Floristic diversity of an eastern Mediterranean dwarf shrubland: the importance of soil pH

Questions: Does plant species richness and composition of eastern Mediterranean dwarf shrubland (phrygana) correlate with soil pH? How important is the effect of pH on species diversity in relation to other environmental factors in this ecosystem? What is the evolutionary background of the diversity–pH relationship? Location: Western Crete, Greece. Methods: Species composition of vascular plants, soil and other environmental variables were sampled in 100‐m² plots on acidic and basic bedrock in phrygana vegetation. The relationships between species composition and environmental variables (including climate) were tested using canonical correspondence analysis, and relationships between species richness and environment using correlation and regression analyses. Data were analysed separately for different plant functional types based on life form and life span. Results: Although soil pH varied across a narrow range (5.9‐8.1), species composition changed significantly along the pH gradient within all plant functional types. For most functional types, the effect of soil pH on species composition was stronger than that of other environmental variables. Species richness of annuals, geophytes and suffruticose chamaephytes increased with soil pH, while richness of hemicryptophytes and shrubs was not correlated with pH. Conclusions: The results are consistent with the evolutionary species pool hypothesis. High numbers of calcicole annuals, geophytes and suffruticose chamaephytes may be a result of the evolution of these groups on base‐rich dry soils in the Mediterranean climate. In contrast, hemicryptophytes, a life form typical of the temperate zone, evolved on both acidic and basic soils and therefore their species numbers do not respond to soil pH across the narrow range studied. The lack of a relationship between shrub species richness and pH is difficult to explain: it may reflect the more diverse or older origin of Mediterranean woody species and their conservative niches.     

The importance of Anatolian mountains as the cradle of global diversity in Arabis alpina, a key arctic-alpine species

Background and Aims

Anatolia is a biologically diverse, but phylogeographically under-explored region. It is described as either a centre of origin and long-term Pleistocene refugium, or as a centre for genetic amalgamation, fed from distinct neighbouring refugia. These contrasting hypotheses are tested through a global phylogeographic analysis of the arctic–alpine herb, Arabis alpina.

Methods

Herbarium and field collections were used to sample comprehensively the entire global range, with special focus on Anatolia and Levant. Sequence variation in the chloroplast DNA trnL-trnF region was examined in 483 accessions. A haplotype genealogy was constructed and phylogeographic methods, demographic analysis and divergence time estimations were used to identify the centres of diversity and to infer colonization history.

Key Results

Fifty-seven haplotypes were recovered, belonging to three haplogroups with non-overlapping distributions in (1) North America/Europe/northern Africa, (2) the Caucuses/Iranian Plateau/Arabian Peninsula and (3) Ethiopia–eastern Africa. All haplogroups occur within Anatolia, and all intermediate haplotypes linking the three haplogroups are endemic to central Anatolia and Levant, where haplotypic and nucleotide diversities exceeded all other regions. The local pattern of haplotype distribution strongly resembles the global pattern, and the haplotypes began to diverge approx. 2·7 Mya, coinciding with the climate cooling of the early Middle Pleistocene.

Conclusions

The phylogeographic structure of Arabis alpina is consistent with Anatolia being the cradle of origin for global genetic diversification. The highly structured landscape in combination with the Pleistocene climate fluctuations has created a network of mountain refugia and the accumulation of spatially arranged genotypes. This local Pleistocene population history has subsequently left a genetic imprint at the global scale, through four range expansions from the Anatolian diversity centre into Europe, the Near East, Arabia and Africa. Hence this study also illustrates the importance of sampling and scaling effects when translating global from local diversity patterns during phylogeographic analyses.     

