Plants living on gypsum: beyond the specialist model

BACKGROUND AND AIMS: Plants from gypsum habitats are classified as gypsophiles and gypsovags. The former include both narrow endemics limited to small gypsum areas and regionally dominant gypsophiles growing in gypsum areas of large regions, whereas gypsovags are plants that can grow both in gypsum and non-gypsum soils. Factors controlling the distribution of gypsum plants are still not fully understood. METHODS: To assess how the different types of gypsum plants deal with the stressful conditions of gypsum substrates, comparisons were made of the leaf chemical composition of four gypsovags, five regionally dominant gypsophiles and four narrow gypsum endemics growing in two massive gypsum areas of the Iberian Peninsula. KEY RESULTS: The chemical composition of gypsovags was clearly different from regionally dominant gypsophiles, while the chemical composition of narrow-gypsophile endemics was more similar to the chemical composition of gypsovags than to that of regionally dominant gypsophiles. Regionally dominant gypsophiles showed higher concentrations of ash, Ca, S, N, Mg P and Na, whereas gypsovags and local gypsophile endemics displayed higher concentrations of C and greater C : N ratios. CONCLUSIONS: Such differences suggest that the three groups of gypsum plants follow diverse ecological strategies. It is suggested that regionally dominant gypsophiles might fit the 'specialist' model, being species specifically adapted to gypsum, whereas both gypsovags and narrow-gypsophile endemics might fit the 'refuge' model, being stress-tolerant species that find refuge on gypsum soils from competition. The analysis of the leaf chemical composition could be a good predictor of the degree of plants specialization to gypsum soils.     

Gypsophile Chemistry Unveiled: Fourier Transform Infrared (FTIR) Spectroscopy Provides New Insight into Plant Adaptations to Gypsum Soils

Gypsum soils are among the most restrictive and widespread substrates for plant life. Plants living on gypsum are classified as gypsophiles (exclusive to gypsum) and gypsovags (non-exclusive to gypsum). The former have been separated into wide and narrow gypsophiles, each with a putative different ecological strategy. Mechanisms displayed by gypsum plants to compete and survive on gypsum are still not fully understood. The aim of this study was to compare the main chemical groups in the leaves of plants with different specificity to gypsum soils and to explore the ability of Fourier transform infrared (FTIR) spectra analyzed with neural network (NN) modelling to discriminate groups of gypsum plants. Leaf samples of 14 species with different specificity to gypsum soils were analysed with FTIR spectroscopy coupled to neural network (NN) modelling. Spectral data were further related to the N, C, S, P, K, Na, Ca, Mg and ash concentrations of samples. The FTIR spectra of the three groups analyzed showed distinct features that enabled their discrimination through NN models. Wide gypsophiles stood out for the strong presence of inorganic compounds in their leaves, particularly gypsum and, in some species, also calcium oxalate crystals. The spectra of gypsovags had less inorganic chemical species, while those of narrow gypsum endemisms had low inorganics but shared with wide gypsophiles the presence of oxalate. Gypsum and calcium oxalate crystals seem to be widespread amongst gypsum specialist plants, possibly as a way to tolerate excess Ca and sulphate. However, other mechanisms such as the accumulation of sulphates in organic molecules are also compatible with plant specialization to gypsum. While gypsovags seem to be stress tolerant plants that tightly regulate the uptake of S and Ca, the ability of narrow gypsum endemisms to accumulate excess Ca as oxalate may indicate their incipient specialization to gypsum.     

