Variation in species composition and vegetation structure of succulent scrub on Tenerife in relation to environmental variation

Abstract. On Tenerife, the occurrence of environmental gradients over short distances provides a unique opportunity to investigate the relationship between vegetation and environmental factors. In the semi‐arid coastal region of Tenerife, floristic composition, species richness and vegetation structure of perennial plants have been studied in 67 locations covering the existing precipitation gradient. On the island as a whole, variation in species composition could be best explained by mean annual precipitation; at coastal sites, substrate age and soil characteristics also played a significant role. On the other hand, substrate chemistry and the type of eruptive material explained little of the floristic variation. Stand biomass was strongly correlated with mean annual precipitation and was, on the youngest lava flows studied, also affected by substrate age. The native stem succulent species made up the bulk of total biomass along the whole precipitation gradient. Disturbed and undisturbed sites differed significantly in stand biomass and cover. Species richness was correlated with precipitation and substrate age. Distribution of plant functional types was also related to the precipitation gradient. The relative abundance of hemicryptophytes and shrubs with non‐hairy leaves increased with increasing precipitation whereas the ratio of shrubs with hairy/non‐hairy leaves and succulent plants decreased. Some alien plants were quite frequent at disturbed sites but, on the whole, they contributed little to the species spectrum and to the stand biomass. Undisturbed sites remained almost free of introduced species not considering annuals.     

Plant composition and diversity in a semi-natural Mediterranean island landscape: the importance of environmental factors

Abstract

Islands crucially contribute to the Mediterranean Basin’s high floristic diversity, which, however, is at risk facing climate and land-use changes. Besides the identification of highly diverse areas, the knowledge about factors favouring diversity is of great importance. We analysed plant species diversity and composition related to environmental factors over varied vegetation units on a former Italian prison island in the northwest of Sardinia. Due to a long history of land use with grazing and later abandonment the nowadays protected island features a semi-natural landscape and can serve as an example for strongly anthropogenic altered insular ecosystems. Floristic composition, soil properties, microclimate and ungulate abundance were assessed. Relationships of vegetation composition and diversity with abiotic variables were examined by Canonical Correspondence Analysis, which indicated the importance of air temperature, soil moisture, slope gradient and C/N ratio for floristic differentiation. Most important abiotic factors for plant species richness were relative air humidity and soil moisture, while floristic diversity was mainly determined by air temperature and pH. Furthermore, observation data pointed to an adverse influence of ungulate abundance for plant species diversity. Regarding nature conservation, grazing intensity thus must be critically taken into account, especially for sensitive vegetation units like the coastal garrigue.     

Air humidity,soil moisture and soil chemistry as determinants of the herb layer composition in European beech forests

Question: What role does air humidity play as an environmental factor for the abundance and distribution of temperate woodland herbs? Location: Beech forests on calcareous soils in southern lower Saxony, central Germany. Methods: The abundance of woodland herb species and total herb cover were investigated in 60 plots with contrasting exposure, slope angle and relief type. On all plots, air humidity, air temperature, soil moisture, photosynthetically active radiation, pH (H₂O) and concentration of salt‐exchangeable Ca, Mg and K were measured. Species‐environment relationships were analysed with multiple regression analysis and CCA. Results: Air humidity (RH), soil moisture and the concentration of exchangeable Ca and K, but not light, C/N ratio and the concentration of exchangeable Mg were identified as the most important abiotic factors influencing the cover of the most abundant plant species and total herb cover. RH varied substantially across the different forest floor site types and influenced species abundance independent of soil moisture. In several species (including Mercurialis perennis and Impatiens noli‐tangere), RH was found to be a key environmental factor. Other species such as Aegopodium podagraria and Lamiastrum galeobdolon depended more on elevated soil moisture, while RH was less important. Conclusions: This study showed that the distribution of widespread temperate woodland herb species depends on high air humidity, and that certain sensitive species do not occur at sites with reduced air humidity even though soil moisture is high. Thus, high air humidity and ample soil moisture are key abiotic factors in beech forests on calcareous soils. Shade level (PAR) was found to be of secondary importance.     

