Spatial patterns of surface soil properties and vegetation in a Mediterranean semi-arid steppe

In arid and semi-arid areas with sparse vegetation cover, the spatial pattern of surface soil properties affects water and nutrient flows, and is a question of considerable interest for understanding degradation processes and establishing adequate management measures. In this study, we investigate the spatial distribution of vegetation and surface soil properties (biological crusts, physical crusts, mosses, rock fragments, earthworm casts, fine root accumulation and below-ground stones) in a semi-arid Stipa tenacissima L. steppe in SE Spain. We applied the combination of spatial analysis by distance indices (SADIE) and geostatistics to assess the spatial pattern of soil properties and vegetation, and correlation analyses to explore how these patterns were related. SADIE analysis detected significant clumped patterns in the spatial distribution of vegetation, mosses, fine root accumulation and below-ground stone content. Contoured SADIE index of clustering maps suggested the presence of patchiness in the distribution of earthworm casts, fine roots, below-ground stone content, mosses and biological crusts. Correlation analyses suggested that spatial pattern of some soil properties such as biological crusts, moss cover, surface rock fragments, physical crusts and fine roots were significantly related with above-ground plant distribution. We discuss the spatial arrangement of surface soil properties and suggest mechanistic explanations for the observed spatial patterns and relationships.     

Effects of isolation,logging and dispersal on woody-species richness of islands

Isolation effects on species richness of woody plants were investigated in a system of islands that were created by the filling of the Clarks Hill Reservoir, Georgia. This reservoir was built between 1946–1954. Some islands were logged and cleared of woody plants prior to the filling of the reservoir and others were not logged. The presence of logged versus unlogged islands in the same system allowed us to test whether and how geographical isolation interacts with island history and species-specific dispersal properties in determining patterns of among-island variation in species number. Thirty-six years after the islands were created, logged islands had significantly fewer species of woody plants than unlogged ones. On logged islands, total number of woody species was negatively correlated with distance to the closest mainland (r=–0.95). On unlogged islands, variation in species number was very low (CV=4.9%) and was not correlated with distance to the mainland. These results indicate that the studied system as a whole has not yet reached equilibrium. However, the mean number of species on unlogged islands was very close to the intercept of the regression obtained for logged islands, suggesting that islands close to the mainland have already reached their equilibrium species richness. This conclusion is consistent with predictions of island biogeography theory. When species representing different dispersal properties were analyzed separately, statistically significant distance effects were obtained for bird-dispersed species (r=0.88) and for species with no adaptations to bird or wind dispersal (r=0.81). Wind-dispersed species did not show a decrease in species number with increasing isolation, but their relative frequency was positively and significantly correlated with distance to the mainland (r=0.94). Historical factors, as well as differences among species in dispersal properties, are important in explaining patterns of among-island variation in species number.     

Spatial associations and patterns of perennial vegetation ina semi-arid steppe: a multivariate geostatistics approach

We investigated the spatial patterns of perennial species (Stipa tenacissima, Anthyllis cytisoidesGlobularia alypum, Brachypodium retusum and chamaephytes) in a 50 m × 50 m semi-arid steppe by using the combination of a linear model of coregionalization (LMC) and sampling units of varying size (1.25 m × 1.25 m, 2.5 m × 2.5 m, and 5 m × 5 m). The data-adjusted LMC showed the patchy structure of the vegetation, which was especially evident with the highest resolution grid. It also detected a periodic pattern in the distribution of S. tenacissima, as well as autocorrelation at two spatial scales for A. cytisoides and G. alypum. The latter species was negatively associated with the other species at both short and long distances. These negative associations were consistent for all sampling grids and suggest the presence of interference between G. alypum and the rest of the evaluated species. Despite species-specific differences, the LMC was fitted satisfactorily to all of them. This suggests a common variation pattern for all the species, which may be caused by an underlying environmental property driving the patterns of all the species or, alternatively, by the dominance of some species’ spatial pattern, or another kind of species association, over the rest. The spatial patterns found were profoundly affected by the observational scale. Our results reveal that the multivariate geostatistical approach introduced in this paper is a suitable technique for the spatial analysis of semi-arid plant communities. It allows plant ecologists to evaluate if the species forming the plant community of interest share a common spatial pattern, and to assess the spatial covariation between the species forming a plant community at different spatial scales independently.     