Plant life on gypsum: a review of its multiple facets

The adaptation of plants to particular soil types has long intrigued biologists. Gypsum soils occupy large areas in many regions of the world and host a striking biological diversity, but their vegetation has been much less studied than that developing over serpentine or saline soils. Herein, we review all aspects of plant life on gypsum ecosystems, discuss the main processes driving their structure and functioning, and highlight the main conservation threats that they face. Plant communities in gypsum habitats typically show distinctive bands at very small spatial scales, which are mainly determined by topography. Plants living on gypsum soils can be classified into three categories: (i) wide gypsophiles are specialists that can penetrate the physical soil crust during early life stages and have physiological adjustments to cope with the chemical limitations imposed by gypsum soils; (ii) narrow gypsophiles are refugee plants which successfully deal with the physical soil crust and can tolerate these chemical limitations but do not show specific adaptations for this type of soils; and (iii) gypsovags are non‐specialist gypsum plants that can only thrive in gypsum soils when the physical crust is absent or reduced. Their ability to survive in gypsum soils may also be mediated by below‐ground interactions with soil microorganisms. Gypsophiles and gypsovags show efficient germination at low temperatures, seed and fruit heteromorphism within and among populations, and variation in seed dormancy among plants and populations. In gypsum ecosystems, spatio‐temporal changes in the composition and structure of above‐ground vegetation are closely related to those of the soil seed bank. Biological soil crusts (BSCs) dominated by cyanobacteria, lichens and mosses are conspicuous in gypsum environments worldwide, and are important drivers of ecosystem processes such as carbon and nitrogen cycling, water infiltration and run‐off and soil stability. These organisms are also important determinants of the structure of annual plant communities living on gypsum soils. The short‐distance seed dispersal of gypsophiles is responsible for the high number of very narrow endemisms typically found in gypsum outcrops, and suggests that these species are evolutionarily old taxa due to the time they need to colonize isolated gypsum outcrops by chance. Climate change and habitat fragmentation negatively affect both plants and BSCs in gypsum habitats, and are among the major threats to these ecosystems. Gypsum habitats and specialists offer the chance to advance our knowledge on restrictive soils, and are ideal models not only to test important evolutionary questions such as tolerance to low Ca/Mg proportions in soils, but also to improve the theoretical framework of community ecology and ecosystem functioning.     

Modularity Reveals the Tendency of Arbuscular Mycorrhizal Fungi To Interact Differently with Generalist and Specialist Plant Species in Gypsum Soils

Patterns in plant–soil biota interactions could be influenced by the spatial distribution of species due to soil conditions or by the functional traits of species. Gypsum environments usually constitute a mosaic of heterogeneous soils where gypsum and nongypsum soils are imbricated at a local scale. A case study of the interactions of plants with arbuscular mycorrhizal fungi (AMF) in gypsum environments can be illustrative of patterns in biotic interactions. We hypothesized that (i) soil characteristics might affect the AMF community and (ii) there are differences between the AMF communities (modules) associated with plants exclusive to gypsum soils (gypsophytes) and those associated with plants that show facultative behavior on gypsum and/or marly-limestone soils (gypsovags). We used indicator species and network analyses to test for differences between the AMF communities harbored in gypsophyte and gypsovag plants. We recorded 46 operational taxonomic units (OTUs) belonging to nine genera of Glomeromycota. The indicator species analysis showed two OTUs preferentially associating with gypsum soils and three OTUs preferentially associating with marly-limestone soils. Modularity analysis revealed that soil type can be a major factor shaping AMF communities, and some AMF groups showed a tendency to interact differently with plants that had distinct ecological strategies (gypsophytes and gypsovags). Characterization of ecological networks can be a valuable tool for ascertaining the potential influence of above- and below-ground biotic interactions (plant-AMF) on plant community composition.     

Centaurea subsect. Phalolepis in Southern Italy: ongoing speciation or species overestimation? Genetic evidence based on SSRs analyses

In this paper, we investigated a set of narrow endemics of Centaurea subsect. Phalolepis from the mountains of South Italy (mainly Calabria and Salento), segregated from the widespread species Centaurea deusta, using microsatellite (SSR) markers. The goal was to analyse the genetic makeup (levels and structure) of C. deusta and the segregated species and verify whether genetic clusters were in agreement with current classification of the species. With C. deusta, we also carried out an ecological niche modelling (ENM) analysis to check its potential distribution under present climatic conditions and to project it to the Last Glacial Maximum (LGM). As also found in former studies with subsect. Phalolepis in Greece and Turkey using the same set of SSRs, genetic diversity for the segregated Italian species was higher than expected for narrow endemics with small populations. Genetic clusters, however, were not correlated with the described species and did not support the segregation of the purported narrow endemics from a widely defined C. deusta. The results of the ENM indicate that the Adriatic Sea was a migration corridor for C. deusta at the LGM.     