Comparative taxonomic structure of the floras of two Mediterranean-climate regions: Iberia and California

Two major events were invoked to understand recent biodiversity patterns in Mediterranean floras: northern hemisphere glaciations and historical human impacts. These two events were considered in this work, where we investigated general patterns in plant species richness and rarity attributes in two different Mediterranean regions: California and Iberia. Our goal was to assess whether comparisons of this sort provided evidence of different extinctions rates, making an effort to decouple anthropogenic from ice age‐related effects in both regions. We employed a taxonomically revised database for eight Mediterranean floras containing information on species richness for 298 families and rarity attributes for 11,834 taxa. We used summary statistics (Gini coefficient) and randomly generated models to test for general patterns of the distribution of diversity within and among taxonomic groups. We then used this general pattern among Mediterranean floras to provide a context in which to evaluate our two focal areas. Results indicated that floras of California and Iberia share the closest taxonomic structure among Mediterranean regions. Differences emerged in rarity attributes and the taxonomic identities of rarity rich groups. These findings were interpreted in the light of Pleistocene changes. In addition, a closer focus on rarity attributes allowed us to pinpoint some segments of these floras where anthropogenic activities may drive variation from general patterns, specifically for rare species in ecologically sensitive habitats.     

Seasonal fires shape the germinable soil seed bank community in eastern Mediterranean woodlands

季节性火烧塑造了东地中海地区林地的可萌发种子库群落 本研究探索了在典型东地中海林地中，季节性火烧是否以及如何与微生境类型(即黄连木属(Pistacia)灌木下、岩蔷薇属(Cistus)灌木下或林冠间隙)相互作用，从而影响可萌发种子库(germinable soil seed bank, GSSB)群落的组成。我们进行了一项野外实验，其中包括预定的春秋火烧，然后利用从火烧地块和相邻未火烧对照地收集的土壤样本对种子萌发的模式进行定量。研究结果表明，秋烧期间，土壤温度比春烧的要高，而春烧期间土壤温度则比秋烧的变化更大。火烧导致可萌发种子库整体的密度、丰富性和多样性降低。秋烧地块内黄连木属和岩蔷薇属灌木的可萌发种子库丰富性和多样性显著降低，这些模式主要表现在一年生植物中。在火烧样地收集的样本中，低矮灌木的可萌发种子库密度更高，在黄连木属和岩蔷薇属灌木下采集的样品中，这种模式更为明显。在季节性火烧与出现的独特物种会共同导致可萌发种子库的群落组成发生显著的变化。这些结果表明，季节性火烧与空间异质性互相作用，主要通过对一年生植物和低矮灌木的可萌发密度的差异效应来影响可萌发种子库群落的组成。这些研究发现意味着，过去几十年东地中海盆地的季节性火烧频率上升，可能转化为进化生态选择压力的变化，并对这一独特生态系统内的植物产生影响。     

Is Cupressus sempervirens native in Italy? An answer from genetic and palaeobotanical data

This study represents the first large-scale analysis using nuclear molecular markers to assess genetic diversity and structure of Cupressus sempervirens L.. Genetic and fossil data were combined to infer the possible role of human activity and evolutionary history in shaping the diversity of cypress populations. We analysed 30 populations with six polymorphic nuclear microsatellite markers. Dramatic reductions in heterozygosity and allelic richness were observed from east to west across the species range. Structure analysis assigned individuals to two main groups separating central Mediterranean and eastern populations. The two main groups could be further divided into five subgroups which showed the following geographical distributions: Turkey with the Greek islands Rhodes and Samos, Greece (Crete), Southern Italy, Northern Italy, Tunisia with Central Italy. This pattern of genetic structure is also supported by samova and Barrier analyses. Palaeobotanical data indicated that Cupressus was present in Italy in the Pliocene, Pleistocene and Holocene. Furthermore, our molecular survey showed that Italian cypress populations experienced bottlenecks that resulted in reduced genetic diversity and allelic richness and greater genetic differentiation. Recent colonization or introduction may also have influenced levels of diversity detected in the Italian populations, as most individuals found in this range today have multilocus genotypes that are also present in the eastern range of the species. The data reveal a new interpretation of the history of cypress distribution characterized by ancient eastern populations (Turkey and Greek islands) and a mosaic of recently introduced trees and remnants of ancient, depauperate populations in the central Mediterranean range.     

