High competitiveness of a resource demanding invasive acacia under low resource supply

Mechanisms controlling the successful invasion of resource demanding species into low-resource environments are still poorly understood. Well-adapted native species are often considered superior competitors under stressful conditions. Here we investigate the competitive ability of the resource demanding alien Acacia longifolia, which invades nutrient-poor Mediterranean sand dunes such as in coastal areas of Portugal. We explore the hypothesis that drought may limit invasion in a factorial competition experiment of the alien invasive versus two native species of different functional groups (Halimium halimifolium, Pinus pinea), under well-watered and drought conditions. Changes in biomass, allocation pattern, and N-uptake-efficiency (via ¹⁵N-labeling) indicated a marked drought sensitivity of the invader. However, highly efficient drought adaptations of the native species did not provide a competitive advantage under water limiting conditions. The competitive strength of H. halimifolium towards the alien invader under well-watered conditions turned into a positive interaction between both species under drought. Further, low resource utilization by native species benefited A. longifolia by permitting continued high nitrogen uptake under drought. Hence, the N-fixing invader expresses low plasticity by continuous high resource utilization, even under low resource conditions. The introduction of novel traits into a community like N-fixation and high resource use may promote A. longifolia invasiveness through changes in the physical environment, i.e., the water and nutrient cycle of the invaded sand dune system, thereby potentially disrupting the co-evolved interactions within the native plant community.     

Autecological traits ofHalimium halimifolium in contrasting habitats under a Mediterranean type climate—A review

Halimium halimifolium (L.)Willk. is a woodyCistaceae species occurring locally in the Western Mediterranean. At the Doñana National Park (S Atlantic coast of Spain),Halimium halimifolium is the main component of stable sand vegetation. It grows in a range of environmental conditions from flood-prone depressions to mobile dunes, but it is most abundant, and dominates the scrub composition on the slopes of fixed dune ridges with a water table depth of 2 to 4 m. The species exhibits not only morphological modifications (hairy leaves and twigs), but also structural and ecophysiological adaptations. The response ofH. halimifolium to stress conditions (leaf water potential, leaf diffusion resistance, and plant growth) have been studied in the field, throughout an annual cycle in four populations growing in different environments, three dune types differing in soil moisture, and one dune slack. Results showed that plants from the hygrophytic area (MN) had the less negative water potentials, the lowest stomatal resistances and the biggest vegetative growth, together with the highest Leaf Area Index, canopy light extinction, the largest leaf area and the lowest sclerophyll index. Plants from the most xerophytic area (MB) had the lowest Leaf Area Index, the smallest leaf area and the highest sclerophyll index. Even though in dune slopes (MI) water table was shallower than in MB, plants in the latter had more negative water potentials and bigger vegetative growth. Water potential values only exhibited significant differences among the four populations at the end of the spring period and over the summer. These results are discussed in relation to the climatic conditions of the study period (3 years of a drought cycle). Plants from the mobile dune system showed features which were intermediate between MN and MI plants.     

Interactions between seedlings of Chrysanthemoides monilifera and Acacia longifolia

Displacement of Acacia longifolia on coastal dunes in New South Wales by the invasive species Chrysanthemoides monilifera may be linked to the greater competitiveness by the latter in the seedling stage, as demonstrated in pot experiments. This occurs despite a lower chlorophyll concentration in shoots of C. monilifera which leads to a lower assimilation rate per unit leaf area and lower carbohydrate concentrations. However, this assimilate is spread over a greater total leaf area. Such a strategy associated with‘quantity’may thus be more important than leaf‘quality’in terms of competitiveness. In A. longifolia, the production of higher quality‘leaves’but of lower total area may be well-suited in the often sparse native populations found in sand dunes, but appears disadvantageous when seedlings of C. monilifera also co-exist. The competitive advantage of C. monilifera over A. longifolia is reduced but not reversed under water stress. Under severe stress, mortality of C. monilifera is greater than that of A. longifolia in monocultures but mortality of both species is similar in mixtures. The reason appears to be that C. monilifera transpires more water per plant even though its rate of transpiration per unit leaf area is reduced under water stress because of early stomatal closure. In mixtures, faster root growth of C. monilifera ensures faster uptake of the available soil water, thus minimizing the inherent advantage in A. longifolia of its lower water use and greater efficiency.     

