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.     

The role of the expansion of native-invasive plant species in coastal dunes: The case of Retama monosperma in SW Spain

Invasion by allochthonous plant species are identified, at present, among the main conservation hazards to coastal dunes. Nevertheless, the role of the expansion, with invasive character, of native species in these ecosystems has received little attention in ecological studies. In recent decades, Retama monosperma, a late colonizing legume shrub found in coastal sandy areas, endemic to the SW of the Iberian Peninsula and NW Morocco, has displayed invasive behavior in coastal dunes in different parts of the world, including its natural area of distribution. Its rapid expansion and increase in coverage has significantly contributed to the dune stabilization process, sometimes involving notable changes in the environment, plant community and shaping local distribution of some associated fauna, thus modifying the functioning of the whole ecosystem. In this review we examine the role of the expansion of R. monosperma in SW Spain coastal dunes, causes and implications, in the context of the ecological theory of invasions, and comparing it with other case studies.     

Multiple introductions and no loss of genetic diversity: invasion history of Japanese Rose, Rosa rugosa, in Europe

The shrub Rosa rugosa (Japanese Rose), native to East Asia, is considered one of the most troublesome invasive plant species in natural or semi-natural habitats of northern Europe and has proven very difficult to control. We aimed at disentangling the species’ invasion history in Europe, including determining the number of introductions and their geographic origin, and at investigating whether populations in the introduced and native ranges differ in genetic diversity, structure and degree of differentiation. We found that introduced (n = 16) and native (n = 16) populations had similar levels of genetic diversity at seven nuclear SSR (microsatellite) loci. European populations lack isolation by distance and are less genetically differentiated than are populations in East Asia. Multiple and at least three independent colonization events, one of which was particularly successful, gave rise to current R. rugosa populations in Europe. The geographic distribution patterns of these three genetic clusters could not be explained by natural dispersal alone, indicating that human mediated secondary dispersal is driving the expansion in Europe. One cluster representing three of the European populations was most likely derived from NW Japan, whereas the origin of the remaining thirteen populations could not clearly be resolved. The introduction and expansion in Europe occurred with no significant loss of genetic diversity. We conclude that high propagule pressure at the primary establishment phase is the most parsimonious explanation for this pattern. A potential for long distance seed dispersal, coastal habitat connectivity and an outcrossing breeding system are factors likely to have enabled populations of R. rugosa to avoid detrimental effects of genetic bottlenecks and will further increase the species’ range size and abundance in Europe. We recommend that human-mediated dispersal should be prevented in order to halt the continued expansion.     

Uprooting and Burial of Invasive Alien Plants: A New Tool in Coastal Restoration?

Invasive alien plants are a problem for conservation management, and control of these species can be combined with habitat restoration. Subsoil burial of uprooted plants is a new method of mechanical control, which might be suitable in disturbed habitats. The method was tested in Rosa rugosa (Japanese Rose), an invasive shrub in north‐western Europe with negative effects on coastal biodiversity. Two months after uprooting and burial in dunes of north‐eastern Denmark, 89% of the 58 shrubs resprouted from roots and rhizomes; on average 41 resprouts per shrub. Resprout density was twice as high at former shrub margins compared with the center; resprouts were taller and originated from more superficial soil layers at the margin than in the center. Resprouting was negatively correlated with fragment depth, and no resprouts were observed from greater than 15 cm depth. The number of resprouts increased with fragment dry mass (0.5–168.5 g). After 18 months with harrowing the species was still resprouting, flowering, and fruiting, albeit with no difference between shrub margin and center. Resprouts were taller (26 cm) and coverage was higher (0–4%) after two compared with three times harrowing, whereas no difference was found in cover of native dune species (1–5%). The results show that even small fragments of R. rugosa resprout, and that resprouting persists despite repeated harrowing. Thus, careful subsoil burial of all fragments is necessary, special attention should be paid to the shrub margin, and follow‐up treatments are needed. The effectiveness of the burial method is discussed for restoration of coastal dunes.     

