Multi-scale phylogenetic structure in coastal dune plant communities across the globe

Aims Studies integrating phylogenetic history and large-scale community assembly are few, and many questions remain unanswered. Here, we use a global coastal dune plant data set to uncover the important factors in community assembly across scales from the local filtering processes to the global long-term diversification and dispersal dynamics. Coastal dune plant communities occur worldwide under a wide range of climatic and geologic conditions as well as in all biogeographic regions. However, global patterns in the phylogenetic composition of coastal dune plant communities have not previously been studied.Methods The data set comprised vegetation data from 18463 plots in New Zealand, South Africa, South America, North America and Europe. The phylogenetic tree comprised 2241 plant species from 149 families. We calculated phylogenetic clustering (Net Relatedness Index, NRI, and Nearest Taxon Index, NTI) of regional dune floras to estimate the amount of in situ diversification relative to the global dune species pool and evaluated the relative importance of land and climate barriers for these diversification patterns by geographic analyses of phylogenetic similarity. We then tested whether dune plant communities exhibit similar patterns of phylogenetic structure within regions. Finally, we calculated NRI for local communities relative to the regional species pool and tested for an association with functional traits (plant height and seed mass) thought to vary along sea–inland gradients.Important findings Regional species pools were phylogenetically clustered relative to the global pool, indicating regional diversification. NTI showed stronger clustering than NRI pointing to the importance of especially recent diversifications within regions. The species pools grouped phylogenetically into two clusters on either side of the tropics suggesting greater dispersal rates within hemispheres than between hemispheres. Local NRI plot values confirmed that most communities were also phylogenetically clustered within regions. NRI values decreased with increasing plant height and seed mass, indicating greater phylogenetic clustering in communities with short maximum height and good dispersers prone to wind and tidal disturbance as well as salt spray, consistent with environmental filtering along sea–inland gradients. Height and seed mass both showed significant phylogenetic signal, and NRI tended to correlate negatively with both at the plot level. Low NRI plots tended to represent coastal scrub and forest, whereas high NRI plots tended to represent herb-dominated vegetation. We conclude that regional diversification processes play a role in dune plant community assembly, with convergence in local phylogenetic community structure and local variation in community structure probably reflecting consistent coastal-inland gradients. Our study contributes to a better understanding of the globally distributed dynamic coastal ecosystems and the structuring factors working on dune plant communities across spatial scales and regions.     

Environmental Factors Influencing Coastal Vegetation Pattern: New Insights from the Mediterranean Basin

Coastal dune ecosystems show strongly dynamic interactions between abiotic and biotic factors. The relationship between plant communities and environmental factors has been previously studied in oceanic dune systems, but few studies have been conducted along Mediterranean coasts. In this study we analyze the relative contribution of environmental factors associated with two of the main drivers of vegetation zonation: soil and wind. We chose two representative coastal dune systems in the western Mediterranean Basin subject to low levels of human disturbance. Within 54 plots we recorded floristic and environmental data. Vegetation zonation and relationships with environmental variables were investigated through unconstrained and constrained ordinations, correlation, and variance partitioning. Environmental factors shift along the gradient from coastal to inland dunes, concomitantly with the pattern of community types from annual beach communities to shrub-covered fixed dunes. This general gradient is similar both in the Mediterranean and in the oceanic coastal ecosystems, with the same factors that show similar trends along the dune profile. However, our results highlight some peculiarities of the Mediterranean dune systems in relation to the amount of variation explained by environmental factors. While most studies conducted in oceanic ecosystems find that wind-related parameters may control the vegetation zonation, in our study areas we observed a minor importance of the wind-related variables when compared to soil properties. In particular, organic matter and grain-size variability were found to be closely correlated with the distribution of plant communities along the gradient.     

Are some communities of the coastal dune zonation more susceptible to alien plant invasion?

