Millipede communities in rehabilitating coastal dune forests in northern KwaZulu/Natal, South Africa

The rehabilitation, after mining, of coastal sand dunes north of Richards Bay by Richards Bay Minerals began some 18 years ago, and resulted in the simultaneous availability of a known-aged series of stands representative of coastal dune forest succession.
A survey of the millipede community in this area revealed increases in species diversity and a decrease in their density, with an increase in stand age. Development of these communities is characterized by replacement and addition of species, typical of ecological succession. Colonization of areas disturbed by mining reflects on the species reservoir present in the surrounding unmined forests with apparent pioneer species being either replaced or complemented by the relatively slow invasion of secondary species.
Comparisons of age-specific millipede community variables on rehabilitating dunes with those recorded in relatively undisturbed dunes suggest that the development of communities results from autogenic succession initiated through habitat rehabilitation. With several community parameters in rehabilitating dune forests being similar to those recorded in undisturbed forests, it is concluded that the millipede community can be restored through management options based on principles relating to ecological succession.     

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.     

Restoration of riverine inland sand dune complexes: implications for the conservation of wild bees

1.  The evaluation of restoration measures is an important task of conservation biology. Inland sand dunes and dry, oligotrophic grasslands have become rare habitat types in large parts of Central Europe and their restoration and management is of major importance for the preservation of many endangered plant and insect species. Within such habitats, it is important to restore key ecosystem services, such as pollination networks. As wild bees are the most important pollinators in many ecosystems, they represent a suitable key group to evaluate restoration measures. Furthermore, the recent decline of many bee species and the potential ecological and economic consequences are currently topics of strong scientific interest.
2.  We studied the succession of bee communities in response to restoration measures of sand dunes and sand grasslands and compared these communities with those of old sand dune complexes.
3.  Our results show that wild bees respond rapidly to restoration measures indicated by a high species richness and abundance. The community structure of bees at restoration sites converged only slightly to those of the target sites. A higher similarity was found between bee communities at the restoration sites (sand dunes and grasslands), indicating that their close proximity was an important determinant of species overlap. Environmental factors such as the number of entomophilous plant species and moisture had a strong influence on wild bee species composition.
4.   Synthesis and applications . The restoration of inland sand dune complexes provides opportunities for colonization by a diverse wild bee community. Although it is difficult to establish a given target community, restoration measures gave rise to a high pollinator diversity and abundance, suggesting that community function can be re-established.     

Climate and coastal dune vegetation: disturbance, recovery, and succession

The sand dune habitats found on barrier islands and other coastal areas support a dynamic plant community while protecting areas further inland from waves and wind. Foredune, interdune, and backdune habitats common to most coastal dunes have very different vegetation, likely because of the interplay among plant succession, exposure, disturbance, and resource availability. However, surprisingly few long-term data are available describing dune vegetation patterns. A nine-year census of 294 plots on St. George Island, Florida suggests that the major climatic drivers of vegetation patterns vary with habitat. Community structure is correlated with the elevation, soil moisture, and percent soil ash of each 1 m² plot. Major storms reduce species richness in all three habitats. Principle coordinate analysis suggests that changes in the plant communities through time are caused by climatic events: changes in foredune vegetation are correlated with temperature and summer precipitation, interdune vegetation with storm surge, and backdune vegetation with precipitation and storm surge. We suggest that the plant communities in foredune, interdune, and backdune habitats tend to undergo succession toward particular compositions of species, with climatic disturbances pushing the communities away from these more deterministic trajectories.     

Notes on Lejeuneaceae: I. Microlejeunea diversiloba (Spr.) K.M.

Abstract

Previous work on the bryophytic component of dune communities has demonstrated changes in the representation of species with the progress of succession. The ability of some to act as pioneers and the restriction of others to later stages may be associated with their reactions to recurrent burial by wind-blown sand.

Eighteen species were selected for experimental investigation of the effects of sand burial, along the following lines:

(a) Transplants. Samples of eight species were transplanted to four sites representing different stages of the dune succession. Where the sand increment over 11 months exceeded about 4 cm. none survived but species found naturally in mobile dune areas emerged from about 3 cm. Those belonging to fixed dune communities survived only in the absence of burial, although on excavation all species showed some upward growth through the covering sand.