Patterns of composition and species richness of crustaceans and aquatic insects along environmental gradients in Mediterranean water bodies

Differences in the dynamics of ecological processes between Mediterranean and colder temperate aquatic systems could imply different patterns in faunal communities in terms of composition and biodiversity (i.e. species richness and rarity). In order to identify some of these patterns the crustacean and aquatic insect composition and biodiversity of four water body types, classified according to their salinity and water permanence, were compared. Moreover, the relationships between species richness and water, pond and landscape variables were analysed. A total number of 91 water bodies located throughout Catalunya (NE Iberian Peninsula) were sampled. Three species assemblages were observed: one for permanent freshwaters, another for temporary freshwaters, and a third one for saline waters (SW), since permanent and temporary saline water bodies had similar composition. Differences in salinity were associated with proportion of crustaceans versus insects and with singularity. Thus, saline ponds had a higher proportion of crustaceans, and lower values of singularity. Conductivity was significantly related to total (crustaceans plus insects) richness, and also related to insect richness. The main difference between the models obtained for crustacean species richness and insect species richness is the significance of landscape variables in the latter, and this fact could be related to the different dispersion types of these two faunal groups: active for insects versus passive for crustaceans. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users. Guest editors: R. Céréghino, J. Biggs, B. Oertli & S. Declerck The ecology of European ponds: defining the characteristics of a neglected freshwater habitat     

Relative influence of size,connectivity and disturbance history on plant species richness and assemblages in fragmented grasslands

Questions: What is the relative influence of size, connectivity and disturbance history on plant species richness and assemblages of fragmented grasslands? What is the contribution of small fragments to the conservation of native species pool of the region? Location: Tandilia's Range, Southern Pampa, Argentina. Methods: Cover of plants was registered within 24 fragments of tall‐tussock grassland remnants within an agricultural landscape using modified Whittaker nested sampling. We analysed the influence of site variables related to disturbance history (canopy height, litter thickness) and fragment variables (size, connectivity) on species richness (asymptotic species richness, slope of the species–area curve) as well as on species assemblages by multiple regressions analysis and canonical correspondence analyses, respectively. Cumulative area was used for analysing whether small fragments or large fragments are more important to species diversity in the landscape. Results: Asymptotic species richness was significantly influenced by site variables, in particular by Paspalum quadrifarium's canopy height, but not by fragment variables. Species assemblages were also affected by site variables (12.2% of total variation), but no additional portion of the species assemblage variability was significantly explained by fragment size and connectivity. Sampling of several small fragments rendered more exotic and native species than sampling of few large fragments of the same total area. Conclusions: Our results agree with previous studies reporting low sensitivity of species diversity to size and isolation of grassland fragments in fragmented landscapes and high sensitivity of species diversity to local variables. The higher capture of regional native species pool by small grassland fragments than by few larger ones of equivalent accumulated area highlights the value of small fragments for conservation.     

Seasonal changes in net photosynthesis rates and photosynthetic capacity in leaves of Cistus salvifolius,a European mediterranean semi-deciduous shrub

Summary During five different periods between Nov. 1982 and Aug. 1983, the diurnal patterns exhibited in photosynthetic CO₂ uptake and stomatal conductance were observed under natural conditions on twigs of Cistus salvifolius, a Mediterranean semi-deciduous shrub which retains a significant proportion of its leaves through the summer drought. During the same periods, net photosynthesis at saturating CO₂ partial pressure was measured on the same twigs as a function of irradiance at different temperatures. From these data, photosynthetic capacity, defined here as the CO₂- and light-saturated net photosynthesis rate, was obtained as a function of leaf temperature. C. salvifolius is a winter growing species, shoot growth being initiated in Nov. and continuing through May. Photosynthetic capacity was quite high in Nov., March and June, exceeding 40 mol m^-2 s^-1 at optimum temperature. In Dec., photosynthetic capacity was somewhat reduced, perhaps due to low night-time temperatures (<5°C) during the measurement period. In Aug., capacity in oversummering shoots at optimum temperature fell to less than 8 mol m^-2 s^-1, due to water trees and perhaps leaf aging. Seasonal changes in maximal photosynthetic rates under ambient conditions were similar, and like those found in co-occurring evergreen sclerophylls. Like the evergreens, Cistus demonstrated considerable stomatal control of transpirational water loss, particularly in oversummering leaves. During each measurement period except Aug. when capacity was quite low, the maximum rates of net photosynthesis measured under ambient conditions were less than half the measured photosynthetic capacities at comparable temperatures, suggesting an apparent excess nitrogen investment in the photosynthetic apparatus.     