Long-term spatio-temporal expansion of the native-invasive Retama monosperma on coastal dunes: Importance of land-use and natural dispersal vectors

Expansion of invasive species inducing sands stabilization is becoming an important ecological problem in coastal dunes in some parts of the world. Retama monosperma, an endemic to coasts of SE Spain and NE of Morocco, late-colonizing shrub occurring in sandy coastal areas, was planted along the coast of SW Spain during the 20th century to stabilize dunes. In recent decades, the species has spread rapidly, becoming invasive, and sometimes incurring notable changes in its environment and plant community. The expansion patterns of R. monosperma from 1956 to 2001 were described and quantified at the landscape scale within a protected dune system, using GIS. The differential effects on expansion patterns of the main factor controlling the population spread, grazing by domestic livestock and the abundance of wild rabbits, were analyzed comparatively. R. monosperma displays an exponential, invasive-type expansion trend, presenting a mean annual coverage increase of 15% and a mean lateral spread rate of 65.5 m yr⁻¹ from the original population nucleus to the Western and Eastern ends of the study area. The dispersal activity primarily of rabbits and the absence of competing woody species contribute to this rapid expansion. The highest local increases in plant coverage could be related to increased rabbit abundance and to the improved germination rates of seeds which have passed through the rabbit gut. In contrast, areas highly grazed by livestock present the lowest increases, and an open-type plant community is maintained there for a longer time.     

Regional and local variation of spider assemblages (Araneae) from coastal grey dunes along the North Sea

Aim This study aims to determine the underlying causes of local and regional patterns of variation in community structure of spiders in coastal grey dunes, and especially whether ecological time constrains the species composition in young and isolated grey dune habitats. Location The study was conducted in coastal dunes from northern France (Boulonnais, Nord‐Pas‐de‐Calais), Belgium (Flemish coastal dunes) and the Netherlands [Amsterdam Water Supply (AWD) dunes and Dunes from the Provincial Water Company North‐Holland]. Methods Spiders were collected with pitfall traps in twenty‐eight grey dune patches in the four areas under investigation. Species composition and environmental parameters (vegetation structure, distance to the sea and the patch‐edge, eolic dynamics and lime richness) were determined. Assemblage composition was related to the regional and local environmental factors with the Primer software package to determine the assemblage‐determining parameters. Differences in species presence were analysed as a function of their habitat preference and distribution range. Results Differences in grey dune spider assemblage structure can mainly be attributed to differences in local sand dynamics and the region. Species from dynamic dunes are mainly present in grey dunes from Belgium and France, while species from non‐dunal xerothermic habitats (chalk grasslands and heathland) occur in both the Boulonnais and the north Holland dune region. These species are absent from geologically young or other xerotherm habitat isolated Flemish coastal dunes. Main conclusion The data show that regional variation in spider assemblage composition results from local landscape characteristics (dynamics in the dune area), the latitude and the connectivity to non‐dunal xerothermic habitats. The strong and moderate geological isolation of dune areas from the Flemish coast and the AWD dunes, respectively, results in the absence of (at least some) species that are primarily bound to heathland and/or chalk grassland. This indicates the importance of ecological time for the assemblage structure. The limited dispersal capacity of the heathland and/or chalk grassland species is probably the main reason for their absence.     

玫瑰种群生物学研究进展

玫瑰(Rosa rugosa Thunb.)原产于我国东部沿海、日本、朝鲜半岛和俄罗斯远东地区,18世纪作为园艺种引入欧洲后逃逸并入侵至北海和波罗的海周边多个国家以及北美沙质海岸,而中国野生种群却在过去30年间持续萎缩,成为珍稀濒危物种。从玫瑰种群生物学角度,通过文献比较和综合,在阐明玫瑰生态学特性和野生分布变化的基础上,全面论述了玫瑰种群在我国的生境退化、种群动态、种子繁殖、遗传多样性、濒危机理、保育方面的成果和悬疑问题;并结合欧洲入侵种群分布范围和敏感生境、对本地群落和物种的影响、种子繁殖、遗传变异、种群扩张和模拟预测、管理和控制方面的研究进展,分析了濒危种群和入侵种群数量动态、群落组成、幼苗更新、遗传变异、管理策略方面的差异及其影响因素;进而提出未来的玫瑰研究可从濒危种群和入侵种群的比较研究、种群和灌丛的动态监测、适合度相关性状的变异及其遗传基础、基于种群生物学的保育或控制4个方面为切入点,集中探索玫瑰种群濒危和入侵动态的规律、遗传基础和主要驱动力,为玫瑰保育和管理提供理论依据,为相似物种的适应和进化机制研究提供例证。     

