High genetic and morphological diversity despite range contraction in the diploid Hieracium eriophorum (Asteraceae) endemic to the coastal sand dunes of south‐west France

Endemic plants inhabiting coastal sand dunes show augmented extinction risks due to the dynamic nature of dunes and strong human pressure on coastal areas. To investigate the survival strategies and threats to long‐term survival of such species, we combined genetic, morphological and biogeographical approaches, using the example of Hieracium eriophorum (Asteraceae) and its putative cryptic sister species H. prostratum, which are endemic to the longest coastal sand dune in Europe. An analysis of amplified fragment length polymorphism revealed high within‐population genetic variability, and slight isolation by distance was the only indication of genetic population structure. Thus, no signs of genetic threats to survival were found. Furthermore, genetic and morphometric data provided no evidence for the existence of two species. Therefore, we propose to synonymize H. prostratum with H. eriophorum and provide a nomenclatural overview with typification. Finally, an analysis of historical distribution records showed that, during the last 100 years, the species was lost from its range margins, where its habitat became fragmented. Taken together, our results suggest that one successful survival strategy of narrow endemics may be the achievement of large local population sizes on a small geographical scale, thus avoiding the genetic problems inherent to small and fragmented populations. Dune management policies should thus take care that the current tendencies to allow more erosion will not result in too severe fragmentation of the remaining continuous stretches of dune habitat. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 169 , 365–377.     

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.     

Plant functional traits and diversity in sand dune ecosystems across different biogeographic regions

Plant species of a functional group respond similarly to environmental pressures and may be expected to act similarly on ecosystem processes and habitat properties. However, feasibility and applicability of functional groups in ecosystems across very different climatic regions have not yet been studied. In our approach we specified the functional groups in sand dune ecosystems of the Mediterranean, Hyrcanian and Irano-Turanian phytogeographic regions. We examined whether functional groups are more influenced by region or rather by habitat characteristics, and identified trait syndromes associated with common habitat types in sand dunes (mobile dunes, stabilized dunes, salt marshes, semi-wet sands, disturbed habitats). A database of 14 traits, 309 species and 314 relevés was examined and trait-species, trait-plot and species-plot matrices were built. Cluster analysis revealed similar plant functional groups in sand dune ecosystems across regions of very different species composition and climate. Specifically, our study showed that plant traits in sand dune ecosystems are grouped reflecting habitat affiliation rather than region and species pool. Environmental factors and constraints such as sand mobility, soil salinity, water availability, nutrient status and disturbance are more important for the occurrence and distribution of plant functional groups than regional belonging. Each habitat is shown to be equipped with specific functional groups and can be described by specific sets of traits. In restoration ecology the completeness of functional groups and traits in a site may serve as a guideline for maintaining or restoring the habitat.     

Craniodental indicators of prey size preference in the Felidae

In the present study, we used linear morphometrics of the crania, mandible and dentition to explore the association between craniodental shape and prey size among 35 species of living felids. To accomplish this, felids were divided into three prey-size groups: (1) large prey specialists; (2) small prey specialists; and (3) mixed prey feeders. From these linear measurements, large prey specialist felids can be distinguished from small and mixed prey feeders by their relatively robust canines and incisors and relatively wide muzzles. These cranial characters are advantageous when dispatching large prey, due to the stranglehold that cats employ during this activity. Robust canines resist the bending and torsional forces applied by struggling prey and a wider muzzle helps to stabilize grip and distribute bite forces more evenly during the killing bite. Small prey specialists had smaller canines, narrower muzzles and slightly longer jaws for a speed advantage when catching small, quick prey. Mixed prey feeders were intermediate between large and small prey specialists, indicating they are adapted to killing both sizes of prey. Given the success of this ecomorphological analysis of living felids that specialize on different prey sizes, we look forward to applying this same approach to extinct species.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 784–799.     

Tracks and trackways of Myotragus balearicus Bate (Artiodactyla, Caprinae) in Pleistocene aeolianites from Mallorca (Balearic Islands, Western Mediterranean)