(b) Experimental burial, Method I: samples of the same eight species, grown in pots standing in water but not watered from above, were buried in sand to depths of 1, 2, 4, 5 and 6 cm. The maximum depth through which emergence took place was 4 cm., from which Ceratodon purpureus, Bryum pendulum and Brachythecium albicans reappeared. From shallower layers, however, a greater rate of emergence was shown by Tortula ruraliformis. Pseudoscleropodiun purum and Climacium dendroides did not survive depths greater than 2 cm., and 1cm. was the maximum for Hylocomium splendns and Rhytidiadelphus squarrosus.

Method II. Sample strips of fourteen species were placed in aquarium tanks and, apart from a small portion len exposed, were buried under a sand slope grading up to 7 cm. in depth. Watering was from above and the humidity was kept high by glass lids. The time taken for the appearance of shoots from increasing depths was recorded, also the periods required for reestablishment from increasing depths of 50% and 100% of the original shoot density. For species tested by both methods the results were in most cases similar. Marked powen of recolonization were also shown by Polytrichum piliferum, P. juniperinum and Pohlia annotina.

In P. annotina, Ceratodon purpureus and Dicramum scoparium re-establishment by means of vertically extending rhizoids producing protonemata at the surface is described.

The maximum depth from which the various species can emerge, the depths from which effective recolonization of the surface is achieved within a fairly short time, the ability to produce rhizoids in the overlying sand and the growth-form of the shoots on emergence are regarded as the most significant aspects of reaction to burial, in relation to the role of the species in their natural habitats.     

Bryophytes in Mediterranean coastal dunes: ecological strategies and distribution along the vegetation zonation

The bryophytic flora has been widely studied in Atlantic coastal dune environments while bryological literature addressing the Mediterranean bryophytes is still scarce and fragmented, although they are an important component of this area, especially considering their ability to colonize and stabilize sandy soils. The aims of this study were: (i) to analyze the taxonomy, chorology and ecology of coastal dunes bryoflora; (ii) to analyze distribution patterns of bryophytic species along a coastal dune vegetation zonation. We used as coastal dune model system a particularly well-preserved study area of the Sardinian coast (Italy), which includes several habitats listed in European Habitats Directive 92/43/EEC. We conducted a general survey of the area and a random sampling. A total of 20 bryophytic species were detected most of which were acrocarpous mosses and, to a lesser extent, pleurocarpous and liverworts. Most bryophytes are pioneer colonizers, reflecting their adaptation to environments with high levels of stress/disturbance conditions such as those of coastal dunes. Finally, in this study we described for the first time how bryophytic species are distributed along a Mediterranean coastal dune vegetation zonation, focusing also on their chorological and ecological characteristics and on how they seem to follow the sea-land gradient, probably in response to different environmental conditions developing along this gradient.     

Effects of fire on the structure and composition of open eucalypt forests

Abstract Fires are integral to the healthy functioning of most ecosystems and are often poorly understood in policy and management, however, the relationship between floristic composition and habitat structure is intrinsically linked, particularly after fire. The aim of this study was to test whether the variability of habitat structure or floristic composition and abundance in forests at a regional scale can be explained in terms of fire frequency using historical data and experimental prescribed burns. We tested this hypothesis in open eucalypt forests of Fraser Island off the east coast of Australia. Fraser Island dunes show progressive stages in plant succession as access to nutrients decreases across the Island. We found that fire frequency was not a good predictor of floristic composition or abundance across dune systems; rather, its affects were dune specific. In contrast, habitat structure was strongly influenced by fire frequency, independent of dune system. A dense understorey occurred in frequently burnt areas, whereas infrequently burnt areas had a more even distribution of plant heights. Plant communities returned to pre‐burn levels of composition and abundances within 6 months of a fire and frequently burnt areas were dominated by early successional species of plant. These ecosystems were characterized by low diversity and frequently burnt areas on the east coast were dominated by Pteridium. Greater midstorey canopy cover in low frequency areas reduces light penetration and allows other species to compete more effectively with Pteridium. Our results strongly indicate that frequent fires on the Island have resulted in a decrease in relative diversity through dominance of several species. Prescribed fire represents a powerful management tool to shape habitat structure and complexity of Fraser Island forests.     