Pleistocene colonization of afro-alpine 'sky islands' by the arctic-alpine Arabis alpina

The afro-alpine region comprises the high mountains of Ethiopia and tropical East Africa, which represent biological 'sky islands' with high level of endemism. However, some primarily arctic-alpine plants also occur in the afro-alpine mountains. It has been suggested that these plants are Tertiary relicts, but a recent worldwide study of Arabis alpina suggests that this species colonized the region twice during the Pleistocene. Here we investigate the detailed colonization history of A. alpina in the afro-alpine region based on chloroplast DNA sequences from 11 mountain systems. The results confirm the twice-into-Africa scenario. The Asian lineage is confined to the mountains closest to the Arabian Peninsula, on opposite sides of the Rift Valley (Simen Mts and Gara Muleta in Ethiopia), suggesting long-distance dispersal of this lineage. The African lineage is divided into two phylogeographic groups with distinct geographic distribution. The observed pattern is consistent with isolation of the African lineage in at least two interglacial refugia, located on separated highlands, followed by range expansion in cooler period(s), when the afro-alpine habitat extended further down the mountains. Several long-distance dispersal events, also across the Rift Valley, are suggested by single haplotypes observed outside the area occupied by the phylogeographic groups they belonged to.     

Genetic consequences of Pleistocene range shifts: contrast between the Arctic, the Alps and the East African mountains

In wide-ranging species, the genetic consequences of range shifts in response to climate change during the Pleistocene can be predicted to differ among different parts of the distribution area. We used amplified fragment length polymorphism data to compare the genetic structure of Arabis alpina, a widespread arctic-alpine and afro-alpine plant, in three distinct parts of its range: the North Atlantic region, which was recolonized after the last ice age, the European Alps, where range shifts were probably primarily altitudinal, and the high mountains of East Africa, where the contemporary mountain top populations result from range contraction. Genetic structure was inferred using clustering analyses and estimates of genetic diversity within and between populations. There was virtually no diversity in the vast North Atlantic region, which was probably recolonized from a single refugial population, possibly located between the Alps and the northern ice sheets. In the European mountains, genetic diversity was high and distinct genetic groups had a patchy and sometimes disjunct distribution. In the African mountains, genetic diversity was high, clearly structured and partially in accordance with a previous chloroplast phylogeography. The fragmented structure in the European and African mountains indicated that A. alpina disperses little among established populations. Occasional long-distance dispersal events were, however, suggested in all regions. The lack of genetic diversity in the north may be explained by leading-edge colonization by this pioneer plant in glacier forelands, closely following the retracting glaciers. Overall, the genetic structure observed corresponded to the expectations based on the environmental history of the different regions.     

Individual and interactive effects of Amur honeysuckle (Lonicera maackii) and white-tailed deer (Odocoileus virginianus) on herbs in a deciduous forest in the eastern United States

Invasive Amur honeysuckle (Lonicera maackii) has reduced diversity, growth, and reproduction of native herbs in the Midwest USA. These effects may be compounded by browsing from overabundant white-tailed deer (Odocoileus virginianus). We used experimental treatments of honeysuckle (present, absent, removed) and deer (present, excluded) to measure their independent and interactive impacts on diversity, richness, and abundance of herbs in a deciduous forest in southwestern Ohio, USA. Species diversity and richness of herbs were not affected by honeysuckle or deer. Honeysuckle reduced abundance of annuals, graminoids, spring perennials, and summer perennials; deer decreased abundance of annuals and spring perennials, but increased abundance of graminoids. A deer × honeysuckle interaction showed that when honeysuckle was absent or removed, browsing by deer kept abundance of annuals and spring perennials low. Effects of honeysuckle and deer also were assessed for the three most abundant herbs: honeysuckle reduced abundance of Carex rosea and Sanicula odorata, and deer reduced abundance of Viola soraria. Herb abundance varied seasonally and annually. Honeysuckle and deer reduced number of leaves/stem of Maianthemum racemosum and a deer × honeysuckle interaction indicated that the negative effect of honeysuckle was released only when deer were excluded. Herb abundance and M. racemosum rebounded to or near to control levels after removal of honeysuckle. Our findings revealed that impacts of invasive honeysuckle or overabundant deer were not the same across all levels of biological organization (i.e., individual species, growth forms, community measures of species diversity/richness). Measuring impacts of these species at multiple levels of biological organization, considering deer × honeysuckle interactions, and collecting data for several years to account for seasonal and annual fluctuations will help guide management plans. The rapid response by herbs to removal of honeysuckle demonstrated the resilience of this community and is a hopeful sign for restoration of native understory herbs.     