Floristic composition in relation to environmental gradients across KwaZulu‐Natal,South Africa

Conservation planning in the face of global change is still in its infancy. A suggested approach is to incorporate environmental gradients into conservation planning as they reflect the ecological and evolutionary processes generating and maintaining diversity. Our study provides a framework to identify the dominant environmental gradients determining floristic composition and pattern. Nonmetric multidimensional scaling was used on 2155 sampling plots in savanna and grassland habitat located across the province of KwaZulu‐Natal, South Africa (94 697 km²), a floristically rich region having steep environmental gradients, to determine the dominant gradients. Hierarchical cluster analysis was used to group similar plots which were then used in a Classification and Regression Tree analysis to determine the environmental delimiters of the identified vegetation clusters. Temperature‐related variables were the strongest delimiters of floristic composition across the province, in particular mean annual temperature. Frost duration was the primary variable in the Classification and Regression Tree analysis with important implications for savanna/grassland dynamics. Soil properties (base, pH status) and moisture variables accounted for most of the variation for the second and third axes of floristic variation. Given that climatic and edaphic variables were well correlated with floristic composition, it is anticipated that a changing climate will have a marked influence on floristic composition. We predict warmer temperatures may facilitate the spread of frost sensitive savanna species into previously cooler, grassland areas. Species associated with specific soil types will not easily be able to move up the altitudinal gradient to cooler climes because geology is aligned in an approximately north‐south direction compared with increasing altitude from east‐west. Future conservation planning should take cognisance of these gradients which are surrogates for ecological and evolutionary processes promoting persistence.     

CONTROL OF VEGETATION ON CONTRASTING SUBSTRATES: HERB PATTERNS ON SERPENTINE AND SANDSTONE

Binary discriminant analysis (BDA) reveals relationships between species distributions and environmental variables. In this study, BDA was applied to transects on serpentine and sandstone in the Wenatchee Mountains, Washington. Species presence and states of ten habitat variables were recorded in each quadrat. Species response patterns significantly different from a random expectation suggested that distributions are controlled by soil moisture far more strongly on serpentine than they are on sandstone, where light and its correlates are more important. Environmental patterns were explored with direct principal components analysis (PCA) of standardized environmental variables and indirectly by PCA of the significant D-values. The latter is biased towards variables to which species respond strongly. The results emphasize the environmental and floristic contrasts between substrates. On serpentine, direct PCA indicates that effective moisture, soil fertility, and insolation control the first three axes, while on sandstone, insolation, fertility, and effective moisture control the first three axes, respectively. The PCA of D-values (Q-mode) is similarly interpreted: moisture is strongly correlated with the first component on the serpentine transect, while insolation and fertility are correlated with the first component on sandstone. Species ordinations also result from Q-mode analysis. They reflect species responses to the identified gradients and indicate mutual species-environment interactions. These analyses conform to the hypothesis that mineral conditions of serpentine select against intolerant species and that survivors respond primarily to moisture conditions that are a result of low productivity, attributable to adverse nutrient conditions. On sandstone, the moisture gradient is less pronounced, and direct canopy effects that create variable light conditions predominate. BDA is a useful tool in ecological survey and pattern analysis. Species responses to environmental factors may be estimated quickly and directly. This method will help to focus subsequent research, improve experimental design, and generate explicit and testable hypotheses about species-habitat interactions.     

Application of multivariate methods to a study of community composition and structure in an escarpment woodland in northeast Ireland

Multivariate models based independently on the floristic and environmental attributes of sample quadrats from an escarpment woodland, were constructed. They were used to interpret the ecology of plant species distribution and to investigate the size structure of tree populations. Site instability, soil water status, and past management and grazing pressure were found to exert major influences on the composition and structure of the woodland. It is suggested that early mortality associated with site instability, plays an important role in maintaining the floristic diversity of vascular plants in the woodland ground flora, by increasing the frequency of tree replacement. This may have the effect of preventing canopy closure and any associated, detrimental effects of low light intensities. The relevance of the work of the conservation management of escarpment woods in Northern Ireland is discussed.     