Aims Coastal areas, and in particular coastal dunes, are ecosystems strongly affected by the invasion of alien plants. However, few attempts have ever been made to quantify alien species incidence in different communities along the coastal zonation. This work aims to analyze the distribution of alien plants along the coastal zonation of sandy shores on the Tyrrhenian coast, addressing specifically differences among plant communities in abundance of alien plants.Methods The study was performed on recent dunes (Holocene) of the central western coast of Italy. We selected dune landscapes where invasion processes were particularly evident. Vegetation plots were randomly sampled and through cluster analysis, we identified six plant communities corresponding to the typical zonation described for the Tyrrhenian sandy coast of Central Italy. We evaluated and compared frequency and abundance of invasion in these different communities. Further, we investigated how propagule pressure (measured using as proxy human structures) contributed to the observed invasion patterns.Important findings We found a relatively low total number of aliens but also a differential distribution pattern and strong abundance of some of the aliens in specific sectors of the vegetation zonation. The perennial community of transition dunes appears most affected by invasion processes, related almost exclusively to the frequent and widespread Carpobrotus aff. acinaciformis. This alien species reaches high cover values, apparently lowering cover of native species of transition dune plant communities. Higher levels of invasion in the transition dune can be partially explained because of greater propagule pressure in this section of the dune profile. Our findings thus have important conservation and management implications since transition dune communities with Crucianella maritima are rare and protected (sensu Habitat 92/43/EEC Directive) along the entire Italian coast.     

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.     

CO2-fluxes in different plant communities of a high-Arctic tundra watershed (Western Spitsbergen)

Abstract. There is little information on (1) whether the net carbon balance is positive or negative in different habitats in high Arctic ecosystems such as Spitsbergen today, and (2) what effect a cloudier, cooler summer could have on carbon balance. To provide data on this subject CO₂-flux measurements in different plant communities were made in the high-Arctic coastal tundra of Spitsbergen, using a mobile macro-cuvette system based on infrared gas analysis. The study area was situated on the exposed west coast, where westerly winds produce precipitation daily in the form of rain, drizzle and fog. During the cold and cloudy measurement period in 1996, light and surface temperatures were limiting for primary producers, resulting in low size and low physiological activity of the plants. Net CO₂-flux measurements showed carbon fluxes from soil to atmosphere in most of the communities even during the brightest hours of the day, when mean photon flux density was 325 mmol m^-2 s^-1. Calculations based on spatial distribution of the plant communities and soils in the watershed combined with C-flux measurements revealed information on daily carbon loss. For instance, the Drepanocladus community, covering 21 % of the catchment area, was responsible for 42.6 % of the catchment carbon loss. Only two of nine investigated plant communities, the Racomitrium and a Salix-Saxifraga community on debris, both adapted to frequent fog situations, were able to compensate for respiratory CO₂-losses under the prevailing low light conditions during daytime. Since there were no significant sunny periods in this area in the summer of 1996, the habitats of the investigated coastal tundra finished the season with a marked carbon loss due to increased cloudiness.     

Does disturbance drive the collapse of biotic interactions at the severe end of a diversity–biomass gradient?

It has been recently proposed that the decrease in diversity towards the severe end of the humped-back diversity–biomass model of Grime was driven by a collapse of facilitation due to extreme conditions of either stress or physical disturbance. In order to test the hypothesis that disturbance is the primary direct factor driving the collapse of interactions occurring along environmental severity gradients, we conducted a removal experiment in the highly stressed French coastal dunes along a gradient of disturbance due to sand burial. Four dune species were used as targets and transplanted with and without neighbours in four communities along the gradient. The experiment was conducted twice, a dry and an average year. Results of the experiment showed that during the dry year the effect of the environment was prominent and only one species was facilitated for survival in the least disturbed community. During the average year, interactions for growth were important only in the same community, with positive or negative responses depending on the natural position of the target species within the coastal dune gradient. In accordance with our hypothesis, most interactions for both survival and growth were observed in the least disturbed community exhibiting the highest diversity. There were no interactions in the most disturbed community with the lowest diversity.     

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.     