Phylogeographic patterns of genetic diversity in eastern Mediterranean water frogs have been determined by geological processes and climate change in the Late Cenozoic

AIM: Our aims were to assess the phylogeographic patterns of genetic diversity in eastern Mediterranean water frogs and to estimate divergence times using different geological scenarios. We related divergence times to past geological events and discuss the relevance of our data for the systematics of eastern Mediterranean water frogs. LOCATION: The eastern Mediterranean region. METHODS: Genetic diversity and divergence were calculated using sequences of two protein-coding mitochondrial (mt) genes: ND2 (1038 bp, 119 sequences) and ND3 (340 bp, 612 sequences). Divergence times were estimated in a Bayesian framework under four geological scenarios representing alternative possible geological histories for the eastern Mediterranean. We then compared the different scenarios using Bayes factors and additional geological data. RESULTS: Extensive genetic diversity in mtDNA divides eastern Mediterranean water frogs into six main haplogroups (MHG). Three MHGs were identified on the Anatolian mainland; the most widespread MHG with the highest diversity is distributed from western Anatolia to the northern shore of the Caspian Sea, including the type locality of Pelophylax ridibundus. The other two Anatolian MHGs are restricted to south-eastern Turkey, occupying localities west and east of the Amanos mountain range. One of the remaining three MHGs is restricted to Cyprus; a second to the Levant; the third was found in the distribution area of European lake frogs (P. ridibundus group), including the Balkans. MAIN CONCLUSIONS: Based on geological evidence and estimates of genetic divergence we hypothesize that the water frogs of Cyprus have been isolated from the Anatolian mainland populations since the end of the Messinian salinity crisis (MSC), i.e. since c. 5.5-5.3 Ma, while our divergence time estimates indicate that the isolation of Crete from the mainland populations (Peloponnese, Anatolia) most likely pre-dates the MSC. The observed rates of divergence imply a time window of c. 1.6-1.1 million years for diversification of the largest Anatolian MHG; divergence between the two other Anatolian MHGs may have begun about 3.0 Ma, apparently as a result of uplift of the Amanos Mountains. Our mtDNA data suggest that the Anatolian water frogs and frogs from Cyprus represent several undescribed species.     

Plant functional group composition and large‐scale species richness in European agricultural landscapes

Question: Which are the plant functional groups responding most clearly to agricultural disturbances? Which are the relative roles of habitat availability, landscape configuration and agricultural land use intensity in affecting the functional composition and diversity of vascular plants in agricultural landscapes? Location: 25 agricultural landscape areas in seven European countries. Methods: We examined the plant species richness and abundance in 4 km × 4 km landscape study sites. The plant functional group classification was derived from the BIOLFLOR database. Factorial decomposition of functional groups was applied. Results: Natural habitat availability and low land use intensity supported the abundance and richness of perennials, sedges, pteridophytes and high nature quality indicator species. The abundance of clonal species, C and S strategists was also correlated with habitat area. An increasing density of field edges explained a decrease in richness of high nature quality species and an increase in richness of annual graminoids. Intensive agriculture enhanced the richness of annuals and low nature quality species. Conclusions: Habitat patch availability and habitat quality are the main drivers of functional group composition and plant species richness in European agricultural landscapes. Linear elements do not compensate for the loss of habitats, as they mostly support disturbance tolerant generalist species. In order to conserve vascular plant species diversity in agricultural landscapes, the protection and enlargement of existing patches of (semi‐) natural habitats appears to be more effective than relying on the rescue effect of linear elements. This should be done in combination with appropriate agricultural management techniques to limit the effect of agrochemicals to the fields.     