Plant community response following the removal of the invasive Lupinus arboreus in a coastal dune system

The removal of invasive species is common in restoration projects, yet the long‐term effects of pest management programs are seldom assessed. We present results of a long‐term program to remove the invasive species Lupinus arboreus (lupin) from sand dunes in New Zealand. We evaluate the response of plant communities to lupin removal, by comparing total plant cover, the cover of non‐native and native plant species, and species richness between sand dune sites where lupin removal has occurred, not occurred, and where lupin has never been present. Neither lupin presence nor removal had a significant impact on the foredune environment. Following removal, total and other non‐native plant cover remained higher, and the cover of several native sand dune species remained lower compared with uninvaded sites in the deflation and backdune environments. These changes can be attributed to persistent effects associated with the invasion of lupin, but have also developed in response to lupin removal. The results of this study have implications for restoration projects in sand dunes. Pest management alone is unlikely to be sufficient to restore plant communities. Given the difficulties in restoring plant communities once an invasive species has established, managers should prioritize actions to prevent the spread of invasive species into uninvaded areas of sand dunes. Finally, the response to lupin invasion and removal differed between dune habitats. This highlights the importance of tailoring a pest management program to restoration goals by, for example, prioritizing areas in which the impacts of the invading species are greatest.     

Ecophysiological differences among Leymus mollis populations across a subarctic dune system caused by environmental, not genetic, factors

Plant species that persist during succession, from the colonization to the stabilization stages, face major environmental changes. Such changes are believed to have significant effects on species performance. In subarctic coastal dune systems, Leymus mollis colonizes the embryo dunes, on the upper limit of the beach. It reaches its maximum density on the foredune, but also grows on older, stabilized ridges. This paper reports on the phenotypic variations of some ecophysiological traits associated with the persistence of L. mollis on a dune system on the east coast of Hudson Bay (northern Quebec). Leymus mollis ramets tend to have a lower net carbon assimilation rate and water use efficiency, and a higher substomatal CO₂ concentration on the stabilized dune than on the foredune. However, these physiological differences cannot be explained by differences in leaf morphology or nitrogen content. Under controlled conditions, ecophysiological differences observed in the field disappear, suggesting that these are not genetic but determined by environmental changes along the foredune-stabilized dune gradient. We propose that higher net carbon assimilation rate on the foredune might be related to higher sink strength in relation to the growth-stimulating effect of sand burial.     

Interactions between mycorrhizal colonization and plant life forms along the successional gradient of coastal sand dunes in the eastern Mediterranean, Turkey

The mycorrhizal status of dune plant species in relation to their plant life forms was surveyed along a successional gradient of sand dune on the southern Mediterranean coast of Turkey. Roots of 64 dune plant species belonging to 30 families were collected from sand dune communities at four different successional stages: embryonic dunes (ED), mobile dunes (MD), fixed dunes (FD), and remnant dunes (RD). Of the plant species surveyed in all successional stages, 54 (84%) had formed mycorrhizal associations. Nonmycorrhizal plants with cryptophyte life forms predominated in the earlier successional stages (ED and MD), whereas the number and percent coverage of mycorrhizal plant species belonging to hemicryptophytes, phanerophytes, and chamaephytes generally increased with the stabilization of sand dunes. Arbuscular mycorrhizal (AM) colonization was found to be the dominant mycorrhizal type in ED, MD, and RD. But phanerophytes with dual colonization, AM and ectomycorrhizal, became the dominant life form with high plant coverage in the FD stage. Total percentage of mycorrhizal root length colonization showed significant positive correlations relating to soil parameters such as organic matter and nitrogen content, while negatively correlating to high soil reaction (pH).     

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。     

The sand dune vegetation of Chrystalls Beach, southern New Zealand, with particular reference to the cushion community

The sand dune system at Chrystalls Beach, New Zealand comprises front dune, dune hollow and rear dune areas. On the front dunes are two communities: one on the seaward face of the dunes, and another on the landward face. Most of the dune hollow is occupied by a series of grassland communities, but there are areas of a distinctive cushion community, still largely native in its species composition. It is suggested that the cushion community is maintained against invasion by a switch, possibly involving sand grain size, soil organic content, water supply, wind and magnesium:calcium ratio. The coarse sandy/pebbly substrate, with its low organic content, causes water stress. The water stress permits only low vegetation, which exacerbates the low soil organic content. The low vegetation results in high wind, again favouring cushion species. The water regime has also allowed a buildup of Mg:Ca ratio, especially in the lower layers of the soil.     