The impact of human pressure on landscape patterns and plant species richness in Mediterranean coastal dunes

We analyze the spatial patterns of natural dune cover patches and their plant richness, comparing coastal sites with different levels of human pressure in central Italy. We created a detailed land cover map of dune sites. The spatial pattern of natural dune cover types was characterized by computing a set of patch-based metrics. To quantify patch plant richness, we used 16 m² vegetation plots, randomly distributed on coastal dune cover types. For each patch, the richness of the entire pool of species and of three guilds (i.e., typical dune, ruderal, and alien species) was considered. We compared different levels of human pressure on coastal dunes focusing on pattern metrics and floristic information by using the nonparametric Kruskal–Wallis test. In sites with high human pressure, we have observed a general simplification in the natural dune spatial pattern and a decline of plant richness but with a specific response for each cover type. Alien and ruderal species presented low richness in all patches. In coastal dunes, the harsh ecological conditions and the strong sea–inland gradient shape the distribution of human activities and control the number of ruderal species. The approach effectively describes fragmentation and biodiversity in dune ecosystems.     

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     

Recruitment limitation of native species in invaded coastal dune communities

Recruitment limitation may limit the ability of sites to regenerate after disturbances such as weed invasion and weed management. We investigated seed bank constraints and dispersal limitation in coastal dune communities on the east coast of Australia. The ability of sites to regenerate naturally following weed removal was assessed in coastal dune communities invaded by the invasive alien, bitou bush (Chrysanthemoides monilifera subsp. rotundata). To investigate recruitment limitation, seed banks and vegetation of invaded, native, intensively managed (selective application of herbicide and some re-vegetation) and extensively managed (large-scale, non-selective herbicide application) sites were compared. We investigated the dispersal mechanisms of species in the seed bank and vegetation to determine if communities might be dispersal-limited, i.e. contain significant numbers of species with only short-distance dispersal capabilities. Species richness and composition of soil seed banks differed from the vegetation in foredunes and hinddunes. Invasion depleted seed banks further. About half of the species had short-distance dispersal mechanisms indicating the potential for dispersal limitation. Secondary weed invasion following management was evident although alien species occurred in both seed banks and vegetation. Our results indicated that coastal dune communities suffer recruitment limitation. Native, managed and invaded dune communities appear to be both seed bank and dispersal-limited although management and invasion exacerbates recruitment. Regeneration of coastal dune communities will require active reintroduction of species, particularly those with short-distance dispersal mechanisms.     

Coastal dune vegetation and pollen representation in south Buenos Aires Province, Argentina

Aim To establish the relationship between coastal dune vegetation and its pollen representation as an aid to interpret Holocene vegetation dynamics and environmental changes from pollen assemblages. Location The study area is situated on the temperate Atlantic coast of south Buenos Aires Province, Argentina (c. 39° S and 61°20′ W). Methods The vegetation of the active dune area adjacent to the beach was described on the basis of its floristic composition from 25 plots. Classification of the vegetation into distinct zones was carried out by cluster analysis. Surface samples were collected from each vegetation stand and analysed for their pollen composition. Pollen percentage data were analysed using principal components analysis in order to investigate the degree to which the different vegetation units can be distinguished by their pollen spectra. Pollen–vegetation relationships for selected taxa were explored using simple scatter plots and indices of association, under‐ and over‐representation. Indices of floristic diversity and palynological richness were used to assess the representation of the vegetation in the pollen spectra. Results and conclusions Five vegetation zones are defined on the basis of species composition and their quantitative variation: back shore, mobile dunes, slacks, semi‐fixed and fixed dunes. Pollen assemblages from back shore, mobile dunes and slacks are clearly differentiated from semi‐fixed and fixed dunes. Pollen assemblages differ considerably from the associated vegetation composition. Major discrepancies are caused by large differences in pollen and vegetation proportion of Hyalis argentea and Discaria americana. There is a considerable proportion of non‐local pollen in every spectrum. Pollen representation in the coastal dunes at Monte Hermoso is influenced by differences in pollen production, dispersal and preservation of individual taxa as well as by the spatial distribution of the vegetation, the topography of the dune system and the wind pattern. The pollen–vegetation relationship established in this study has important implication for understanding and interpreting fossil pollen records from coastal dune environments.     