Invasive Congeners Differ in Successional Impacts across Space and Time

Invasive species can alter the succession of ecological communities because they are often adapted to the disturbed conditions that initiate succession. The extent to which this occurs may depend on how widely they are distributed across environmental gradients and how long they persist over the course of succession. We focus on plant communities of the USA Pacific Northwest coastal dunes, where disturbance is characterized by changes in sediment supply, and the plant community is dominated by two introduced grasses – the long-established Ammophila arenaria and the currently invading A. breviligulata. Previous studies showed that A. breviligulata has replaced A. arenaria and reduced community diversity. We hypothesize that this is largely due to A. breviligulata occupying a wider distribution across spatial environmental gradients and persisting in later-successional habitat than A. arenaria. We used multi-decadal chronosequences and a resurvey study spanning 2 decades to characterize distributions of both species across space and time, and investigated how these distributions were associated with changes in the plant community. The invading A. breviligulata persisted longer and occupied a wider spatial distribution across the dune, and this corresponded with a reduction in plant species richness and native cover. Furthermore, backdunes previously dominated by A. arenaria switched to being dominated by A. breviligulata, forest, or developed land over a 23-yr period. Ammophila breviligulata likely invades by displacing A. arenaria, and reduces plant diversity by maintaining its dominance into later successional backdunes. Our results suggest distinct roles in succession, with A. arenaria playing a more classically facilitative role and A. breviligulata a more inhibitory role. Differential abilities of closely-related invasive species to persist through time and occupy heterogeneous environments allows for distinct impacts on communities during succession.     

Spatial indicators of plant community assembly from a 453-year sand-dune chronosequence

Background : We explored evidence for spatial patterning in vegetation across a sand-dune chronosequence spanning 453 years of primary succession to test for indications of a temporal signature of niche versus neutral processes during community assembly.

Methods : The study was conducted on a series of dune-capped beach ridges located in Wilderness State Park in Emmet County of northern Lower Michigan, United States (45° 43′ N, 84° 56′ E). Nearest-neighbour survey data were subjected to species dispersion and association analyses.

Results : Both species richness and species evenness showed the classic ‘hump-shaped’ relationship across successional time, suggesting that competition is an important force in structuring this community. However, the only evidence of intra-specific aggregation occurred in the youngest dune. Older dunes displayed random spatial structure and the oldest dunes tended toward a mix of random patterns and intra-specific segregation, i.e. with nearest neighbours more likely than by chance to be inter-specific. In addition, the frequency of species having lower neighbour-species richness than expected by chance was higher in younger, not older dunes. Species with significantly low neighbour richness (based on absolute nearest neighbours) on later dunes tended to have a high proportion of nearest neighbours that were intra-specific.

Conclusions : The data provided no evidence that resident species in later succession have greater intra-specific aggregation or more niche differentiation based on horizontal niche space. The data are more consistent with theories of community assembly involving competitive equivalence and/or effects of facilitation.     

两类植物型沙丘上植物群落的异同及其对沙丘形态的响应

为探究相同环境条件下发育的两类植物型沙丘上植物群落的异同点,以及相异点与沙丘形态的关系,对毛乌素沙地南缘盐碱地上相间分布的抛物线形沙丘和白刺灌丛沙丘分别进行形态学、植物群落学及土壤理化性质的调查分析。结果显示:抛物线形沙丘的水平尺度虽为白刺灌丛沙丘的12-23倍,但两类沙丘上物种数相当,其植物分属12科31属39种和12科30属33种,均以菊科、藜科、禾本科、豆科植物为主(占70%以上),其中共有植物17种,群落相似度0.66;两类沙丘上的群落建群种不同,优势种的重叠度较低,抛物线形沙丘的不同部位共统计到7个植物群丛,可分为沙生植物群落和喜湿耐盐碱群落,白刺灌丛沙丘上均以白刺为建群种,油蒿、冰草、雾冰藜、沙蓬、狗尾草为主要优势种;沙丘形态造成其不同部位风沙活动及土壤水分、PH值和全盐含量的差异是两类沙丘上植物群落相异的重要影响因素。     

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.     