Variable flowering phenology and pollinator use in a community suggest future phenological mismatch

Recent anthropogenic climate change is strongly associated with average shifts toward earlier seasonal timing of activity (phenology) in temperate-zone species. Shifts in phenology have the potential to alter ecological interactions, to the detriment of one or more interacting species. Recent models predict that detrimental phenological mismatch may increasingly occur between plants and their pollinators. One way to test this prediction is to examine data from ecological communities that experience large annual weather fluctuations. Taking this approach, we analyzed interactions over a four-year period among 132 plant species and 665 pollinating insect species within a Mediterranean community. For each plant species we recorded onset and duration of flowering and number of pollinator species. Flowering onset varied among years, and a year of earlier flowering of a species tended to be a year of fewer species pollinating its flowers. This relationship was attributable principally to early-flowering species, suggesting that shifts toward earlier phenology driven by climate change may reduce pollination services due to phenological mismatch. Earlier flowering onset of a species also was associated with prolonged flowering duration, but it is not certain that this will counterbalance any negative effects of lower pollinator species richness on plant reproductive success. Among plants with different life histories, annuals were more severely affected by flowering–pollinator mismatches than perennials. Specialized plant species (those attracting a smaller number of pollinator species) did not experience disproportionate interannual fluctuations in phenology. Thus they do not appear to be faced with disproportionate fluctuations in pollinator species richness, contrary to the expectation that specialists are at greatest risk of losing mutualistic interactions because of climate change.     

Macroecological patterns of species and genetic diversity in vascular plants of the Mediterranean basin

Aim We address the question of whether broad scale biogeographical structure of species diversity (SD) matches that of genetic diversity (GD) of vascular plants. Location The Mediterranean basin. Methods We normalized vascular plant species richness (SD) estimates per country using the Med‐Checklist taxonomic database. We used a linear regression analysis to correlate normalized country estimates with country longitudinal position. We also compiled published and geo‐referenced within‐population GD data for tree species, which had populations in the Mediterranean. We normalized GD estimates for each population across species. Again, we used a linear regression analysis to correlate GD with population longitudinal position. We then compared the populations’ geographical and bioclimatic trends for GD with Last Glacial Maximum (LGM) palaeo‐climate data and the species current ecological requirements. Results The eastern Mediterranean and the coast of former Yugoslavia had higher SD than other regions. There was no overall spatial structure of SD in the Mediterranean, whereas there was an east–west trend of decreasing GD. This trend for GD tended to covary with an east–west warm/wet–cold/dry trend detected during the LGM. Low elevation xerothermic pine species displayed significantly less GD than higher elevation mesothermic or mountain pine species. Main conclusions We suggest that LGM climate may have significantly shaped the current longitudinal and altitudinal patterns of GD we observed in woody taxa across the Mediterranean, although it did not affect comparable SD patterns. In particular, colder LGM summer temperatures in the western Mediterranean may have reduced population sizes significantly more than in the eastern Mediterranean. As plant species richness and GD did not covary, SD and GD may not be used as surrogates of one another in the Mediterranean basin. As they contain comparatively less GD, conservation priorities in the Mediterranean should focus on hot spots of endemism and Western Mediterranean populations and species.     