Invasive species can handle higher leaf temperature under water stress than Mediterranean natives

Thermal tolerance of Photosystem II (PSII) highly influences plant distribution worldwide because it allows for photosynthesis during periods of high temperatures and water stress, which are common in most terrestrial ecosystems and particularly in dry and semi-arid ones. However, there is a lack of information about how this tolerance influences invasiveness of exotic species in ecosystems with seasonal drought. To address this question for Mediterranean-type ecosystems (MTE) of the Iberian Peninsula, we carried out an experiment with fifteen phylogenetically related species (8 invasive and 7 native, Pinus pinaster Ait., Pinus radiata D. Don, Schinus molle Linn., Elaeagnus angustifolia L., Eucalyptus globulus Labill., Acacia melanoxylon R. Br., Gleditsia triacanthos L., Pistacia terebinthus L., Rhamnus alaternus L., Anagyris foetida L., Colutea arborescens L., Oenothera biennis L., Epilobium hirsutum L., Achillea filipendulina Lam. and Achillea millefolium L). Seedlings were grown and maximal photochemical efficiency of PSII (Fv/Fm) was measured at two water availabilities (well-watered and with water stress). PSII thermal tolerance measurements were related to specific leaf area (SLA), which varied significantly across the study species, and to the mean potential evapotranspiration (PET) of the month with the lowest precipitation in the native areas of both groups and in the invaded area of the Iberian Peninsula. Additionally, PSII thermal tolerance measurements under water stress were phylogenetically explored. Invasive and native species neither differed in SLA nor in their thermal tolerance under well-watered conditions. For well-watered plants, SLA was significantly and positively related to PSII thermal tolerance when all species were explored together regardless of their invasive nature. However, this relationship did not persist under water stress and invasive species had higher plastic responses than Mediterranean natives resulting in higher leaf temperatures. Higher PSII thermal tolerance could explain invasiveness because it allows for longer periods of carbon acquisition under water stress. In fact, PSII thermal tolerance was positively related to the PET of the invaded and native areas of the Iberian Peninsula. PSII thermal tolerance was not related to PET at the native range of the invasive species, suggesting that successful invasive species were plastic enough to cope with novel dry conditions of the Iberian Peninsula. Moreover, our phylogenetic results indicate that future scenarios of increased aridity in MTE associated to climate change will filter invasion success by taxonomic identity. This study reveals the importance of studying ecophysiological traits to understand and better predict future biological invasions.     

Linkages between woody plant proliferation dynamics and plant physiological traits in southwestern North America

Aims Woody plant encroachments in arid and semiarid ecosystems are widely reported but the physiological mechanisms still need to be further revealed. In the current study, we aim to determine whether differences in leaf physiological traits help explain grassland susceptibility to woody plant encroachment and whether distinctive physiological adaptations allow some shrub species to invade grasslands.Methods We compared physiological traits (photosynthesis, leaf water status, pigment compositions and leaf antioxidant capacities) of six species representing three functional groups: woody encroachers (Prosopis velutina, Larrea tridentata), woody non-encroachers (Acacia greggii, Lycium fremontii) and C4 grasses (Bouteloua curtipendula, Bouteloua barbata) which are naturally growing in a botanical garden in University of Arizona, USA.Important findings We infer that P. velutina (encroacher) but not A. greggii or L. fremontii (non-encroachers) is encroaching in grasslands because the former species has higher water and light utilization efficiencies (instantaneous water use efficiency, instantaneous light use efficiency, and Fv/Fm). The extremely high carotenoid and total antioxidant capacity in its leaves appears to help the shrub L. tridentata (encroacher) survive high ambient oxidative damage caused by both drought and high light stresses in this grassland. The two C₄ grass species, B. curtipendula and B. barbata, grow well in the arid ecosystem but may be susceptible to disturbances.     

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.     

Ecology of Indian desert. III. Survival adaptations of vegetation in dry environment

Summary A preliminary investigation was made on the survival adaptations of 45 plant species growing in Indian arid zone. A number of species sprang up soon after rains in July when they grew fast and many of them disappeared by December because of dry soil conditions and desiccating atmosphere. An account is presented here on the ecophysiological and morpho-physiological adaptations for survival in drought escaping, evading, enduring and resisting species. The stomatal behaviour during the day, water loss, leaf water deficit, leaf water content and relative water content have been examined for each species during late October till early December 1970. An account of their survival qualities with particular reference to the long dry period following after monsoons in relation to drought resisting species was considered. The significance of adaptive features for each and every species was correlated with its drought tolerance under the prevailing conditions. On the basis of their water loss, the species have been grouped into six categories.     

Genetic diversity of rhizobia associated with Acacia longifolia in two stages of invasion of coastal sand dunes

We examined the genetic diversity of root nodule bacteria associated with the Australian legume Acacia longifolia in two stages of invasion of a coastal sand dune system. All isolates belonged to the genus Bradyrhizobium. A higher diversity was found in the long-established trees. The results suggest the introduction of exotic bradyrhizobia with the plant.     