Patterns of phylogenetic community structure of sand dune plant communities in the Yucatan Peninsula: the role of deterministic and stochastic processes in community assembly

ABSTRACT

Background: Tropical sand dunes are ideal systems for understanding drivers of community assembly as dunes are subject to both deterministic and stochastic processes. However, studies that evaluate the factors that mediate plant community assembly in these ecosystems are few.

Aims: We evaluated phylogenetic community structure to elucidate the role of deterministic and stochastic processes in mediating the assembly of plant communities along the north of the Yucatan Peninsula, Mexico.

Methods: We used plastid genetic markers to evaluate phylogenetic relationships in 16 sand-dune communities. To evaluate the role of climate in shaping plant community structure we carried out linear regressions between climatic variables and mean phylogenetic distance. We estimated the Net Relatedness Index and Nearest Taxon Index to identify ecological processes mediating community assembly.

Results: Observed phylogenetic structure was not different from random, suggesting that stochastic processes are the major determinants of community assembly. Climate was slightly correlated with phylogenetic diversity suggesting that abiotic environment plays a minimal role in community assembly.

Conclusions: Random assembly appears to be the primary factor structuring the studied sand dune plant communities. Environmental filters may represent a secondary factor contributing to the observed phylogenetic structure. Thus, both processes may act simultaneously to mediate the assembly of sand-dune plant communities.     

臭柏群落在不同演替阶段某些群落特征的研究

毛乌素沙地臭柏群落自发演替的基本过程为：半流动沙地沙竹 +沙米（有时是白沙蒿）群落→半固定沙地油蒿+臭柏群落→固定沙地臭柏+硬质早熟禾群落→固定沙地臭柏群落→老固定沙地臭柏+苔藓群落,最后可能发展成为地带性的本氏针茅草原。演替早期向演替亚顶级群落发展过程中物种多样性逐步增加,生态环境逐渐变湿。随着臭柏的衰退,某些喜湿植物的退出,物种多样性开始减少,群落生境向旱生化发展。     

Characterization of arbuscular mycorrhizal fungal communities with respect to zonal vegetation in a coastal dune ecosystem

Coastal dune vegetation distributes zonally along the environmental gradients of, e.g., soil disturbance. In the preset study, arbuscular mycorrhizal fungal communities in a coastal dune ecosystem were characterized with respect to tolerance to soil disturbance. Two grass species, Elymus mollis and Miscanthus sinensis, are distributed zonally in the seaward and landward slopes, respectively, in the primary dunes in Ishikari, Japan. The seaward slope is severely disturbed by wind, while the landward slope is stabilized by the thick root system of M. sinensis. The roots and rhizosphere soils of the two grasses were collected from the slopes. The soils were sieved to destruct the fungal hyphal networks, and soil trap culture was conducted to assess tolerance of the communities to disturbance, with parallel analysis of the field communities using a molecular ecological tool. In the landward communities, large shifts in the composition and increases in diversity were observed in the trap culture compared with the field, but in the seaward communities, the impact of trap culture was minimal. The landward field community was significantly nested within the landward trap culture community, implying that most members in the field community did not disappear in the trap culture. No nestedness was observed in the seaward communities. These observations suggest that disturbance-tolerant fungi have been preferentially selected in the seaward slope due to severe disturbance in the habitat. Whereas a limited number of fungi, which are not necessarily disturbance-sensitive, dominate in the stable landward slope, but high-potential diversity has been maintained in the habitat.     