Combined effects of enhanced UV-B radiation and additional nutrients on growth of two Mediterranean plant species

Seedlings of two Mediterranean plants, the slow-growing, evergreen sclerophyll Ceratonia siliqua L. and the fast growing drought semi-deciduous Phlomis fruticosa L., were grown for one year in the field at ambient or ambient plus supplemental UV-B radiation (equivalent to a 15% ozone depletion) and two levels of applied fertilizers (NPK). The effects on growth, morphological, anatomical and physiological parameters were measured at final plant harvest. Additional nutrients increased leaf nitrogen, improved growth and reduced the root/shoot ratio in both plants, yet these effects were more pronounced in the fast growing P. fruticosa. A nutrient-induced increase in chlorophyll content was also observed in this plant. The growth responses to UV-B radiation were different for the two species. Growth in C. siliqua was not affected by UV-B radiation at both nutrient levels and the same was true for P. fruticosa at low nutrients. However, at the high nutrient level, supplemental UV-B radiation improved growth in P. fruticosa, indicating a strong interaction between the treatments. Photosystem II (PSII) photochemical efficiency, methanol-extractable UV-B absorbing capacity, total phenolics and tannins were not affected by either treatment in both plants. It is concluded that nutrient levels can strongly modify the UV-B radiation effects on growth of P. fruticosa. We presume that this may be correlated to the fast growing habit of this species.     

Long-term impacts of goat browsing on bush-clump dynamics in a semi-arid subtropical savanna

The effects of 16 years of continuous browsing by goats in a South African savanna at stocking rates intended for bush control were compared with plots unbrowsed for the same period of time. Differences in bush-clump density, structure and species composition were recorded. Bush-clump density did not differ between browsed and unbrowsed plots. Within individual bush-clumps, browsing was shown to impact more on structure than species composition, with smaller, shorter bush-clumps, containing fewer species but much greater stem-densities. Although species presence/absence was little affected by browsing, many species showed differences in abundance, growth and location within browsed and unbrowsed bush-clumps. Species reduced in abundance in browsed plots included Cussonia spicata, Ehretia rigida, Grewia occidentalis, Jasminum angulare and Senecio linifolius. Several species showed reduced growth in browsed plots, particularly those located at bush-clump edges. The relatively unpreferred Aloe ferox was a notable exception. Although browsing had little effect on the composition of the main clump founding species, emergents or late arrivals, there were twice as many single plants in browsed plots and emergence of several species was restricted to the middle of bush-clumps. Comparison of our findings with aerial photographic evidence and other literature suggest that browsing alone is unlikely to significantly reduce scrub cover, although it can clearly control further expansion. Combinations of fire and browsing, rather than one factor alone, are considered likely to act fastest and most effectively to significantly reduce or remove scrub cover altogether.     

Effect of substrate treatments on survival and growth of Mediterranean shrubs in a revegetated quarry: An eight-year study

Loss of soil is a major problem that restoration actions must solve in Mediterranean quarries. Soil replacement by a low-quality substrate is often inadequate for revegetation and limits plant establishment and growth in these degraded areas, which are also subject to drought recurrent stress.In 1998, a field experiment was performed in a limestone quarry at Outão (Serra da Arrábida, SW Portugal) to test different types of substrate improvement: NPK fertilizer, water-holding polymer, mycorrhizal inoculum, and combinations of these. Two-year old plants of three native woody species were planted - carob (Ceratonia siliqua), wild-olive tree (Olea europaea var. sylvestris) and mastic (Pistacia lentiscus). Reports concerning the short-term results showed some differential effects of the assayed treatments on plant growth and physiology. The monitoring program was maintained, and here we report on the survival and growth of the introduced plants over eight years after planting. This prolonged study showed that, with the limited exception of fertilization, none of the assayed treatments added major advantages for plant survival or growth. Regardless of the tested substrate treatment, mortality was low and these native species became established in the revegetated area.     