Fragment size does not matter when you are well connected: effects of fragmentation on fitness of coexisting gypsophiles

Most habitat fragmentation studies have focused on the effects of population size on reproductive success of single species, but studies assessing the effects of both fragment size and connectivity, and their interaction, on several coexisting species are rare. In this study, we selected 20 fragments along two continuous gradients of size and degree of isolation in a gypsum landscape in central Spain. In each fragment, we selected 15 individuals of each of three dominant gypsophiles (Centaurea hyssopifolia, Lepidium subulatum and Helianthemum squamatum, 300 plants per species, 900 plants in total) and measured several reproductive traits: inflorescence number, fruit set, seed set and seed mass. We hypothesised that plant fitness would be lower on small and isolated fragments due to an interaction between fragment size and connectivity, and that response patterns would be species‐specific. Overall, fragment size had very little effect on reproductive traits compared to that of connectivity. We observed a positive effect of fragment connectivity on C. hyssopifolia fitness, mediated by the increased seed predation in plants from isolated fragments, resulting in fewer viable seeds per capitulum and lower seed set. Furthermore, seed mass was lower in plants from isolated fragments for both C. hyssopifolia and L. subulatum. In contrast, few reproductive traits of H. squamatum were affected by habitat fragmentation. We discuss the implications of species‐specific responses to habitat fragmentation for the dynamics and conservation of gypsum plant communities. Our results highlight the complex interplay among plants and their mutualistic and antagonistic visitors, and reinforce the often‐neglected role of habitat connectivity as a key component of the fragmentation process.     

Additive effects of a potentially invasive grass and water stress on the performance of seedlings of gypsum specialists

Question: What is the combined effect of two drivers of local biodiversity changes (presence of a potentially invasive species and seasonal drought) on the performance of seedlings of plants from gypsum habitats under experimental conditions? Location: A controlled microcosm reconstruction of natural assemblages of gypsum plant communities from central Spain. Methods: We evaluated the effects of a potentially invasive grass (Lolium rigidum) and water stress on the survival, height growth and biomass of five woody species (Colutea hispanica, Gypsophila struthium, Thymus lacaitae, Lepidium subulatum and Helianthemum squamatum) from semi‐arid gypsum ecosystems. Seedlings of the five species were grown with or without the potential invader and under three watering regimes: early stress — simulating an advanced summer, late stress — simulating the characteristic timing of current summer drought and well‐watered. Results: Seedling survival and performance were negatively affected by the presence of the potential invader. Early stress had larger impacts on the gypsum species than late stress. No interactions were found between factors for any of the study variables, and responses to both factors were found to be species‐specific. Conclusions: The lack of interactions between factors indicates that the presence of the potentially invasive grass and water stress had additive effects in our study system. The negative impact of early water stress draws attention to the possible consequences of the advances of summer drought predicted for Mediterranean ecosystems. Finally, the differential responses found for the study species suggest that plant communities will not respond as a unit to global change, leading to significant changes in species composition and dominance.     

Gypsophile vegetation patterns under a range of soil properties induced by topographical position

Iberian gypsophile plant communities are considered a priority for conservation by the European Community because of their highly specialized flora in gypsum outcrops in arid and semiarid regions. Despite the ecological importance of these ecosystems, the edaphic factors that constrain plant communities on gypsiferous soils remain unclear. It has been proposed that both the chemical and physical restrictive conditions of gypsum soils determine gypsophily in plants. Here we hypothesize that the rigors of the gypsum soil environment depends on topography, decreasing from flat areas on hilltops to south-oriented slopes and finally to slopes oriented to the north. We also hypothesized that the relaxation of the rigors of the gypsum soil environment with topography affects both to individual plant and community characteristics of gypsophile vegetation: we expect a reduction of gypsophyte abundance, an increase of diversity and the amelioration of facilitative interactions of plant species. We analysed the physical and chemical properties of gypsum soils that have been proposed that determine the rigors of the gypsum soil environment (i.e.: unbalanced ion concentrations and superficial soil crust). The predicted rigor gradient along topographical locations was confirmed and was mainly caused by superficial soil crust. The decreasing rigor gradient was accompanied by a fall in the abundance of gypsophytes. However, when gypsophytes were considered separately, several patterns were observed, indicating distinct tolerance to relaxation of rigor of the gypsum soil conditions and different competition abilities between gypsophytes. Plant species were more clumped, and gypsophile communities presented higher diversity, evenness and richness values where rigor of gypsum soil conditions were maximum (flat hilltop positions). Relaxation of rigor (north-oriented slopes) was characterized by loss of facilitative interaction between species and the dominance of the gypsovag Rosmarinus officinalis L., although richness was still very high, which can be attributed to the coexistence of gypsophytes and gypsovags. We conclude that the rigor of gypsum soil environment gradient with topography is mainly determined by superficial soil crust, and it is a crucial determinant of gypsophile plant communities.     