Multi-scale segregations and edaphic determinants of marsh plant communities in a western Pacific estuary

Whether and how the roles of environmental factors in producing vegetation patterns in coastal marshes vary with spatial scale is not well understood. We investigated the relationship between plant communities and edaphic factors in the Yangtze estuary at three spatial scales. Plant communities and edaphic factors were quantified at high and low tidal levels in both freshwater and salt marshes. Canonical correspondence analyses were conducted to examine the relationship between plant communities and edaphic factors at the landscape scale (freshwater vs. salt marsh), the zonation scale (high vs. low tidal level) and the patch scale (dominant vs. other species). Soil salinity, moisture content, pH, bulk density, and organic carbon could well explain segregations of plants at the landscape and zonation scales. However, the same factors exhibited only very weak relationships to plant communities at the patch scale. These results suggest that plant communities in the Yangtze estuary are segregated at different spatial scales by different environmental factors. As spatial scale is often not explicitly addressed investigating community assembly rules, our study underscores the importance of scaling for an improved understanding of community organization in coastal wetlands.     

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.     

Trampling affects the distribution of specialised coastal dune arthropods

From a conservation point of view, species’ tolerances towards disturbance are often generalised and lack reference to spatial scales and underlying processes. In order to investigate how average typical species react to habitat fragmentation and disturbance, we adopted a multi-species approach to address occupancy patterns of five specialised dune arthropods (butterflies Hipparchia semele, Issoria lathonia; grasshopper Oedipoda caerulescens; spiders Alopecosa fabrilis, Xysticus sabulosus) in recently fragmented coastal dune habitats which are subjected to varying levels and modes of local disturbance, i.e. trampling by cattle or people.Occupancy patterns were assessed during two successive years in 133 grey dune fragments of the Flemish coastal dunes (Belgium, France). By treating species as a random factor in our models, emphasis was placed on generalisations rather than documenting species-specific patterns.Our study demonstrates that deteriorating effects of local disturbance on arthropod incidence cannot be interpreted independent of its landscape context, and appear to be more severe when patch area and connectivity decrease. When controlled for patch area and trampling intensity, the probability of species occupancy in poorly connected patches is higher under cattle trampling than under recreation. Incidences additionally decrease with increasing intensity of cattle trampling, but increases with trampling by tourists.This study provides evidence of mode- and landscape-dependent effects of local disturbance on species occupancy patterns. Most importantly, it demonstrates that trampling of sensitive dune fragments will lead to local and metapopulation extinction in landscapes where trampling occurs in a spatially autocorrelated way, but that the outcome (spatial patterns) varies in relation to disturbance mode, indicating that effects of disturbance cannot be generalised.     

Impact of anthropogenic disturbances on beetle communities of French Mediterranean coastal dunes

In coastal dunes, influenced by anthropogenic activities such as tourism, it is important to determine the relative influence of environmental factors at different spatial scales to evaluate the sensitivity of local communities to disturbances. We analyzed beetle communities of 14 dunes of the French Mediterranean coast: four in the relatively preserved Camargue area, and ten in the Var department, where tourism is intensive. Beetle communities were studied three times in early spring using sand sampling. Species-environment relationships were evaluated at the regional, landscape and local scale using redundancy analysis (RDA) and variability partitioning. About 28 species were identified, of which 15 were sand-specialist species, which accounted for more than 93% of total abundance. The beetle communities of Camargue were significantly different from those of the Var department owing to the pullulation of a Tenebrionid species (Trachyscelis aphodioides Latr.) in the Var, except for one restored dune where the community was very similar to those of Camargue. Our results showed no longitudinal gradient between the two regions. Local factors (dune height, preservation and disturbance index) significantly explained most of the variation in the dominance of T. aphodioides, while some other local factors were important for other psammophilous species. This study also suggests that dune beetle communities are strongly affected on beaches intensively managed for tourism, but beetles are still abundant in much disturbed sites.     