Importance of historical factors on species richness and composition of butterfly assemblages (Lepidoptera: Rhopalocera) in a northern Iberian mountain range

ABSTRACT. The butterfly assemblages of three vegetation types (pasture, heathland and rocky outcrop with Genista spp.) and two geographical orientations (a northern slope and a sheltered gorge) were studied along an altitudinal gradient in the Picos de Europa in northern Spain. In order to study the effects or historical climatic changes on current butterfly assemblages, it was examined if changes in richness and faunal composition of assemblages were joined with changes in their biogeographical composition. The three vegetation types showed characteristic butterfly assemblages, but no difference in species richness and biogeographical composition was found. Species richness in the sheltered gorge was higher than in the northern slope. Faunal composition also varied between orientations and there were more widespread and Mediterranean species in the sheltered gorge than in the northern slope. Species richness declined with increasing altitude, though a midaltitudinal peak was observed. There was a faunal segregation between high and low localities. Species richness of widespread and Mediterranean butterflies decreased as altitude increased, whereas the reverse was true for montane species. Moreover, biogeographical elements differed in their climatic tolerances measured as altitudinal ranges. Therefore, changes in richness and composition of butterfly assemblages between both slopes and along the altitudinal gradient were joined in part with variation in their biogeographical composition. These results suggest that current species richness and composition of butterfly assemblages in the Picos de Europa might be the consequence of differential colonization of refuges during the past climatic changes.     

Determinants of woody species richness in Scot pine and beech forests: climate, forest patch size and forest structure

We analysed patterns of woody species richness in Pinus sylvestris and Fagus sylvatica forests in Catalonia (NE Spain) from forestry inventory databank in relation to climate and landscape structure. Both types of forests are found within the same climatic range, although they have been managed following somewhat different goals. Overall, woody species richness significantly increased when conditions get closer to the Mediterranean ones, with milder temperatures. Differences between the two types of forests arose when comparing the relationship between richness and forest patch size. Woody species richness increased in pine forests with patch size, while the opposite trend was observed in beech forests. This pattern is explained by the different behaviour of structural canopy properties, since leaf area index and canopy cover showed a steeper increase with increasing forest patch size in Fagus forests than in Pinus ones. Accordingly, richness decreased with canopy cover in Fagus plots, but not in Pinus ones. We suggest that these differences would be related to management history, which may have enhanced the preservation of beech stands in larger forest landscape units.     

Hierarchical large-scale to local-scale influence of abiotic factors in summer-fragmented Mediterranean rivers: structuring effects on fish distributions,assemblage composition and species richness

Summer rainfall shortage and low or no flows in Mediterranean rivers make it difficult for fish to attain their final preferenda and hinder the assembling of fish communities. We investigated in this study the degree to which large-scale features of the watersheds and reach-scale habitats correlated with a non-random structure of fish species' richness, composition and distribution under these conditions by intensively sampling a river system during summer and using Geographical Information Systems data. The results of multivariate ordination and association analyses showed that multi-scale factors were hierarchically related and that they correlated with non-random fish distributions and assemblage structure. They also highlighted large-scale factors shaping reach-scale water availability for fish during summer. Fish species' richness, composition and distribution varied along a gradient of water shortage and connectivity loss which were more severe with the smaller drainage area into the reaches upstream. Fish species' distributions along these gradients were strongly correlated with both life-history traits (adult size) and biogeographical origin (native vs. exotic). The results for the summer environmental segregation of exotic versus small endemic richness characteristics indicate strong species' interactions forcing the latter into the upstream, harsher seasonal habitats. Special fitness costs are to be expected in drought years, in regulated rivers, and in general, in a climate change scenario in Mediterranean river networks. The study emphasizes that the conservation of the threatened Mediterranean freshwater fish faunas requires operating at the proper spatial scale. In particular, reach-scale habitat improvements are adequate but not enough during the summer in fragmenting river networks, unless there is proper watershed-scale management of the flow regimes.     