Does salt stress increase the ability of the exotic legume Acacia longifolia to compete with native legumes in sand dune ecosystems?

Sand dune ecosystems are one of the areas most affected by the introduction of invasive species which represents a threat for biodiversity conservation. Their invasion patterns and spread may depend on their salinity tolerance, besides other factors. To test this hypothesis, we investigated the effects of salt stress on seed germination and on the activity of antioxidant enzymes (catalase, CAT; ascorbate peroxidase, APX; peroxidase, POX; and glutathione reductase, GR) in two legume species, an invasive, Acacia longifolia (Andrews.) Willd., and a native, Ulex europaeus (L.), very common in the sand dunes of the coast of Portugal. Salt stress was induced by adding NaCl at different concentrations, 0, 50, 100 and 200 mM, for 15 days. Results showed that the highest germination percentages were obtained in distilled water (control) and that, with increasing salt concentration, seed germination was delayed and decreased in both species. Inhibition of germination was higher in the native species, only 3% of seeds germinated at 100 mM and no seeds germinated at 200 mM NaCl. In the invasive species, the reduction was higher at 200 mM NaCl (16%). Considering the coefficient of germination velocity, a decrease in both species with increasing NaCl concentration was observed. The CAT and GR activities decreased in A. longifolia with increasing salinity. In turn, APX activity significantly increased as NaCl concentration increased while the POX activities declined at the highest NaCl concentration. On the other hand, at 50 mM NaCl lower activity of CAT and APX and higher GR and POX were found in U. europaeus. In both species, protein content increased as NaCl concentration increased. In addition, it seems that APX activities play an essential role in the scavenging reactive oxygen species (ROS). These results suggest that the seeds of the invasive legume A. longifolia are more tolerant to salinity than the native legume U. europaeus, and seem better equipped to handle the physiological stress of high salinity, which may contribute to its invasive ability in sand dunes.     

不同沙丘生境主要植物比叶面积和叶干物质含量的比较

研究了生长在不同沙丘生境中 (流动沙丘 ,半固定沙丘和固定沙丘 ) 2 0个植物种 (10个 1年生植物种和 10个多年生植物种 )的比叶面积 (SL A)和叶干物质含量 (L DMC)的变化 ,并且分析了各个沙丘生境的土壤养分特征。结果表明 ,各个植物种的平均 SL A和 L DMC在植物种之间差异显著 ;多数在两种或 3种沙丘生境均有分布的植物其 SL A在不同沙丘生境之间差异显著 ,但是仅有 6个植物种的 L DMC在不同沙丘生境之间表现出差异 (p<0 .0 5 )。与许多研究结果类似 ,1年生植物的 SL A显著大于多年生植物的 SL A,而且两者之间 L DMC存在一定的差异。 1年生植物 SL A和 L DMC之间相关性不显著 ,但多年生植物SL A和 L DMC之间呈显著负相关。综合所有 2 0个植物种可以发现 ,SL A增大时 ,L DMC有下降的趋势     

Ecophysiological traits of invasive alien Acacia cyclops compared to co‐occuring native species in Strandveld vegetation of the Cape Floristic Region

Tree invasions of Mediterranean‐climate ecosystems pose a significant threat to both biodiversity and functioning, by excluding native species, altering soil nutrient status and depleting water resources. In order to attain greater relative biomass associated with successful invasion in these characteristically resource‐poor environments, invasive species must have novel traits that enable better acquisition (e.g. deep roots) or exploitation of different resources (e.g. N₂ fixation) and/or more efficient use of available resources than native species. We compared the ecophysiological and morphological traits of three abundant native species to those of the invasive Australian tree species, Acacia cyclops. This species is widely invasive in the Mediterranean‐climate coastal vegetation of South Africa that includes the Strandveld vegetation type. A. cyclops had 30–50% greater foliar N concentrations (P < 0.001) in comparison with the native species and lower foliar δ¹⁵N values that may indicate N₂ fixation. Additionally, A. cyclops maintained higher photosynthetic rates over the dry summer season (ca. 15 μmol m^?2 s^?1) than the native species. These higher photosynthetic rates may result from sustained access to water due to deeper rooting abilities as indicated by the more negative δD values (P < 0.001) of A. cyclops (?43‰) in comparison with the some native species (?29 to ?37‰). Acacia cyclops did not, however, exhibit greater water use efficiencies or photosynthetic nitrogen use efficiencies (P > 0.05) compared to native species. Invasiveness of A. cyclops into this resource‐limited Mediterranean‐climate ecosystem appears to be supported by greater resource acquisition, possibly partially through N₂ fixation and greater rooting depth, rather than greater resource use efficiency or conservation.     