Effects of coppicing on demographic structure, fruit and seed set in Orchis mascula

Opening of the forest canopy may result in higher fitness of understory plants due to increased light intensity on the forest soil and increased nutrient availability. Especially, tuberous orchids may profit from increased light as photosynthetic assimilation products and nutrients are accumulated in their belowground storage organs. We investigated the effects of coppicing on demographic structure and fruit and seed set in 15 populations of the tuberous, perennial orchid Orchis mascula, seven of which were located in undisturbed, shaded woodland and eight in recently coppiced woodland. Coppicing resulted in a massive increase in flowering and in increased fruit set. On average, 42.9% of all individuals consisted of flowering plants in coppiced woodland, whereas in undisturbed woodland only 20.8% of all individuals flowered. The percentage fruit set varied between 20% and 55% in coppiced woodland and was strongly related to population size (measured as the number of flowering individuals). Mean fruit weight and fruit size were also significantly higher in populations located in coppiced woodland than in undisturbed woodland, whereas the proportion of viable seeds was not related to woodland type. Given that O. mascula is dependent on seed set to fill space with new individuals, these results suggest that the long-term survival of this species strongly depends on frequent opening of the canopy.     

Environmental heterogeneity and interspecific interactions influence nest occupancy by key seed-dispersing ants

The complex interplay between species along environmental gradients ultimately shapes their distributions and additional community interactions. Ant-mediated seed dispersal fails in the wettest habitat of deciduous forest in eastern North America, and we examine whether this pattern corresponds with colony distributions for seed-dispersing ants and associated heterogeneity in abiotic and biotic variables. Specifically, we used spatial variation in soil moisture, temperature and diffuse light along natural habitat gradients and experimentally manipulated soil moisture gradients to examine ant habitat selection. We also examined niche segregation between effective (Aphaenogaster spp.) and ineffective (Lasius alienus Foerster) seed-dispersing ants across these environmental gradients. Whereas most research links ant foraging and nesting with temperature gradients, we find niche segregation between Aphaenogaster spp. and L. alienus by soil moisture along naturally occurring gradients and in experimentally irrigated upland habitat. The failure of Aphaenogaster spp. to occupy the wettest habitats, where L. alienus is present, is consistent with observed seed dispersal failure in these habitats. These results indicate that environmental heterogeneity drives niche segregation between effective (Aphaenogaster spp.) and ineffective (L. alienus) seed dispersers so each occupies distinct habitat. Most forest understory plants rely on ants for seed dispersal. Our research implies that climate-mediated interactions between effective and ineffective seed dispersing ant species may structure the microhabitat distributions for woodland herbs.     

Vascular plant diversity and climate change in the alpine zone of the Lefka Ori,Crete

The aim of this study is to analyse the vascular flora and the local climate along an altitudinal gradient in the Lefka Ori massif Crete and to evaluate the potential effects of climate change on the plant diversity of the sub-alpine and alpine zones. It provides a quantitative/qualitative analysis of vegetation-environment relationships for four summits along an altitude gradient on the Lefka Ori massif Crete (1664–2339 m). The GLORIA multi-summit approach was used to provide vegetation and floristic data together with temperature records for every summit. Species richness and species turnover was calculated together with floristic similarity between the summits. 70 species were recorded, 20 of which were endemic, belonging to 23 different families. Cretan endemics dominate at these high altitudes. Species richness and turnover decreased with altitude. The two highest summits showed greater floristic similarity. Only 20% of the total flora recorded reaches the highest summit while 10% is common among summits. Overall there was a 4.96°C decrease in temperature along the 675 m gradient. Given a scenario of temperature increase the ecotone between the sub-alpine and alpine zone would be likely to have the greatest species turnover. Southern exposures are likely to be invaded first by thermophilous species while northern exposures are likely to be more resistant to changes. Species distribution shifts will also depend on habitat availability. Many, already threatened, local endemic species will be affected first.     

Vegetation-environment relations of a Middle Zambezi floodplain

Detrended correspondence analysis was used to study the relationships between environmental factors and the species composition of vegetation on Zambezi River alluvium downstream of the Kariba hydroelectric dam. Grass, sedge and woody species were recorded in 73 stands in Mana Pools National Park, Zimbabwe. Grass and sedge species composition was related to the soil moisture regime, as indexed by soil texture and flooding frequency. The first woody plant ordination axis was related to a stand development gradient; Acacia albida was a pioneer species on lowlying sandbanks and the woody species richness of stands increased with their height above the Zambezi River. Two-way indicator species analysis identified 7 vegetation types which could be separated on the basis of their topsoil texture and flooding frequency. The types were: sandbanks; young A. albida woodland; A. albida woodland; A. albida dominated mixed woodland; mixed riverine woodland with understory; mixed riverine woodland; and grassland on clay soils.     