Environmental tuning of an insect ensemble: The tenebrionid beetles inhabiting a Mediterranean coastal dune zonation

Few studies have investigated insect ensembles, i.e. phylogenetically bounded groups of species that use a similar set of resources within a community. The zonation of dune vegetation makes these ecosystems ideal for the study of insect ensembles in a short space. In this study, we investigated if the tenebrionid beetles forming an ensemble on a dune zonation showed variations in community organization (relative abundances and species diversity) in different but spatially associated biotopes defined by different plant communities. Three biotopes (corresponding to European Commission habitat 2110, 2120 and 2230) of a well-preserved Mediterranean dune were sampled using square plots of 2 × 2 m at three places. To investigate if there was some association between species and habitat we applied a χ² test. Variations in community structure parameters were investigated using Shannon index. The three biotopes host tenebrionid communities with similar species composition and overall abundances, confirming that they form a single ensemble. However, tenebrionid species are differently associated with different biotopes along the zonation, with some species occurring with different proportions among the biotopes. A local selection process can be postulated as a mechanism responsible for these differences.     

植物群落动态的模型分析

植物群落的动态是植物群落学的中心问题之一,包括更新、波动、演替、进化等主要内容。空间格局对种群和群落的动态起着至关重要的作用,种群空间格局和群落空间结构是群落中各种过程相互作用的产物。模型是描述群落动态、认识植物群落组建和维持机理的有效工具。本文阐述和比较了描述群落动态的四种具有代表性的经验模型,即镶嵌循环模型、随意游走模型、同资源种团比例模型、空间抢先占有模型及其机理。四种经验模型的空间性及缺陷分别是:(1)“镶嵌循环模型”考虑到了相邻斑块之间的植被空间结合在群落动态中的作用,而另外三种模型没有考虑到这一点;(2)在一定程度上,四种植物群落动态模型对各自针对的植物群落可能是适合的,但要作为描述群落动态发展的一般性模型还需要不断完善和发展;因为四种模型均没有考虑到自然干扰和人类干扰对植物群落动态的影响。作者对将来植物群落动态的研究及实践意义做出以下展望:(1)在不同空间尺度上,更加有效地评价控制群落动态变化的各种过程的相对重要性,并进一步将它们之间的复杂相互作用整合到群落动态模型中;(2)充分认识植物群落中存在的各种自然环境条件和生物群体的结构配置对植物群落动态发展的重要性;(3)重视植物群落动态发展中自然干扰过程和人类干扰过程的整合以     

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.     

Coastal sand dune vegetation of Tabasco and Campeche,Mexico

Abstract. Sand dune vegetation along the coast of the Mexican states of Tabasco and Campeche was sampled by means of 1501 releves and the data were analyzed with classification and ordination programs. 36 community types were distinguished, which were grouped into 10 higher-order groups. The communities are described in a sequence reflecting the main vegetation zonation. Beach, embryo dune and foredune community groups include 12 community types, the sheltered zone includes 16 types and the fixed dunes 8 types. The ordination results reveal one main vegetation gradient corresponding to the increase in mean distance from the shoreline and elevation, and being covariant with species richness, mean vegetation cover and mean vegetation height. Additional differences are related to the geographical transition between the Gulf and Caribbean coastal vegetation and environmental conditions, mainly type of sand, occurring in the study area. Siliceous sand and a Gulf climate are characteristic for most of Tabasco, and calcareous sand and a Caribbean-type climate are characteristic for Campeche. Disturbance caused by coconut plantations, cattle grazing and tourism favours the invasion of ruderal species from waste places behind the dunes, which attain local dominance in the dunes described here. Consequently some separate community types had to be distinguished. The distribution of community types is analyzed in the context of the transitions between calcareous and siliceous substrates in the study area.     