Reevaluating sponge diversity and distribution in the Mediterranean Sea

The aim of this paper was (1) to update sponge diversity and distribution in the Mediterranean and (2) to re-examine faunal relationships among the Mediterranean areas on the basis of their sponge fauna. The Mediterranean demosponge faunal list was updated to 629 species by taking into consideration recent data from previously poorly studied areas. The species lists of 14 Mediterranean areas were compared on the basis of their sponge species richness, species composition, and taxonomic relatedness of species using multivariate analyses and diversity measures, such as PD, Delta+, and Lambda+. The 14 Mediterranean areas examined for their diversity and affinities were assembled into four major zoogeographic groups: the northwestern, northeastern, the central zone, and southeastern areas. Richest in species numbers were the areas belonging to the two northern groups. The species richness comparisons and similarity analyses performed at the generic level showed that it can be safely used as a surrogate for sponge species diversity in the Mediterranean. The results of this study showed that the simple traditional division of the Mediterranean Sea into a western, central, and eastern basin cannot reliably describe the distribution of sponges in the area. Thus, the W to E faunal decline previously presented for several faunal groups shifts to a general NNW-SSE pattern when one examines separately the northern and the southern parts of the traditional basins. This gradient seems to be in agreement with differences in key environmental variables, such as latitude, salinity, temperature, and water circulation, besides the typically examined distance from Gibraltar. Handling editor: T. P. Crowe     

Interactive effects of grazing and shrubs on the annual plant community in semi‐arid Mediterranean shrublands

Question: We studied the interactive effects of grazing and dwarf shrub cover on the structure of a highly diverse annual plant community. Location: Mediterranean, semi‐arid shrubland in the Northern Negev desert, Israel. Methods: Variation in the biomass and plant density of annual species in the shrub and open patches was monitored during four years, inside and outside exclosures protected from sheep grazing, in two contrasting topographic sites: north and south‐facing slopes that differed in their dominant dwarf shrubs species: Sarcopoterium spinosus and Corydothymus capitatus, respectively. Results: Above‐ground biomass, density and richness of annual species were lower under the canopy of both shrub species compared to the adjacent open patches in the absence of grazing. Grazing reduced the biomass of annuals in open patches of both topographic sites, but not in the shrub patches. On the north‐facing slope, grazing also reduced plant density and richness in the open patches, but increased plant density in the shrub patches. At the species level, various response patterns to the combined effects of grazing and patch type were exhibited by different annuals. Protection against the direct impacts of grazing by shrub cover as well as species‐specific interactions between shrubs and annuals were observed. A conceptual mechanistic model explaining these interactions is proposed. Conclusion: In semi‐arid Mediterranean shrublands grazing and dwarf shrub cover interact in shaping the structure of the annual plant community through (1) direct impacts of grazing restricted to the open patches, (2) species‐specific facilitation/ interference occurring in the shrub patches and (3) subsequent further processes occurring among the interconnected shrub and open patches mediated through variation in seed flows between patches.     

Multiple dispersal out of Anatolia: biogeography and evolution of oriental green lizards

The oriental green lizards of the Lacerta trilineata group are widely distributed in Greece, Anatolia, the eastern Mediterranean, the southern Caucasus, and the Zagros mountains in Iran. We studied their phylogeography using three mitochondrial markers with comprehensive sampling from most representatives of the group. Their phylogeny and divergence times (implementing fossil‐based molecular clock calibrations) were inferred using Bayesian methods, and haplotype networks were reconstructed to assess how genetic diversity and current distributional patterns were shaped. According to our phylogenetic analyses, the group constitutes a well‐supported monophylum containing several distinct evolutionary lineages with high haplotype diversity. Vicariance might explain the divergences within most lineages that have accumulated by range restriction and expansion of populations as a result of Quaternary climate oscillations and glacial refugia. However, niche divergence appears to be a major force promoting speciation, and large scale distributional patterns between lineages were shaped earlier by multiple, independent dispersals out of Anatolia during the Pliocene and early Pleistocene. The results of the present study also suggest that the group is in need of a taxonomical revision because the identified lineages and genetic diversity are not congruent with the currently recognized subspecies. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 398–408.     