Variation in floristic and trait composition along environmental gradients in the herb layer of temperate forests in the transition zone between Central and SE Europe

Species- and trait-environment linkages in forest plant communities continue to be a frequent topic in ecological research. We studied the dependence of floristic and functional trait composition on environmental factors, namely local soil properties, overstory characteristics, climatic parameters and other abiotic and biotic variables. The study area comprised 50 monitoring plots across Slovenia, belonging to the EU ICP Forests monitoring network. Vegetation was surveyed in accordance with harmonized protocols, and environmental variables were either measured or estimated during vegetation sampling. Significant predictors of species composition were identified by canonical correspondence analysis. Correlations between plant traits, i.e. plant growth habit, life form, flowering features and CSR signature, were examined with fourth-corner analysis and linear regressions. Our results show that variation in floristic composition was mainly explained by climatic parameters (mean annual temperature, mean annual precipitation), soil properties (pH) and tree layer-dependent light conditions. Trait composition was most closely related with tree layer characteristics, such as shade-casting ability (SCA, a proxy for light availability in the understory layer), tree species richness and tree species composition. Amongst soil properties, total nitrogen content and soil texture (proportion of clay) were most frequently correlated with different species traits or trait states. The CSR signature of herb communities was associated with tree layer SCA, soil pH and mean annual temperature. The floristic composition of the studied herb-layer vegetation depended on temperature and precipitation, which are likely to be influenced by ongoing climate change (warming and drying). Trait composition exhibited significant links to tree layer characteristics and soil conditions, which are in turn directly modified by forest management interventions.

     

Variability in Mediterranean annual grassland diversity driven by small-scale changes in fertility and radiation

Using the small-scale gradients in light and fertility imposed by the presence of trees on Mediterranean open oak woodlands, we studied differences in the ecology, floristic diversity and species composition of an annual grassland, and modeled plant relationships with soil properties and radiation using generalized linear models. The strong intra-annual variability in the annual grassland greatly affected overall floristic diversity estimates. Therefore, sampling at least twice during the annual growing period was important to avoid underestimating floristic composition. As has been found in previous studies, different groups of grasses were clearly shaped as a response to the ecological gradient created by the influence of the tree. However, when analyzing specific relationships, only a few individual species directly responded to the studied soil and light availability factors analyzed. Among them, those species considered ‘ruderal’ such as Carduus tenuiflorus, Echium plantagineum, Lolium rigidum, Rumex bucephalophorus, and Tolpis barbata showed a clear response to some nutrients and to light availability but displayed no relationship to N soil content. Our results suggest that individual species are responding to a combination of ecological factors. Therefore, some of the generally accepted assumptions regarding the ecology of specific taxa may not be as straightforward as traditionally considered.     

Floristics and soil relations of upland swamp vegetation near Sydney

Abstract We examined relations between vegetation and soils, using multivariate methods, in hitherto poorly-known upland swamps on the Woronora Plateau, south of Sydney. A major trend in floristic composition was related to the height and cover of the herbaceous stratum and reflected a gradient in soil moisture and nutrients. A second trend in floristic composition was related to the height and cover of the shrub stratum, and may reflect the influence of recurring fires on certain dominant shrub species. Five plant communities were recognized on the basis of floristic composition and were distinguishable by their different soil habitats and/or structural characteristics. We report some of the highest species-richness values in the world (at scales of 1–15 m₂) for shrub/sedge-dominated vegetation, with up to 70 vascular plant species in 15 m2. Variation in species richness is inversely related to the resource gradient and positively related to the penetration of light through the vegetation canopy. This pattern is consistent with resource-competition models and warrants further investigation.     