Influence of Environmental Factors on Essential Oil Variability in Origanum compactum Benth. Growing Wild in Morocco

The present study aimed to evaluate the influence of environmental factors on essential oils (EOs) composition of Origanum compactum populations sampled all over the distribution area of the species in Morocco, and to determine the extent of the chemical profiles throughout the geographical distribution of the species. The chemical compositions were submitted to canonical correlation analysis and canonical discriminant analysis that indicated a significant relationship between oil components and some environmental factors. According to their chemical composition and edapho‐climatic characteristics, two major groups of populations were differentiated. The first group was composed of samples growing in regions with humid climate, clayey, sandy, and alkaline soils. These samples showed high thymol, α‐terpineol, linalool, and carvacryl methyl oxide content. The second group consisted of plants belonging to semi‐arid climate, and growing at high altitudes and silty soils. These samples were characterized by high carvacrol, α‐thujene, α‐terpinene, and myrcene content. However, populations exposed to sub‐humid climate, appeared less homogeneous and belong mainly either to the first or second group. A significant correlation between some edaphic factors (pH, K₂O content, soil texture) and the EOs yield of O. compactum plants was evidenced. In spite of the correlation obtained for the oil composition with edapho‐climatic factors and the variance explained by the environmental data set, the observed EO diversity might be also genetically determined.     

Soil–plant relationship along a semiarid gypsum gradient (Rio de Aguas,SE Spain)

Gypsum outcrops of southeastern Spain (Almeria) have been highlighted as the most outstanding for the conservation of Iberian gypsum flora by flora rarity and richness, as vascular as cryptogamic plants. However, plant community distribution patterns according to soil chemical properties have been little studied in these gypsum areas. Spatial distribution pattern of plant communities in gypsum hills and its relation to soil chemical properties was surveyed in this study. Twenty-one plots (5 × 5 m) were settled along a semiarid gypsum gradient in Rio de Aguas Basin. Soil samples were taken from each plot’s superficial layer for chemical analysis. Plant canopy cover was sampled at species level. Three plant community bands are identified (from bottom to top) as level I (Flat Piedmont Zone), level II (Hill Slope Zone), and level III (Hill Top Zone). Gypsophyte species (mainly found in level II) appear to be specifically adapted to nutrient-stressed environments (high sulfate content and deficiency in some soil nutrients). Nutrients play an essential ecological role in determining species distribution and community composition. Since this area is a very important site for extracting very high quality gypsum, the pattern described here can be used as a useful tool for ecological restoration of gypsum quarries. Considering environmental heterogeneity of gypsum areas (as an “ecosystem of reference”) is crucial for a successful ecological restoration.     

Disentangling water sources in a gypsum plant community. Gypsum crystallization water is a key source of water for shallow-rooted plants

Background and AimsGypsum drylands are widespread worldwide. In these arid ecosystems, the ability of different species to access different water sources during drought is a key determining factor of the composition of plant communities. Gypsum crystallization water could be a relevant source of water for shallow-rooted plants, but the segregation in the use of this source of water among plants remains unexplored. We analysed the principal water sources used by 20 species living in a gypsum hilltop, the effect of rooting depth and gypsum affinity, and the interaction of the plants with the soil beneath them.MethodsWe characterized the water stable isotope composition, δ ²H and δ ¹⁸O, of plant xylem water and related it to the free and gypsum crystallization water extracted from different depths throughout the soil profile and the groundwater, in both spring and summer. Bayesian isotope mixing models were used to estimate the contribution of water sources to plant xylem sap.Key ResultsIn spring, all species used free water from the top soil as the main source. In summer, there was segregation in water sources used by different species depending on their rooting depth, but not on their gypsum affinity. Gypsum crystallization water was the main source for most shallow-rooted species, whereas free water from 50 to 100 cm depth was the main source for deep-rooted species. We detected plant–soil interactions in spring, and indirect evidence of possible hydraulic lift by deep-rooted species in summer.ConclusionsPlants coexisting in gypsum communities segregate their hydrological niches according to their rooting depth. Crystallization water of gypsum represents an unaccounted for, vital source for most of the shallow-rooted species growing on gypsum drylands. Thus, crystallization water helps shallow-rooted species to endure arid conditions, which eventually accounts for the maintenance of high biodiversity in these specialized ecosystems.     