Species–area relationships in Mediterranean-climate plant communities

Aim To determine the best‐fit model of species–area relationships for Mediterranean‐type plant communities and evaluate how community structure affects these species–area models. Location Data were collected from California shrublands and woodlands and compared with literature reports for other Mediterranean‐climate regions. Methods The number of species was recorded from 1, 100 and 1000 m² nested plots. Best fit to the power model or exponential model was determined by comparing adjusted r² values from the least squares regression, pattern of residuals, homoscedasticity across scales, and semi‐log slopes at 1–100 m² and 100–1000 m². Dominance–diversity curves were tested for fit to the lognormal model, MacArthur's broken stick model, and the geometric and harmonic series. Results Early successional Western Australia and California shrublands represented the extremes and provide an interesting contrast as the exponential model was the best fit for the former, and the power model for the latter, despite similar total species richness. We hypothesize that structural differences in these communities account for the different species–area curves and are tied to patterns of dominance, equitability and life form distribution. Dominance–diversity relationships for Western Australian heathlands exhibited a close fit to MacArthur's broken stick model, indicating more equitable distribution of species. In contrast, Californian shrublands, both postfire and mature stands, were best fit by the geometric model indicating strong dominance and many minor subordinate species. These regions differ in life form distribution, with annuals being a major component of diversity in early successional Californian shrublands although they are largely lacking in mature stands. Both young and old Australian heathlands are dominated by perennials, and annuals are largely absent. Inherent in all of these ecosystems is cyclical disequilibrium caused by periodic fires. The potential for community reassembly is greater in Californian shrublands where only a quarter of the flora resprout, whereas three quarters resprout in Australian heathlands. Other Californian vegetation types sampled include coniferous forests, oak savannas and desert scrub, and demonstrate that different community structures may lead to a similar species–area relationship. Dominance–diversity relationships for coniferous forests closely follow a geometric model whereas associated oak savannas show a close fit to the lognormal model. However, for both communities, species–area curves fit a power model. The primary driver appears to be the presence of annuals. Desert scrub communities illustrate dramatic changes in both species diversity and dominance–diversity relationships in high and low rainfall years, because of the disappearance of annuals in drought years. Main conclusions Species–area curves for immature shrublands in California and the majority of Mediterranean plant communities fit a power function model. Exceptions that fit the exponential model are not because of sampling error or scaling effects, rather structural differences in these communities provide plausible explanations. The exponential species–area model may arise in more than one way. In the highly diverse Australian heathlands it results from a rapid increase in species richness at small scales. In mature California shrublands it results from very depauperate richness at the community scale. In both instances the exponential model is tied to a preponderance of perennials and paucity of annuals. For communities fit by a power model, coefficients z and log c exhibit a number of significant correlations with other diversity parameters, suggesting that they have some predictive value in ecological communities.     

Microsatellite diversity of the agriculturally important alpine grass Poa alpina in relation to land use and natural environment

BACKGROUND AND AIMS: The Alpine Meadow Grass Poa alpina is common in subalpine and alpine natural sites and agriculturally used land, where it is an important fodder grass. Natural factors and human land use are supposed to have been shaping its genetic diversity for hundreds of years. The species comprises sexually and vegetatively reproducing plants. The aim of this study was to investigate the effects of agricultural land use, environmental factors and the mode of reproduction on the distribution of its microsatellite diversity within and among populations and to analyse whether its genetic diversity is correlated with plant species diversity in grassland parcels. METHODS: Genetic diversity of P. alpina was assessed with five microsatellite markers for 569 plants originating from 20 natural sites and from 54 grassland parcels of different cultural tradition, land use and altitude in the Swiss Alps. Due to polyploidy and frequent aneuploidy of the species, data analyses were based on the presence of microsatellite bands. KEY RESULTS: A low but significant differentiation was found in microsatellite bands among natural sites and agriculturally used parcels, while their microsatellite band diversity within populations did not differ. An increased differentiation was found in microsatellite bands with increasing geographic distance among parcels, and a differentiation among grazed and mown parcels, and among sexually and vegetatively reproducing populations. Band richness of sampled plants per village was higher for villages where parcels represented more different land-use types. Within populations, microsatellite band diversity was higher in grazed than in mown parcels. CONCLUSIONS: The diversity of human land use in the Alps was associated with genetic diversity of P. alpina. Therefore, the ongoing socio-economically motivated land-use changes, which reduce the number of different land-use types, will affect the genetic diversity of P. alpina negatively.     