Tracks and trackways of the ruminant goat, Myotragus balearicus Bate, 1909 are abundant in the Late Pleistocene aeolianites of the south coast of Mallorca, Spain. The carbonate aeolianites form impressive dunes and related sand ramps that were initiated as echo dunes in front of a 20–40 m high paleocliff of Miocene reefal limestone. Where the dunes built up to the cliff top, they covered it as an aeolian ramp. The dunes and sand ramps were active during the dry summer months but were passive during the wet winter period. The small, goat-like animals were abundant on the island, and for unknown reasons crossed the aeolian accumulations in large numbers. This study deals with the tracks formed in the dune deposits. All preserved trackways indicate impression into moist sand. Special features of the tracks include the structure produced by the withdrawal of the foot, here called the ‘axis’, and a disturbance zone of plastic deformation around this. On dune crests, the disturbance zone surrounds the axis more or less symmetrically. However, in addition, a ‘pressure pad’ of dislocated, slightly rotated sediment bound by curved microfaults is commonly produced posterior to the axis by propulsive pressure of the foot. On steep stoss and lee slopes, the pressure pad becomes oriented in a down-slope position as a result of gravitational slip of the walking animal. Combination of disturbance of the sediment in this way by manus followed by overprinting of similar disturbance by pes produces highly complicated track structure. This structure may be characteristic of artiodactylous mammals in soft sand, particularly aeolian deposits. The structure is designated as Bifidipes aeolis isp. nov.     

Effects of long‐term grazing management on sand dune vegetation of high conservation interest

Question: Can long‐term grazing management maintain and restore species‐rich sand dune plant communities within a sand dune site of high conservation interest? Location: Newborough Warren, North Wales, UK. Methods: Vegetation changes that occurred between 1987 and 2003, subsequent to grazing by domestic livestock being introduced to the site after decades with little or no stock grazing, were analysed using data collected from permanent monitoring quadrats over a 16‐year period. Results: At the plant community level, grazing brought about a shift from a tall‐grass dominated, species‐poor community to a more species‐rich community in the dry dunes, but did not change community type in dune slacks. However, at the species level, grazing enhanced the abundance of some desired perennial, annual and biennial species, graminoids and bryophytes in both habitat types. The increased frequency of positive indicator species for habitat condition suggests that grazing was beneficial for species of conservation interest. Ellenberg nitrogen (N) values decreased after grazing in dry habitats but showed no long‐term change independent of grazing, suggesting no increase in site fertility over the study period. Surprisingly, light (L) values also decreased in the dry dunes after grazing. Conclusions: Long‐term grazing management can play an important role for the conservation of dune communities and associated species. Because of its positive effects on species diversity, plant communities and habitat condition in sand dunes, livestock grazing is recommended for conservation management.     

If you build it,will they come? Use of restored dunes by beach mice

Restoration of coastal habitat fragmented, degraded, or destroyed by development and climate‐related processes such as sea level rise and storm surge usually involves planting native plants to restore habitat structure, but whether and how restored areas benefit taxa other than plants is rarely reported. Installing restoration plantings is one method used to build habitat such as beach dunes where dunes have been lost, potentially creating habitat for dune‐dependent species. We compared use of natural vegetated dunes, open sand gaps, and restoration plantings (habitat treatment) by Perdido Key beach mice (Peromyscus polionotus trissyllepsis) over 3 years using tracking tubes to assess the value of restoration plantings for beach mice. Tubes were monitored in two seasons (early and mid‐summer), and under new and full moon conditions. Mice used restoration plantings less than natural vegetated dunes but more than open sand gaps, which suggests restoration plantings may facilitate movement of mice across fragmented areas. Both season and moon phase influenced the effect of habitat treatment, interactions which may be attributable to perceived risk associated with movement under a combination of different conditions of ambient light, vegetation cover, and habitat novelty. Our results show restoration plantings provide habitat for movement and foraging, and may ameliorate some consequences of sea level rise and storms for beach mice and potentially other dune‐dependent species into the future.     

沙丘微地形生境蚁丘分布特征及其对恢复过程的响应

为了阐明沙丘微地形生境蚁丘特征及分布规律对沙丘固定过程的响应规律,以科尔沁沙地为研究区,选择不同固沙阶段沙丘的4个方位（西北、东南、西南、东北）布设调查样地,测定了不同样地的蚁丘密度、直径和高度,分析了不同样地蚁丘特征及空间分布规律。结果表明：（1）流动沙丘背风向（即东南方位）蚁丘密度显著高于其他方位,但从流动沙丘到半流动沙丘,背风向（即东南方位）蚁丘密度显著低于其他方位;从半固定沙丘开始,半固定和固定沙丘微地形对蚁丘密度分布的影响较小。（2）在流动沙丘、半流动沙丘、半固定沙丘,不同微地形生境中蚁丘直径和高度均呈现出相似的分布规律,即东南方位均小于其他方位。在固定沙丘,蚁丘高度表现为东南方位亦较低,但蚁丘直径表现为西南和东北方位显著小于西北方位。（3）在流动沙丘,西北、西南和东南方位上蚁丘空间分布均表现为均匀分布型,而在半流动、半固定和固定沙丘,4个方位上蚁丘空间分布均表现为聚集分布型。（4）相关性分析表明,不同固沙阶段不同微生境下蚁丘密度、高度、直径与植物密度、多样性间的相关性不同。随着固沙阶段演替,不同微生境下蚁丘特征与植物密度、多样性的相关性增强。研究表明,在科尔沁沙地,随着流动沙丘固定和生境逐渐恢复,沙丘不同空间方位上蚁丘密度分布差异缩小,而蚁丘直径和高度仍存在沙丘微地形间显著差异性,并且蚁丘空间方位上均呈现聚集分布状态。     

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.     