Chemical transformation of Kamchatka soils after input of products of volcanic eruption

It has been revealed that changes in the chemical composition of soils after the input of volcanic ash under conditions of a cold humid climate depend on granulometric and lithogeochemical compositions of eruption products, the ways and forms of the input of chemical elements to soil, the period of the year, and weather conditions at ash falls. The lithogeochemical composition of ash mainly determines the type of change in the total composition of soils. Variations in the contents of mobile forms of elements depend mainly on the period of year and meteorological conditions during ash falls. Most mobile compounds (99.3%) of many elements are transported to soil upon ash falls not as components of ash particles, but in gas- and watersoluble forms directly from the dispersion medium of the volcanic aerosol. The following regularities were revealed in soils at a distance of 25 km from the centers of eruption: (i) the increase in the total contents of elements and the removal of their mobile forms in winter and (ii) the accumulation of mobile compounds, a drop in their total compounds, and soil acidification with a significant (about 1.5 times) decrease in the degree of base saturation of soils in summer.     

Patterns and traits of the endemic plants of Greece

Greece is characterized by high plant diversity (5800 species) and endemism (15.6%). This study attempts a first overall assessment of the taxonomy, distribution, traits and conservation status of the Greek endemic plants. The endemic species belong to 56 families and 242 genera. Most of the endemic plants have a narrow geographical and altitudinal distribution range. The southern floristic regions are richer in endemic species. The species area relationships for endemics (EARs) for island and continental floristic regions explain over 50% of the variation in number of species and are characterized by steep curves. Analysis of the distributional pattern of the endemics by similarity coefficients offers useful insights into the palaeogeography and biogeography of Greece. The endemic species occur at all altitudes, but the altitudinal distribution shows a predominance of local endemics at 0–600 m in the island regions and in higher zones in the continental regions. The life form spectra show a predominance of hemicryptophytes and chamaephytes. This trait seems indicative of their habitat and adaptive strategy and may be related to speciation processes. The overview of the conservation status of the Greek endemics indicates that over 40% of the taxa are threatened or near threatened. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 162 , 130–422.     

The diversity of life-form type, habitat preference and phenology of the endemics in the Goa region of the Western Ghats, India

Aim To carry out (1) a floristic survey of endemic flowering plants of the Western Ghats occurring in Goa, (2) identify their habitat preference and diversity of life‐form type, (3) observe flowering phenology of the endemics and (4) to correlate factors that affect their phenological pattern. Location Goa state is located between 15°48′ N and 14°53′54′′ N and 74°20′13′′ E and 73°40′33′′ E, in the northern part of the Western Ghats, India. Method A list of endemic plants from the study area was prepared using available floristic works and checklists of endemic plants of India. Based on preliminary field observations carried out in the study area, major habitats such as plateaus, moist deciduous forests, semi‐evergreen forests, evergreen forests and mangroves were identified for the subsequent intensive survey of endemics. Voucher specimens for all the endemic species were collected, processed using conventional herbarium techniques and deposited in the Herbarium, Department of Botany, Goa University. Species were identified using local and regional floras and their identity was confirmed at various herbaria along the Western Ghats. Data on their life‐form types, habitat and phenology was recorded in the field. Phenological observations were made every fortnight. A computerized data base was generated incorporating details on their life‐form type, phenology and habitat. Results A floristic survey of endemic plants of the Western Ghats in Goa resulted in the collection of 113 endemic species. Life‐form analysis reveals that herbaceous endemics are the most dominant followed by trees, shrubs and climbers. Plateaus in the study area harbour the largest number of endemic species, especially herbs. Endemic trees are distributed in the semi‐evergreen and evergreen forests. Endemic species in the study area show different peak and lean seasons of flowering depending on their life‐form type, habitat and ecological factors like temperature and rainfall/moisture content in the soil. Main conclusions The plateaus in the northern part of the Western Ghats are unique, being species rich with herbaceous endemics. These ephemerals are closely associated with the rainfall patterns thus; any change of moisture regime over the long‐term will have an impact on the distribution of these endemics.     