Gender, season and habitat: Patterns of variation in photosynthetic activity, growth and fecundity in Thymelaea velutina

Changes in the ecophysiological performance of a plant species due to different environmental conditions generally reflect adaptations to the habitat where the plant grows and are often related to its survival capacity in a particular place. We examined this with the dioecious shrub Thymelaea velutina, in two contrasting populations representing the extremes of the altitudinal gradient where the species lives (coastal dunes and mountain habitats over 1000 m). We measured net photosynthetic rates and stomatal conductance, estimated the level of plant stress by chlorophyll fluorescence, and assessed their correlations with growth rate, plant size, flower production and fruit set. We hypothesized that plants at high altitude were more photosynthetically stressed than at sea level and expected a gender × habitat interaction in performance as females need more resources than males. Plants in the mountain experienced chronic photoinhibition during winter and a reduced photosynthetic performance both in winter and spring compared to plants in coastal dunes. However, there was no association between any of the fluorescence variables and either plant growth or fecundity, suggesting that other factors are involved determining performance. Mountain plants showed also an apparent lower capacity of heat dissipation to excessive radiation than dune plants. In the dunes, the greater leaf area and mass can lead to a higher photosynthetic carbon gain by whole individuals compared to plants in the mountain. No effect of gender was detected on the ecophysiological performance of this species, which we partly attribute to the small size of fruits of the female plants.     

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     

Woody plant encroachment impacts on soil carbon and microbial processes: results from a hierarchical Bayesian analysis of soil incubation data

Quantitative methods were used to examine soil properties and their spatial heterogeneity in a 0-year fenced mobile dune (MD0), an 11-year fenced mobile dune (MD11) and a 20-year fenced mobile dune (MD20) in Horqin Sandy Land, Northern China. The objective of the study was to assess the effect of vegetation restoration on heterogeneity of soil properties in sand dunes and to provide a concept model to describe the relationship between vegetation succession and spatial heterogeneity variation of soil properties in the dunes. The results showed that the average values of vegetation cover, species number and diversity, soil organic carbon (C), total nitrogen (N), and electrical conductivity (EC) increased with the increase in fenced age of mobile dunes, while soil water content (0–20 cm) showed the reverse trend. Geostatistical analysis revealed that the spatial heterogeneity of soil organic C, total N, EC, very fine sand content, and soil water content (0–20 cm) increased from MD0 to MD11 with succession from sand pioneer plant to shrub species then decreased from MD11 to MD20 due to continuous development of herbaceous plants. Canonical correspondence analysis (CCA) showed that there was a relatively high correspondence between vegetation and soil factors, suggesting that the major gradients relating soil organic C, total N, EC, pH, slope, very fine sand content, and soil water content are the main factors for the distribution of dune plants and account for 68.1% of the species-environment relationship among the three sites. In addition, the distribution of the sand pioneer plant was positively related to the relative height of the sampling site and soil water content, and that of most herbaceous plants were determined by soil organic C, total N, EC, pH, and very fine sand content in mobile dunes. The conceptual model of relationship between vegetation succession and spatial heterogeneity of soil properties in mobile dunes suggests spatial patterns of soil properties are most strongly related to plant-induced heterogeneity in dune ecosystems prone to wind erosion, and conversely, the magnitude and degree of spatial heterogeneity in soil properties can influence the plant distribution pattern and vegetation succession of mobile dunes.     

Ecological Assessment of Dune Restorations in the Great Lakes Region

Because of the economic and environmental importance of stabilizing fragile sand dune habitats, restoration of dunes has become a common practice. Restoration efforts in the Great Lakes and East Coast regions of North America often consist of planting monocultures of the dominant native grass species, Ammophila breviligulata. We evaluated 18 dune restoration projects in the Great Lakes region conducted over the past 25 years. We characterized attributes of diversity (plants and insects), vegetation structure (plant biomass and cover), and ecological processes (soil nutrients and mycorrhizal fungi abundance) in each restoration, and we compared these measures to geographically paired natural dune communities. Restoration sites were similar to reference sites in most measured variables. Differences between restorations and reference sites were mostly explained by differences in ages, with the younger sites supporting slightly lower plant diversity and mycorrhizal spore abundance than older sites. Plant community composition varied little between restored and reference sites, with only one native forb species, Artemisia campestris, occurring significantly more often in reference sites than restored sites. Although it remains unclear whether more diverse restoration plantings could accelerate convergence on the ecological conditions of reference dunes, in general, traditional restoration efforts involving monoculture plantings of A. breviligulata in Great Lakes sand dunes appear to achieve ecological conditions found in reference dunes.