Sandstone vegetation pattern in the Jim Jim Falls region,Northern Territory,Australia

Analysis of 58 floristic quadrats from two areas near Jim Jim Falls on the Arnhem Land Plateau revealed seven plant communities: wetland; monsoon forest; Allosyncarpia forest; broadleaf woodland; shrubby Eucalyptus woodland; Eucalyptus arnhemensis woodland; and sandstone scrub. The communities inter-grade with each other and form a complex mosaic. The patterning of vegetation is related to topography, rockiness, degree of fire protection and moisture supply. Surface soil nutrient concentrations were similar in all the communities with the exception of the monsoon forest which was substantially more fertile. We suggest that moisture supply is the primary determinant of vegetation and soil fertility patterns. Fire in the preceding dry season stimulated some herbaceous species, while a long unburnt shrubby Eucalyptus woodland was invaded by some monsoon forest seedlings. Conservation of the present mosaic of vegetation presents a major challenge to land managers because of this variable impact of fire.     

Floristic diversity patterns in the White Carpathians biosphere reserve,Czech Republic

The flora of the White Carpathians, a mountain range in the south-east of the Czech Republic, is documented by about 485,000 records of vascular plant occurrences collected since the mid-19^th century. A total of 1299 species recorded in 93 grid cells of 2.8 × 3.1 km were used for an analysis of spatial patterns of floristic diversity in the White Carpathians. Multivariate statistical techniques such as ordination and classification were used to reveal the main gradients in floristic composition and species richness, and measured environmental data and Ellenberg indicator values were used to assess underlying environmental factors. There is a striking floristic contrast between the western and eastern part of the study area, which is associated with differences in climate, mean altitude, topographic heterogeneity measured as altitudinal range, and land use. The western part is characterised by thermophilous, continental and calcicolous species of open habitats. In contrast, the more forested eastern part along the state border with Slovakia and the north-eastern part of the area are characterised by acidophilous species with higher moisture requirements. This pattern is consistent with the established phytogeographical division of the Czech Republic into the phytogeographical regions of Thermophyticum and Mesophyticum. The further division of the area into four regions, based on classified grid data, is also similar to the current division into phytogeographical districts, except for the Javorníky district. There are two distinct hot spots of species richness, in the western and the extreme north-eastern part. A poorer flora was found in landscapes with intensive agriculture. Species richness is associated with different environmental factors than species composition, namely with soil types and land-use categories. Alien species are more common in areas with a higher incidence of arable land and built-up areas, and less common in areas dominated by grasslands and forests.     

北方典型沼泽湿地高低土壤水分下植物群落特征

北方沼泽湿地在水源供给、缓解水土流失、遏制草地沙化等方面具有重要作用,明确其植物群落物种组成和多样性特征对提升其生态系统服务功能具有重要意义。目前,在北方地区开展大尺度湿地植被调查的研究仍相对较少。土壤水分是驱动植物群落发展的主导环境因素之一,为了解高低土壤水分背景下湿地植物群落特征差异及关键驱动要素,对我国7个北方典型沼泽湿地的植物群落物种组成及多样性特征进行了调查,分析了植物群落物种组成及多样性特征与环境因子的关系,以及沼泽湿地植物群落内克隆植物的分布特征。研究结果发现不同沼泽湿地的植物群落物种组成和多样性差异显著,但无明显的地带性分布规律,物种分布呈现区域性。群落物种多样性受降水、温度、土壤养分等多种环境因素的共同影响。沼泽湿地高低土壤水分背景下植物群落的物种组成和多样性差异显著,低土壤水分下植物群落物种多样性指数显著高于高土壤水分下植物群落。低土壤水分下物种多样性主要受降水和总氮影响,而高土壤水分下物种多样性主要受温度和总磷的影响。高土壤水分下克隆植物物种数和盖度在沼泽湿地植物中占有较高的比例,表明克隆植物比非克隆植物更适应高土壤水分环境。研究结果表明了7个沼泽湿地植被的区域性...     