Effects of repeated burning on Mediterranean communities of the northeastern Iberian Peninsula

Question: In relation to a single fire, do repeated wildfires in Mediterranean type ecosystems (1) reduce plant species richness or diversity; (2) modify patterns of abundance or dominance of plant species or (3) alter plant composition? Location: Pinus halepensis dominated communities of Catalonia, northeastern Iberian Peninsula, western Mediterranean Basin. Methods: Regional, paired design with 14 study sites, each consisting of a once burnt area (1994) and a twice burnt area (1975–1993 and 1994). Ten years after the last fire, we recorded all vascular plant species present in nested plots and quantified their relative abundances on transects. We compared species richness, diversity, dominance and relative abundance and species‐area correlations between paired once and twice burnt areas and assessed their floristic composition similarity. Results: No statistically significant differences were found in species richness or diversity. Slopes of species‐area correlations were higher in once burnt areas. In twice burnt areas, dominance by one or two species was higher. P. halepensis showed lower relative abundance and nanophanerophytes showed higher relative abundance. No differences were found for resprouter, seeder or resprouter‐seeder species. Floristic composition similarity between paired areas tended to be higher in less productive sites. Conclusions: Fire recurrence had contrasting effects on species richness at different spatial scales. Repeated burning reduced the relative abundance of the dominant tree species, which resulted in a higher relative abundance of shrubs. It also promoted the dominance of herbs, particularly Brachypodium retusum. However, it did not change the relative abundance of regenerative groups. Paired areas were more similar as they were more Mediterranean in terms of climatic conditions.     

Phytogeographical and community similarities of alpine tundras of Changbaishan Summit,China, and Indian Peaks,USA

Abstract. We compared the diversity, phytogeography, and plant communities in two mid-latitude alpine tundras with comparable aerial and elevational extents: Changbaishan Summit in eastern Asia and Indian Peaks in western North America. Despite wide separation, the two areas shared 72 species. In all, 43% of the species on Changbaishan Summit are also distributed in the alpine zones of western North America, while 22% of the species on Indian Peaks are also distributed in the alpine zones of eastern Asia. Almost all the shared species also occur in the Beringian region. Phytogeographical profiles of species and genera showed that 69% of species and over 90% of genera in both alpine tundras belong to the three phytogeographical categories: cosmopolitan, circumpolar, and Asian-North American. We attributed the current floristic relationship between these widely separated areas to the periodic past land connection between the two continents during the Tertiary and Pleistocene. Indian Peaks has a closer floristic relationship with the Arctic tundra than does Changbaishan Summit. Indian Peaks also has 45% higher species richness and lower vegetation cover than Changbaishan Summit. Plant communities from the two areas were completely separated in the two-way indicator species analysis and non-metric multidimensional scaling on floristic data at both species and generic levels, whereas ordination of communities by soil data produced a greater overlap. The plant communities on Changbaishan Summit in general have lower alpha diversity, higher beta diversity (lower between-community floristic similarity), and more rare species than does Indian Peaks. Mosaic diversity does not differ in the two alpine tundras, although the analysis suggests that Changbaishan Summit communities are more widely spaced on gradients than the Indian Peaks communities.