Positive responses of coastal dune plants to soil conditioning by the invasive Lupinus nootkatensis

Invasive nitrogen-fixing plants drive vegetation dynamics and may cause irreversible changes in nutrient-limited ecosystems through increased soil resources. We studied how soil conditioning by the invasive alien Lupinus nootkatensis affected the seedling growth of co-occurring native plant species in coastal dunes, and whether responses to lupin-conditioned soil could be explained by fertilisation effects interacting with specific ecological strategies of the native dune species. Seedling performance of dune species was compared in a greenhouse experiment using field-collected soil from within or outside coastal lupin stands. In associated experiments, we quantified the response to nutrient supply of each species and tested how addition of specific nutrients affected growth of the native grass Festuca arundinacea in control and lupin-conditioned soil. We found that lupin-conditioned soil increased seedling biomass in 30 out of 32 native species; the conditioned soil also had a positive effect on seedling biomass of the invasive lupin itself. Increased phosphorus mobilisation by lupins was the major factor driving these positive seedling responses, based both on growth responses to addition of specific elements and analyses of plant available soil nutrients. There were large differences in growth responses to lupin-conditioned soil among species, but they were unrelated to selected autecological indicators or plant strategies. We conclude that Lupinus nootkatensis removes the phosphorus limitation for growth of native plants in coastal dunes, and that it increases cycling of other nutrients, promoting the growth of its own seedlings and a wide range of dune species. Finally, our study indicates that there are no negative soil legacies that prevent re-establishment of native plant species after removal of lupins.     

Shrubs as ecosystem engineers in a coastal dune: influences on plant populations,communities and ecosystems

Question: How do two shrubs with contrasting life‐history characteristics influence abundance of dominant plant taxa, species richness and aboveground biomass of grasses and forbs, litter accumulation, nitrogen pools and mineralization rates? How are these shrubs – and thus their effects on populations, communities and ecosystems – distributed spatially across the landscape? Location: Coastal hind‐dune system, Bodega Head, northern California. Methods: In each of 4 years, we compared vegetation, leaf litter and soil nitrogen under canopies of two native shrubs –Ericameria ericoides and the nitrogen‐fixing Lupinus chamissonis– with those in adjacent open dunes. Results: At the population level, density and cover of the native forb Claytonia perfoliata and the exotic grass Bromus diandrus were higher under shrubs than in shrub‐free areas, whereas they were lower under shrubs for the exotic grass Vulpia bromoides. In contrast, cover of three native moss species was highest under Ericameria and equally low under Lupinus and shrub‐free areas. At community level, species richness and aboveground biomass of herbaceous dicots was lower beneath shrubs, whereas no pattern emerged for grasses. At ecosystem level, areas beneath shrubs accumulated more leaf litter and had larger pools of soil ammonium and nitrate. Rates of nitrate mineralization were higher under Lupinus, followed by Ericameria and then open dune. At landscape level, the two shrubs – and their distinctive vegetation and soils – frequently had uniform spatial distributions, and the distance separating neighbouring shrubs increased as their combined sizes increased. Conclusions: Collectively, these data suggest that both shrubs serve as ecosystem engineers in this coastal dune, having influences at multiple levels of biological organization. Our data also suggest that intraspecific competition influenced the spatial distributions of these shrubs and thus altered the distribution of their effects throughout the landscape.     