Ecohydrological impacts of woody-plant encroachment: seasonal patterns of water and carbon dioxide exchange within a semiarid riparian environment

Across many dryland regions, historically grass‐dominated ecosystems have been encroached upon by woody‐plant species. In this paper, we compare ecosystem water and carbon dioxide (CO₂) fluxes over a grassland, a grassland–shrubland mosaic, and a fully developed woodland to evaluate potential consequences of woody‐plant encroachment on important ecosystem processes. All three sites were located in the riparian corridor of a river in the southwest US. As such, plants in these ecosystems may have access to moisture at the capillary fringe of the near‐surface water table. Using fluxes measured by eddy covariance in 2003 we found that ecosystem evapotranspiration (ET) and net ecosystem exchange of carbon dioxide (NEE) increased with increasing woody‐plant dominance. Growing season ET totals were 407, 450, and 639 mm in the grassland, shrubland, and woodland, respectively, and in excess of precipitation by 227, 265, and 473 mm. This excess was derived from groundwater, especially during the extremely dry premonsoon period when this was the only source of moisture available to plants. Access to groundwater by the deep‐rooted woody plants apparently decouples ecosystem ET from gross ecosystem production (GEP) with respect to precipitation. Compared with grasses, the woody plants were better able to use the stable groundwater source and had an increased net CO₂ gain during the dry periods. This enhanced plant activity resulted in substantial accumulation of leaf litter on the soil surface that, during rainy periods, may lead to high microbial respiration rates that offset these photosynthetic fluxes. March–December (primary growing season) totals of NEE were ?63, ?212, and ?233 g C m^?2 in the grassland, shrubland, and woodland, respectively. Thus, there was a greater disparity between ecosystem water use and the strength of the CO₂ sink as woody plants increased across the encroachment gradient. Despite a higher density of woody plants and a greater plant productivity in the woodland than in the shrubland, the woodland produced a larger respiration response to rainfall that largely offset its higher photosynthetic potential. These data suggest that the capacity for woody plants to exploit water resources in riparian areas results in enhanced carbon sequestration at the expense of increased groundwater use under current climate conditions, but the potential does not scale specifically as a function of woody‐plant abundance. These results highlight the important roles of water sources and ecosystem structure on the control of water and carbon balances in dryland areas.     

Quantifying the influence of environmental texture on the rate of species turnover: evidence from two habitats

The environmental texture hypothesis (ETH) proposes that the spatial geometry or texture of the environment influences the rate at which species are accumulated in space or time. Specifically, the ETH suggests that regions, and spatial scales, that exhibit a larger rate of environmental distance decay (DD) should exhibit more rapid rates of species turnover. The ETH should apply over any range of scales where the environment is driving species distributions. To examine the relevance of the ETH at local spatial scales, we tested for a positive relationship between the rate of change in soil chemical properties and vascular plant species composition in grassland and woodland habitats. We recorded presence–absence data along a 1.883 km transect in each habitat and estimated the rate of turnover and environmental DD for spatial lags of 1–41 m. We found that the soil environment explained spatial patterns of species composition more accurately in the grassland habitat compared to the woodland habitat. Consequently the rate of change in soil properties as a function of spatial distance was significantly positively correlated with the rate of species turnover in the grassland but not the woodland. Our study suggests that one of the central premises of the ETH is relevant for local patterns of species turnover if the environment appears to influence species composition.     

The comparative ecology of Polystichum aculeatum, P. braunii, and P. lonchitis in Hordaland, western Norway

The comparative ecology of the three species of Polystichum in Hordaland, western Norway, has been studied. Species frequency data and environmental variables were recorded in 132 sample units from 10 different localities. The numerical methods used to analyse these data are two-way indicator species analysis, detrended correspondence analysis, canonical correspondence analysis, and Gaussian logit regression. The main floristic gradient in the data is closely related to altitude. The second floristic gradient follows patterns in precipitation and soil pH. The ordination and classification results show that P. braunii is primarily a plant of deciduous forests, P. aculeatum occurs in open or woodland habitats near the coast, and P. lonchitis favours open habitats with cool summers at either low or high altitudes. The Gaussian logit regression results suggest that low altitude, short distance from the coast, and high soil pH may be important in influencing the growth of P. aculeatum , whereas P. braunii appears to depend most strongly on high soil depth and humidity and an adequate but not too extensive snow-cover in winter. P. lonchitis is found to favour high soil and air humidity, high light intensity, and not too high summer temperatures. The results contribute to our understanding of the local, national, and European distributions of the three species. In western Norway, conditions appear to be generally favourable for P. braunii and P. lonchitis , but are clearly less favourable for P. aculeatum