Edge and shape effects on ant (Hymenoptera: Formicidae) species richness and composition in forest fragments

In this paper, we tested four hypotheses relative to edge and shape effects on ant communities: (i) forest edges have lower species richness than the remnant core; (ii) species richness increases with distance from the edge; (iii) irregularly shaped remnants have lower species richness than more regular remnants; (iv) there is a higher similarity of species composition between edge and core in irregular than in regular remnants. We sampled litter ant communities on the edge and core of ten remnants, in Viçosa, Minas Gerais, Brazil. Species richness was larger at the forest core than at the edges, although did not increase with distance from the edge. Species richness did not vary with shape complexity. The similarity of species composition between edge and core showed a decreasing trend with remnant area, and did not vary with shape complexity. The observed differences of species richness between forest core and edge may be due to higher harshness of edges, caused by environmental changes. The absence of relationship between species richness and distance from the edge might indicate the range of edge effects, which would be smaller than the smallest distance of core sampled. Therefore, edges would affect litter-dwelling ant species richness in a distance smaller than 50 m. The observation of species composition allowed us to notice an effect of fragmentation that would not be noticed if we were considering only species richness. Edge may serve as step to generalist species, which may use it to colonise forest remnants. Furthermore, small remnants are more colonisation-prone by such species, and have a more homogeneous species composition than large remnants.     

Irrigation and fertilization effects on seed number,size, germination and seedling growth: implications for desert shrub establishment

Plants with limited resources adjust partitioning among growth, survival, and reproduction. We tested the effects of water and nutrient amendments on seed production, size, and quality in Sarcobatus vermiculatus (greasewood) to assess the magnitude and importance of changes in reproductive partitioning. In addition, we assessed interactions among the environment of seed-producing plants (adult plant scale), seed size, and seedling microenvironment (seedling scale) on successful seedling establishment. Interactions of these factors determine the scale of resource heterogeneity that affects seedling establishment in deserts. Both total number of seeds produced per plant and seed quality (weight and germination) increased significantly in the enriched treatment in a 3-year field experiment. Seedling length 3 days after germination and seed N concentration, other measures of seed quality, were higher for seed from both irrigated and enriched plants than for seed from control plants. Field S. vermiculatus seed production and quality can be substantially increased with irrigation and nutrient enrichment at the adult plant scale and this allows management of seed availability for restoration. However, based on a greenhouse study, seedling environment, not the environment of the seed-producing plant or seed size, was the most important factor affecting seedling germination, survival, and growth. Thus it appears that production of more seed may be more important than improved seed quality, because higher quality seed did not compensate for a low-resource seedling environment. For both natural establishment and restoration this suggests that heterogeneity at the scale of seedling microsites, perhaps combined with fertilization of adult shrubs (or multi-plant patches), would produce the greatest benefit for establishing seedlings in the field.     

Floristic composition and floral phenology of the Mediterranean batha of Ariel,Samaria

Field observations over a three-year period were used to document the floristic composition, flowering and fruiting sequences of the Sarcopoterium spinosa Mediterranean batha of Ariel. This plant community was found to be species-rich, with high percentages of annuals, bushes and geophytes, low percentages of herbaceous plants, climbers and trees, and ten species considered rare in Israel. Typical Mediterranean characteristics were observed in the S. spinosa batha: spring flowering reaching a peak in April, small between-year variation in flowering and fruiting, most species having Mediterranean distribution, and partial resistance to fire.     

Patterns in species richness,species density,and evenness in groundfish assemblages on the continental slope of the U.S. Pacific coast

For many taxa, diversity, often measured as species richness, decreases with latitude. In this report patterns of diversity (species richness, species diversity, and evenness) in groundfish assemblages were investigated in relation to depth (200–1200 m) and latitude (33–47°N) on the continental slope of the U.S. Pacific coast. The data originated from the 1999–2002 upper continental slope groundfish surveys conducted by the National Marine Fisheries Service. When the data were pooled across depths, species density and evenness were found to decline with latitude. All three diversity measures declined with depth, with the lowest overall diversity in the 600- to 900-m depth range where longspine thornyhead Sebastolobus altivelis constituted close to 70% of the catch. When latitudinal gradients were examined within four depth zones (200–300 m, 400–500 m, 600–900 m, and 1000–1200 m) more complex patterns emerged. At depth species richness and evenness were inversely correlated with latitude as longspine thornyhead dominated catches to the north. However, in shallower areas of the slope, species richness and evenness were positively correlated with latitude. Latitudinal patterns of diversity in the deeper zones and when pooled across depths were positively correlated with temperature and broadly consistent with the Ambient Energy hypothesis discussed by Willig et al. [Annu Rev Ecol System 34:273–309 (2003)].     