Impacts of a woody invader vary in different vegetation communities

The impact of an exotic species in natural systems may be dependent not only on invader attributes but also on characteristics of the invaded community. We examined impacts of the invader bitou bush, Chrysanthemoides monilifera ssp. rotundata , in fore and hind dune communities of coastal New South Wales, Australia. We compared invader impacts on vegetation structure, richness of both native and exotic growth forms and community variability in fore and hind dunes. We found that impacts of bitou invasion were context specific: in fore dune shrublands, functionally distinct graminoid, herb and climber rather than shrub growth forms had significantly reduced species richness following bitou invasion. However, in forested hind dunes, the functionally similar native shrub growth form had significantly reduced species richness following bitou invasion. Density of vegetation structure increased at the shrub level in both fore and hind dune invaded communities compared with non-invaded communities. Fore dune ground-level vegetation density declined at invaded sites compared with non-invaded sites, reflecting significant reductions in herb and graminoid species richness. Hind dune canopy-level vegetation density was reduced at invaded compared with non-invaded sites. Bitou bush invasion also affected fore dune community variability with significant increases in variability of species abundances observed in invaded compared with non-invaded sites. In contrast, variability among all hind dune sites was similar. The results suggest that effects of bitou bush invasion are mediated by the vegetation community. When bitou bush becomes abundant, community structure and functioning may be compromised.     

Landscape genetics of an endangered lemur (Propithecus tattersalli) within its entire fragmented range

Habitat fragmentation may strongly reduce individuals’ dispersal among resource patches and hence influence population distribution and persistence. We studied the impact of landscape heterogeneity on the dispersal of the golden‐crowned sifaka (Propithecus tattersalli), an endangered social lemur species living in a restricted and highly fragmented landscape. We combined spatial analysis and population genetics methods to describe population units and identify the environmental factors which best predict the rates and patterns of genetic differentiation within and between populations. We used non‐invasive methods to genotype 230 individuals at 13 microsatellites in all the main forest fragments of its entire distribution area. Our analyses suggest that the Manankolana River and geographical distance are the primary structuring factors, while a national road crossing the region does not seem to impede gene flow. Altogether, our results are in agreement with a limited influence of forest habitat connectivity on gene flow patterns (except for North of the species’ range), suggesting that dispersal is still possible today among most forest patches for this species. Within forest patches, we find that dispersal is mainly among neighbouring social groups, hence confirming previous behavioural observations.     

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。     

Hard times for Italian coastal dunes: insights from a diachronic analysis based on random plots

Multi-year temporal studies are invaluable tools for monitoring changes in biodiversity through time. However, their applications in coastal ecosystems are still scarce. We investigated temporal trends in coastal dunes analyzing a set of 858 randomly-sampled georeferenced relevés performed between 2002 and 2015 along Central Italy’s sandy coastlines. Specifically, we explored changes in species richness and cover of targeted sandy habitats, we investigated trends in the cover of selected psammophilous native species and we assessed patterns of invasion by means of regression techniques. We observed a significant decrease in species richness and cover of the dune grasslands habitat. The species-level analysis confirmed a negative trend for two characteristic species of dune grasslands, Cutandia maritima and Medicago littoralis, while revealing a similar decline for Crucianella maritima and for Ammophila arenaria subsp. australis, key species of mobile dunes. The most striking trends emerged analyzing patterns in the cover of an invasive alien species, Carpobrotus sp., which showed a concerning increase in shifting dunes. In conclusion, our analyses reveal concerning changes involving dune grasslands, and at the same time hint at “early warnings” of degradation processes traceable in shifting dunes.     

Patch quality and connectivity influence spatial dynamics in a dune wolfspider

The spatial population dynamics of the wolfspider Pardosa monticola, inhabiting patchily distributed grasslands in the Flemish coastal dunes of Belgium and Northern France were investigated with incidence function models using field survey data from 1998 and 2000. Vegetation height and patch size were related to habitat quality. Mark-recapture experiments revealed maximum cursorial dispersal distances of 280 m for moss dunes and 185 m for higher dune grassland. Higher shrub vegetation appeared to be dispersal barriers. These habitat-dependant cursorial distances and the theoretically estimated ballooning distance were included with patch distances into a connectivity index for both dispersal modes. Forward multiple regression indicated that patch occurrence was influenced by habitat quality and ballooning connectivity. Habitat quality and cursorial connectivity explained patterns in short-term colonisation. Extinction appeared to be stochastic and not related to habitat quality and connectivity. Genetic differentiation and variability was low. The discrepancy between the estimated low dispersal capacity and the indirect estimate of gene flow ( F(ST)) indicates that historical population dynamics and/or historical ballooning dispersal influence the genetic structure in this species.