科尔沁沙地植物群落圆环状分布成因地统计学分析

调查发现,在科尔沁沙地中部部分梁窝状沙丘的丘顶、丘坡和丘间地,植物群落依生境梯度而分布,呈典型圆环状分布格局．沙层水分状况分析表明,同一沙丘不同部位的沙层含水量为丘顶低于丘坡地,且二者均低于丘间地．地统计学分析表明,无论是流动沙丘还是固定沙丘,从丘顶、丘坡到丘间地,沙层含水量的空间异质性呈规律性的变化,块金值与基台值之比逐渐减小,变程逐渐减小,分维数逐渐增大,表明随机部分引起的空间异质性逐渐降低,而由空间自相关部分引起的空间异质性逐渐增高．因此,沙丘不同部位沙层含水量及其空间异质性的有规律变化是科尔沁沙地中部部分梁窝状沙丘植物群落呈圆环状分布格局的一个重要原因．     

Conservation management of coastal dunes for Orthoptera has to consider oviposition and nymphal preferences

Dune and heathland ecosystems can harbour a multitude of specialized insect species. To assess the habitat quality of these ecosystems, the presence of specialized insect species may act as a useful indicator. The Orthoptera species Myrmeleotettix maculatus, Decticus verrucivorus, and Platycleis albopunctata present such umbrella species. Because knowledge of oviposition and nymphal habitats of these species is insufficient, we conducted a combined study consisting of an outdoor oviposition experiment and a field survey analyzing nymphal habitat preferences during summer 2009 on the Baltic island of Hiddensee, Germany. The oviposition experiment showed, that all three species mostly avoid oviposition under lichens (= mature grey dunes). M. maculatus preferred bare ground for oviposition, D. verrucivorus favoured both bare ground and mosses, and P. albopunctata laid most eggs into mosses. Young nymphs of both M. maculatus and P. albopunctata preferred initial grey dunes with a high proportion of bare ground and moss-rich grey dunes. Old nymphs were related to moss-rich and lichen-rich grey dunes with more dense vegetation. Based on our results, early seral stages of dune succession with bare ground and mosses as keystone structures are crucial for the conservation of the three studied umbrella species. Because old nymphs and adults additionally require more dense grey dune vegetation or adjacent heath stands, practical dune and heathland management measures should aim to maintain a mosaic-like pattern of different grey dune and dwarf-shrub vegetation stands.     

栖息地荒漠化对草原沙蜥食性的影响

栖息地退化会使环境中蜥蜴的可利用食物资源发生改变。采用剖胃法对内蒙古自治区鄂尔多斯市准格尔旗和达拉特旗地区采集的草原沙蜥样本进行了食性分析。对固定沙丘、半固定沙丘和流动沙丘三类样地中77个实胃样本的652只(套)食物进行鉴定和分析的结果表明,这些食物主要为节肢动物,分为35类,隶属3纲11目32科;也在少部分个体(4.69%)的胃容物中发现少量(1.95%)植物碎片。上述三类栖息地中,草原沙蜥对蚁科、蚜科、瓢虫科、叶蝉科和茧蜂科昆虫均有较大的捕食比例;但相比固定沙区栖息地,草原沙蜥在半固定及流动沙丘中增减的食物种类均比较多,导致草原沙蜥在流动沙区与其他两类样地之间有较大的食物相异性。从固定沙丘、半固定沙丘到流动沙丘,随栖息地荒漠化程度的增加,草原沙蜥捕食食物类群的丰富度逐步减少,辛普森优势度指数相应地下降,但营养生态位宽度、香农威纳多样性指数和皮洛均匀性指数却在流动沙丘栖息地上明显增大。这表明鄂尔多斯地区的草原沙蜥是以捕食昆虫等节肢动物为主、兼食极少量植物的杂食性动物。栖息地荒漠化所致的环境变化显著地影响草原沙蜥的食物组成,使其觅食的食物种类逐步减少。     

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.     

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.     

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.     