Functional spatial scale of community composition change in response to windthrow disturbance in a deciduous temperate forest

Community dynamics in local habitats are affected by landscape characteristics such as the area and connectivity of surrounding habitats at a functional spatial scale where the community responds to landscape structure. However, the functional spatial scale at which community composition is affected by landscape structure has never been explored. We assessed the functional spatial scales of composition change in birds and in three types of arthropod communities (canopy, forest-floor and flying ones) with regard to landscape heterogeneity resulting from a large typhoon in a temperate forest of Japan. We examined the effects of tree-fall disturbance on the communities at various spatial scales, with special attention to compositional evenness. The spatial scale of the best-fitting model, which was selected from models fitted to the disturbance area at stepwise spatial scales, was interpreted as the community-specific functional spatial scale. The composition of all communities studied was all significantly dependent on gap area. The functional spatial scale was highest in birds (370 m in radius), intermediate in flying arthropods (90 m) and lowest in canopy and forest-floor arthropods (10 m). This result may reflect typical dispersal ability and the spatial range of resource use in the community. Compositional changes in each community were consistent with theory regarding traits and responses of component taxa, although the enhancement of evenness was observed only in the arthropod communities. These results imply that management and reserve selection based on functional spatial scales can be effective in the conservation of biodiversity and ecosystem services at the community level.     

The relationship between vegetation dynamics and water table in tropical dune slacks

Abstract. Inundation of tropical dune slacks is an irregular phenomenon produced by consecutive years with high precipitation and also by intense tropical storms. Our hypothesis was that the distribution, structure and composition of vegetation in tropical dune slacks have changed over time as a result of various factors, principally their topographical position with respect to the water table, frequency and duration of inundation, and historical fluctuations in climate. We tested this hypothesis in two different slacks that represent the slack community at the study site. Inundation caused changes in species composition and distribution, as well as in vegetation structure. In one slack, inundation was considered as disturbance because its intensity increased when areas remained inundated for longer periods of time, causing death of vegetation cover and favouring invasion of more aggressive, secondary species (Pluchea odorata, Ambrosia artemisiifolia, Panicum maximum). When inundation lasted less than three months, rhizomatous herbs remained (Cyperus articulatus, Lippia nodiflora, Hydrocotyle bonariensis) inhibiting the succession process. The results suggest that the different degrees of inundation and the irregular frequency and intensity of inundation in the slacks under study, have favoured different successional tendencies.     

Patterns of abundance and co-occurrence in aquatic plant communities

Aquatic plants are well suited as subjects for studies on the distribution and abundance of co-occurring species, especially due to the simple structure of their communities, well defined toposequences and relatively easily measurable environmental factors. Here we show that underwater plants occurring in semi-natural lakes form stable communities, where species interactions dominate over dispersal dynamics to form a modular community structure with a high degree of zonation (turnover) and low within-module species richness. In turn, human-induced disturbance largely destroyed the modular structure. Our results indicate that (1) species abundance distributions (SADs) of underwater plant communities are well described by the lognormal model; (2) environmental characters did not significantly influence the SADs of underwater plant communities; (3) log-series SADs do not indicate specific types of community organization; (4) in our lake communities only few satellites (tourists) occur; (5) the co-occurrence of species is highly dependent on the turnover across lakes and water depth zones; and (6) species zonation is a function of lake properties.     

Population biology of salt marsh and sand dune annuals

Annuals represent a significant component of the vegetation of coastal salt marshes and sand dunes. From many points of view, the two habitats might appear to have little in common. Yet both are characterized by episodes of low water potential, marked spatial and temporal heterogeneity and a zonation which, within certain limits, reflects successional change.There are also similarities of distribution. Annuals are dominant usually in the pioneer stages; the Salicornia-dominated low marsh areas are perhaps analogues with strandline ephemeral populations (e.g. Cakile maritima) on the fore-dunes. In mature stages, annuals are associated with small gaps in the matrix of perennials, at least some of these arising from drought or disturbance. Nevertheless populations can reach very high densities.The most striking contrast is phenological; only summer annuals are found on marshes, whereas winter annuals predominate on dunes (except for the strandline). Similarly there is a difference in species richness. Rather few species of annual are typical of marshes while a great many are found on dunes.Properties of the seed bank, survival, reproduction and population regulation are compared in marsh and dune annuals, with special reference to Cakile, Salicornia, Rhinanthus and Vulpia. Interpretations are suggested which take account of environmental predictability and heterogeneity. Finally, the general applicability of simple mathematical models of these populations in the different coastal habitats is considered.Nomenclature follows Clapham, Tutin & Warburg (1981) except where otherwise stated.