Distribution and abundance of trees in floodplain forests of the Wisconsin River: Environmental influences at different scales

Questions: 1. How do physiography, flooding regime, landscape pattern, land‐cover history, and local soil conditions influence the presence, community structure and abundance of overstorey trees? 2. Can broad‐scale factors explain variation in the floodplain forest community, or are locally measured soil conditions necessary? Location: Floodplain of the lower 370 km of the Wisconsin River, Wisconsin, USA. Methods: Floodplain forest was sampled in 10 m × 20 m plots [n= 405) during summers of 1999 and 2000 in six 12‐ to 15‐km reaches. Results: Species observed most frequently were Fraxinus pennsylvanica, Acer saccharinum and Ulmus americana. Physiography (e.g. geographic province) and indicators of flooding regime (e.g. relative elevation and distance from main channel) were consistently important in predicting occurrence, community composition, and abundance of trees. Correspondence analysis revealed that flood‐tolerant and intolerant species segregated along the primary axis, and late‐successional species segregated from flood‐tolerant species along the secondary axis. Current landscape configuration only influenced species presence or abundance in forests that developed during recent decades. Land‐cover history was important for tree species presence and for the abundance of late‐successional species. Comparison of statistical models developed with and without soils data suggested that broad‐scale factors such as geographic province generally performed well. Conclusions: Physiography and indicators of flood regime are particularly useful for explaining floodplain forest structure and composition in floodplains with a relatively high proportion of natural cover types.     

The microbe–mineral environment and gypsum neogenesis in a weathered polar evaporite

Evaporitic deposits are a globally widespread habitat for micro‐organisms. The microbe–mineral environment in weathered and remobilized gypsum from exposed mid‐Ordovician marine evaporite beds in the polar desert of Devon Island, Nunavut, Canadian High Arctic was examined. The gypsum is characterized by internal green zones of cyanobacterial colonization (dominated by Gloeocapsa/Aphanothece and Chroococcidiopsis spp. morphotypes) and abundant black zones, visible from the surface, that contain pigmented cyanobacteria and fungi. Bioessential elements in the gypsum are primarily provided by allochthonous material from the present‐day polar desert. The disruption, uplift and rotation of the evaporite beds by the Haughton meteorite impact 39 Ma have facilitated gypsum weathering and its accessibility as a habitat. No cultured cyanobacteria, bacteria and fungi were halophilic consistent with the expectation that halophily is not required to persist in gypsum habitats. Heterotrophic bacteria from the evaporite were slightly or moderately halotolerant, as were heterotrophs isolated from soil near the gypsum outcrop showing that halotolerance is common in arctic bacteria in this location. Psychrotolerant Arthrobacter species were isolated. No psychrophilic organisms were isolated. Two Arthrobacter isolates from the evaporite were used to mediate gypsum neogenesis in the laboratory, demonstrating a potential role for microbial biomineralization processes in polar environments.     

A preliminary account of patterns of endemism in Namibia's Sperrgebiet – the succulent karoo

