 4 March 1999     

新月形沙丘稳定性机理——以甘肃河西沙区为例

新月形沙丘是一种重要的沙丘类型,一般存在于绿洲边缘。研究新月形沙丘的稳定性,对于揭示绿洲缘的风沙运动规律有重要学术价值,而到目前新月形沙丘的稳定性还是沙漠生态学领域中一个的尚未解决的科学问题。以河西沙区新月形沙丘为例,通过调查其分布环境、测定沙丘各部位风速和风蚀积沙等,分析了新月形沙丘顶部稳定机理。结果表明:1)甘肃河西沙区的新月形沙丘分布在沙漠边缘的下风向,一般为斑块状分布。分布区域为平坦的沙粘质或沙砾质滩地,丘间地较为开阔且具有明显的主风向。2)主风向(NW)过程是新月形沙丘顶点与沙脊线重合的过程,也是新月形沙丘前移和高度降低的过程;反主风向(SE)过程是新月形沙丘顶点与沙脊线分离的过程,也是新月形沙丘背风坡上部风蚀与沙丘增高的过程。3)新月形沙丘越高大,两翼越长、夹角越小。4)较为开阔的前后滩地是新月形沙丘及新月形沙丘链存在所必须的环境条件,相对单一的主风向是新月形沙丘及新月形沙丘链维持其稳定性的关键因子,反向风及两翼的廊道效应是新月形沙丘增高增大的关键因子,主、反两个方向的风速及其分布频率是控制该区域新月形沙丘及新月形沙丘链高度的限制因子。     

Reproductive consequences of interactions between clonal growth and sexual reproduction in Nymphoides peltata: a distylous aquatic plant

Distyly is a sexual polymorphism in which plant populations contain two floral morphs differing in morphology and physiology. The dimorphism serves to promote animal-mediated cross-pollination between the floral morphs. Clonal propagation can interfere with the functioning of distyly by compromising intermorph pollinations, resulting in reduced fertility. Here, we investigate the relations between clonal growth and sexual reproduction in the aquatic macrophyte Nymphoides peltata (Menyanthaceae). Surveys of morph representation in 30 populations from five regions of China revealed that most populations exhibited strongly biased morph ratios and 30% contained a single floral morph. Experimental pollinations indicate that N. peltata possesses a strong dimorphic incompatibility system preventing self and intramorph fertilizations. An experiment involving the manipulation of morph ratios in an experimental population and an investigation in a natural population with strong morph substructure both provided evidence that compatible pollen dilution limits fertility. Despite constraints on the functioning of distyly in N. peltata we found no evidence for evolutionary changes to the heterostylous syndrome, as reported in Nymphoides, including populations of N. peltata in other parts of its geographical range.     

Estimating the abundance of Long‐billed Curlews in Nebraska

ABSTRACT Long‐billed Curlews (Numenius americanus) are shorebirds of high conservation concern in North America. Populations have declined in the last 150 yr primarily due to habitat loss and conversion. We conducted a 2‐yr study to estimate the density and statewide abundance of breeding Long‐billed Curlews in Nebraska during 2008 and 2009. Surveys were conducted during the prenesting period in April when Long‐billed Curlews were likely to be detected. We used a simple random sample of roadside survey routes (N= 39), each consisting of 40 5‐min point‐counts at 800‐m intervals. We modeled detection probability and found that wind speed negatively affected detectability, but found no evidence of either year effects or observer bias. We estimated there were 0.0038 Long‐billed Curlews per hectare (0.38 Long‐billed Curlews per km²) along survey routes and, by extrapolation, estimated there were 23,909 (SE = 1685; 95% CI: 20,810—27,471) Long‐billed Curlews in Nebraska. Our population estimate suggests that ～15 to 22% of the United States population of Long‐billed Curlews is found in Nebraska. Curlews were not evenly distributed within our survey area, with the highest densities in the central Sandhills, an area dominated by grass‐stabilized sand dunes and shallow wetlands, and the lowest densities in predominantly agricultural areas. Because Long‐billed Curlews in Nebraska face many potential threats resulting from land‐use changes, monitoring of the breeding population should be continued. Our survey method was efficient and yielded interpretable results; other states or regions should consider adopting this approach for estimating curlew abundance.     

A comparison of seed banks across a sand dune successional gradient at Lake Michigan dunes (Indiana,USA)

In habitats where disturbance is frequent, seed banks are important for the regeneration of vegetation. Sand dune systems are dynamic habitats in which sand movement provides intermittent disturbance. As succession proceeds from bare sand to forest, the disturbance decreases. At Indiana Dunes National Lakeshore, we examined the seed banks of three habitat types across a successional gradient: foredunes, secondary dunes, and oak savanna. There were differences among the types of species that germinated from each of the habitats. The mean seed bank density increased across the successional gradient by habitat, from 376 to 433 to 968 seeds m⁻², but with foredune and secondary dune seed bank densities being significantly lower than the savanna seed bank density. The number of seeds germinated was significantly correlated with soil organic carbon, demonstrating for this primary successional sequence that seed density increases with stage and age. The seed bank had much lower species richness than that of the aboveground vegetation across all habitats. Among sites within a habitat type, the similarity of species germinated from the seed banks was very low, illustrating the variability of the seed bank even in similar habitat types. These results suggest that restoration of these habitats cannot rely on seed banks alone.