Aphid herbivory as a potential driver of primary succession in coastal dunes

Herbivory is a major factor affecting both the performance and the fitness of the species composing a plant community and, ultimately, conditioning its temporal and spatial dynamics. Coastal dunes are a typical example of primary succession where different biotic and abiotic factors determine plant species occurrence; however, the effect of insect herbivory herein has remained little explored. To address this matter, we combined an observational study along a successional gradient with a green-house experiment to determine the occurrence and the impact of plant–aphid interactions. We focused on the species Schizaphis rufula, a widespread and abundant aphid associated with dune grasses in early stages of primary succession in Europe. Firstly, we studied aphid infestation rates on the dune grass Ammophila arenaria along a succession gradient in three locations of the North Sea coast to address the relationship between plant community composition and aphid occurrence; secondly, we tested the effect of aphid herbivory on a set of dune species typical for the different stages of succession. We found that the degree of aphid infestation was inversely correlated with the degree of dune fixation. The results of the experiment showed that aphid multiplication was significantly higher and its effect more pronounced on two early successional grass species, i.e. A. arenaria and Leymus arenarius. Here aphid multiplication resulted in a severe decrease in plant biomass; in late successional grass species, there was limited multiplication and no effect on biomass. The results of the field survey and the green-house experiment indicate that aphids show a clear preference for plants from early successional stages and, moreover, they have a greater impact on these plant species. All this supports the hypothesis of aphid herbivory as a driving factor of primary succession in coastal dunes.     

Environmental factors and ground disturbance affecting the composition of species and functional traits of ground layer lichens on grey dunes and dune heaths of Estonia

A unique, species‐rich and endangered lichen biota can be found on European coastal and inland sand dunes. However, it is increasingly affected by natural succession as well as by anthropogenic disturbances. We studied lichen diversity on the grey dunes and dune heaths of coastal and inland regions of Estonia. A total of 28 study plots were investigated; in each 0.1 ha study plot general environmental variables and anthropogenic disturbances were described and all epigeic lichen species were identified. We found 66 lichenized fungus (lichen) species, including several rare and ten red‐listed lichens. Multivariate analysis (DCA, CCA) was performed to examine gradients in species composition and to relate variation in species data to environmental factors. In addition, we used redundancy analysis (RDA) to relate variation in species’ trait composition to environmental factors. Species composition on grey dunes differed significantly from that on dune heaths. The characteristic species for grey dunes are, besides several Cladonia species, foliose lichens, e.g. Hypogymnia physodes, Parmelia sulcata and Peltigera spp. Also species’ traits composition was different for either habitat, indicating that sorediate lichens, foliose lichens, lichens with cyanobacterium as the main photobiont, and sparsely branched Cladonia species dominate on grey dunes, while esorediate, green‐algal, crustose and richly branched fruticose lichens are common on dune heaths. Soil pH was the most essential environmental variable for determining both species composition and species’ traits composition. The composition of lichen species was also significantly influenced by forest closeness, soil Mg content and cover of bare sand; the effect of ground disturbances was low compared to the effect of these environmental factors. To protect and conserve the species‐rich lichen biota, it is necessary to protect the dune habitats from building activity, to avoid overtrampling in recreation areas and to regularly remove shrubs and trees.     

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.     

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.     

Development of vegetation in dune slack wetlands of Cape Cod National Seashore (Massachusetts,USA)

Interdunal seasonal wetlands, known as dune slack wetlands, were mapped and their vegetation surveyed across a large region of dunes within the Cape Cod National Seashore. Wetland sizes and ages were estimated from digitized, georeferenced aerial photographs available from 1938, 1947, 1960, 1986, 1994, and 2001, and from LIDAR elevation data. A total of 346 sites were found, covering an area of ∼45.4 ha., in which 97 species of vascular plants were identified. Vegetation structure and composition exhibited a distinct sequence of development with age, following a pattern of succession from herbaceous, graminoid-dominated communities to shrub- and tree-dominated communities. Floristic variables were not related to proximity to the coast and although wetland size appeared to have some bearing on species richness, the correlation was statistically weak. Soil organic matter determined for a subset of 60 wetlands was positively correlated with age and woody cover but showed no relationship with water depth. The results suggest that vegetation development is primarily driven by the internal mechanisms of succession. Notwithstanding, any changes in the environment that alter the process of succession will collectively influence these wetlands. In addition, stabilization of the dunes resulting in a reduction in the formation of new wetlands, may translate into permanent loss of early and mid-successional dune slack communities. Maintenance of these communities will depend on succession being periodically reset by disturbance or active management.     

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.