沙丘生态系统种子库研究现状、趋势与挑战

风蚀和沙埋赋予沙丘生态系统独特的自然物理和生物过程。沙丘生态系统种子库研究对于沙区植物多样性保护和生态恢复至关重要。但是人们对沙丘生态系统种子库的时空格局、形成机制和生态功能的认识还很肤浅。近些年, 沙丘生态系统种子库开始受到关注。该文对20年来沙丘生态系统种子库研究进行了回顾总结, 论述了沙丘生态系统种子库研究现状, 剖析了沙丘生态系统种子库现有结论, 阐述了沙丘生态系统种子库研究趋势与面临的挑战。沙丘生态系统种子库研究已获得以下认识: 1)沙丘生态系统类型、部位和深度决定种子库大小与组成; 2)沙丘生态系统种子库季节、年际变化受风沙活动和降水调控; 3)沙丘生态系统种子库受各种干扰的综合影响; 4)生境对沙丘生态系统种子库具有重要影响; 5)沙丘生态系统种子库结构和组成与植物繁殖对策密切关联; 6)种子库对沙丘生态系统植被恢复贡献率因沙丘类型而异。沙丘生态系统种子库研究应强化如下方面: 1)种子库区域分异规律研究; 2)种子库与植物生活史阶段的联系的研究; 3)多种干扰方式综合影响下的种子库研究; 4)种子库对沙丘生态系统植被恢复贡献的研究; 5)沙丘生态系统自然物理过程与种子库耦联关系的研究。     

Genetic connectivity and inter-population seed dispersal of Banksia hookeriana at the landscape scale

Background and Aims

Landscape genetics combines approaches from population genetics and landscape ecology, increasing the scope for conceptual advances in biology. Banksia hookeriana comprises clusters of individuals located on dune crests (geographical populations) physically separated by uninhabitable swales, with local extinctions common through frequent fire and/or severe drought.

Methods

A landscape genetics approach was used to explore landscape-scale genetic connectivity and structure among geographical populations of B. hookeriana on 18 physically separated dunes located within a heterogeneous landscape of 3 × 5 km. These geographical populations were separated by approx. 0·1 to >1 km of unsuitable intervening swale habitat. Using 11 highly variable microsatellite loci, we utilized a Bayesian approach to identify genetic discontinuities within and between these geographical populations. Population allocation tests were then used to detect inter-dune seed dispersal inferred from assignment of individuals to a source population other than that from which they were collected.

Key Results

For the modal number of genetically distinct clusters (n = 17 genetic populations), two coincided with the geographical (dune) populations, eight spanned two to four geographical populations, and the remaining seven were spread among various parts of the sampled dunes, so that most geographical populations were spatially defined mosaics of individuals (subpopulations) belonging to two or more genetic populations. We inferred 25 inter-dune immigrants among the 582 individuals assessed, with an average distance between sink and source dunes of 1·1 km, and a maximum of 3·3 km.

Conclusions

The results show that genetic structure in an apparently strongly spatially structured landscape is not solely dependent on landscape structure, and that many physically defined geographical populations were genetic mosaics. More strikingly, there were physically separated individuals and groups of individuals that were part of the same genetically defined populations. We attribute this mismatch between spatially and genetically defined population structure to the varying closeness of the dunes and the ability of seeds to disperse long distances.     

Soil requirements of central Australian sandridge plants in relation to the duneswale soil catena

The soil requirements of seven central Australian sandridge plant species were examined by comparing the soils associated with individuals of each species in their typical sandridge habitats and in anomalous habitats such as dry watercourses, grader banks on road verges, disturbed ground near boreholes, and sand-filled rock crevices. The species studied were Zygochloa paradoxa, Chamaesyce wheeleri, Calotis erinacea, Helichrysum ambiguum, Dampiera cinerea, Triodia basedowii and Monachather paradoxa. The first two of these are characteristic of open dunecrests, the next three of crests and upper dune slopes, and the last two of lower slopes and swales. The first five species grow in a range of anomalous habitats with nutrient status comparable to swales and lowerslopes, but only if the soils are loosened or have enhanced moisture status. The deep wellsorted sands of dry watercourses form the only anomalous habitat for Zygochloa paradoxa and Chamaesyce wheeleri. Calotis erinacea, Helichrysum ambiguum and Dampiera cinerea also grow on poorly sorted graderbank and borehole soils in the swales, but not on unmodified swale soils. The only anomalous habitats in which the last two have been found are very similar to their typical sandridge habitats and so reveal little of soil requirements.     

毛乌素沙地沙丘土壤含水量特点——兼论老固定沙地上油蒿衰退原因

土壤表面干沙层在冬春少雨季节较厚,流动沙地的又明显地大于固和地的。流动沙地地表干沙层以下的土壤含水量一般在于同深化度固定沙地的,特别是在报系密集分布的土层。科季或早春土壤含水量比较低,雨季的明显较高,雨季结束后,土层中的水分在重力的作用下会向深层运动,导致根系层土壤含水量下降。在植物生长季,缺少降雨时,因少地的土土壤水量会因植物的蒸腾消耗而变得很低（１％～２％）,而流动沙地的不受或受此影响很小。地     

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.