Aim This study investigated spatial patterns of endemism in the flora of Namibia's succulent karoo in order to generate information for conservation planning. Location The study area, the Sperrgebiet, comprises the majority of Namibia's portion of the succulent karoo biome which is the south‐west corner of the country. This is an arid area that has been off limits to public access, farming and tourism for nearly a century due to restrictions imposed by the diamond industry. Methods Based on existing distribution records, areas of high concentrations of endemic plants were identified using numbers of endemics and weighted endemics according to area of occupancy. The resolution of the available data was quarter degree squares (15‐min intervals of latitude and longitude grids). Results At the scale of this study straight numbers of endemics generated similar results to the endemics weighted according to area of occupancy, which gives sparsely distributed species a higher weighting. Based on the current distribution records, 17.7% (184 species) of the Sperrgebiet's spermatophyte flora is endemic. The ‘hotspots of endemism’ comprised from north to south: Lüderitz‐Kowisberge, Klinghardt Mountains, Aurusberge‐Heioab, Witpütz, Skorpion and Obib‐Schakalsberge. Taking also areas into account that stand out because of their high proportion of local endemics, this adds Grillental and the central coastal area from Pomona to Baker's Bay to the areas of importance for plant endemism. Main conclusions The Sperrgebiet's endemic flora is special in taxonomic composition in that it does not present a subset of the total flora of this area, but shows a remarkably high representation of the families Mesembryanthemaceae and Liliaceae (sensu lato). Compared with other arid areas, the level of endemism in the Sperrgebiet is high, but not compared with the succulent karoo in general or other hotspots in the succulent karoo biome, such as the Richtersveld. The proportion of local endemics (13.5%) is high compared with some endemism hotspots in southern Africa. Hotspots of plant endemism provide an important tool to contribute to conservation planning studies. This study also highlighted the importance of centralized data bases without which these analyses would not have been possible. Further plant collecting is required to fill presently data‐deficient areas and studies at a finer spatial resolution taking habitat requirements into account are needed to elucidate some of the factors contributing to plant endemism in this area.     

Effects of experimental water addition depend on grassland community characteristics

Plant community biomass and composition on low-productivity soils, such as serpentine, may be more resistant to climate change because they host stress-tolerant species that may respond slowly to change. These communities also host a number of endemic taxa that are of special interest because of their narrow distributions. In a 3-year study, we experimentally tested the response of serpentine and non-serpentine communities to water addition in spring. We also compared the responses of endemics and generalists to water addition, with and without biomass (competitor) removal. In the non-serpentine grassland, peak biomass was significantly greater in the water addition plots compared with control plots, but this effect depended on the year. In the serpentine grassland, there was no effect of water addition on biomass. Survival, biomass, growth rates, and seed production of soil endemics and generalists were all significantly reduced by competition, but were unaffected by water addition. Overall, endemics tended to perform better in serpentine soil and generalists in non-serpentine soil, suggesting that soil is an important factor for the establishment and survival of endemics and generalists. For endemics, the effect of biomass removal was stronger on non-serpentine soil, but for generalists this effect was similar on both soils, indicating that competition can be important in low-resource habitats. In conclusion, our results suggest that low-fertility plant communities may be slow to respond to changes in precipitation compared to communities on more fertile soil.     

Patterns of endemism of the eastern North American cave fauna

Aim Over 250 species of obligate terrestrial cave‐dwelling animals (troglobionts) are known from single caves in the eastern United States. We investigate their geographical distribution, especially in relation to other troglobionts. We relate these patterns to taxonomic group, opportunities for dispersal and geographical location. Location Caves of the United States east of the Mississippi River. Methods We associated over 3000 records of more than 450 troglobiotic species and subspecies with hexagons of 1000, 5000 and 10,000 km² in size. We calculated Moran's I, black–white joins and cubic regression of endemics on non‐endemics at all three spatial scales. For 5000 km² hexagons, we modelled the spatial autocorrelation of the residuals of the cubic regression of endemics on non‐endemics. Results Differences among orders in percentage single‐cave endemism were not significant, except for Pseudoscorpionida, which was higher (69%) than any other order. At all three scales, Moran's I and black–white joins were significant, indicating a clumped distribution of both single‐cave endemics and other troglobionts. Spatial patterns were similar at all three scales and Moran's I was highest at 5000 km². The cubic fit of endemics to non‐endemics was consistently better, with less systematic error or residuals, than were linear or quadratic models. Residuals showed a significant geographical pattern with excess endemics in more southerly locations. Main conclusions There was both a non‐spatial and spatial component to the pattern of single‐cave endemism. The non‐spatial component was the association of high levels of single‐cave endemism with areas of high diversity of non‐endemics. It may be that both are high because of high secondary productivity. Spatially, single‐cave endemism is high in central rather than peripheral areas and in the southern part of the range. It is not higher in areas of more dissected limestone, which would reduce migration rates; if anything endemism is lower. Regional spatial effects are important, indicating that cave communities cannot be understood (or protected) in isolation.