Demographic consequences of drought in the herbaceous perennial Cryptantha flava: effects of density,associations with shrubs,and plant size

The demographic consequences of a severe drought year were examined for two experimental plantings of the herbaceous desert perennial Cryptantha flava(Boraginaceae) in northeastern Utah, United States. A total of 6680 nutlets were planted individually or in clusters of four both under shrubs and in open microhabitats within two natural populations. Survival, growth, and flowering as a function of density and microhabitat were followed for 7 years, including 1 year when precipitation just before and during the growing season was 74.5% below normal. The design permitted assessment of how intraspecific density and shrub cover affect demographic response to drought. Mortality increased and flowering decreased dramatically during drought but neither varied with density or between shrub and open microhabitats. For plants growing under shrubs, survival (at Site 1) and growth (at Site 2) varied with shrub species. Average aboveground plant size also decreased during drought. Population size hierarchies were rearranged because larger plants lost leaf rosettes while many smaller plants grew. Density and microhabitat affected plant performance in non-drought years but more often at Site 1 than at Site 2. Individuals growing alone often were more likely to flower and/or produced more inflorescences when they did flower than did individuals growing with at least one other plant. However, for 2 years, survival rates at Site 1 were higher for plants growing in clumps than for single individuals. Shrubs also had mixed effects on plant performance. In some years, survival was higher under shrubs, but at Site 1 plants in the open often were more likely to flower and/or produced more inflorescences. Thus despite severe demographic consequences of drought, the study provided no evidence that intraspecific competition, interference by shrubs, or facilitation by shrubs increases under limited soil water.     

Veronica: Parallel morphological evolution and phylogeography in the Mediterranean

The genus Veronica s. lat. comprises about 450 species (including about 180 species from the southern hemisphere Hebe-complex), many of which grow in the Mediterranean area. Their extreme variability in morphology, life form and habitats has led to many suggestions regarding evolution and biogeography. Difficulties arise from parallel syndromes, widespread among alpine species and lowland perennials, and particularly among annual species of the genus. We have used sequences of the plastid trnL-F region and nuclear ribosomal ITS sequences to differentiate between different clades of Veronica and reveal cases of parallel evolution. Based on this data, cases of parallel evolution have been found in biogeographical patterns among the alpine species of Veroniceae, in which species from European mountains have affinities to those in the Central Asian/Himalayan region whereas alpine species from Turkey are probably more recently derived from lowland southwestern Asian taxa. Different subspecies of Veronica bombycina gained their characteristic morphology independently and parallel in adaptation to their alpine environment. Pinnatifid leaves have been gained parallel in perennial grassland species of Veronica. Finally, parallel evolutionary trends in many characters, not only morphological but also molecular characters, are common among annual species of Veronica.We wish to thank the Studienstiftung des deutschen Volkes for a doctoral scholarship to DCA and the Spanish Ministerio de Ciencia y Tecnologia for a postdoctoral grant (programa FPI) to MMO. This work was partially supported by the Junta de Castilla y Leon and the European Union (FSE) through the research project SA117/01. We also thank Prof. F. Ehrendorfer for critical comments and valuable discussions. Further, DCA thanks Daniela Hanfland and Yoshiki Nakamoto for help in the field. Manfred A. Fischer, Lena Struwe, Bernhard Dickore, Christoph Dobes, Eberhard Fischer, Niels Köster, Gerald Schneeweiss, Tod Stuessy, Tim Utteridge and Matsugo Yokota have provided plant material for this study.     

An experimental study of factors affecting the distribution of yellow perch and central mudminnows along a species richness gradient

Synopsis Effects of environmental factors on the distribution and abundance of yellow perch and central mudminnows in northern Wisconsin were examined by holding populations of these fishes within single-species enclosures in a series of three small forest lakes having a species-richness gradient of one to four species. These enclosures allowed each species to experience environmental conditions within each lake without directly interacting with each other. In the four-species lake, two other sets of enclosures addressed the effects of intra- and interspecific competition. Changes in total biomass of enclosure stocks (from growth and mortality) indicated that in the absence of other species, perch did best in the lake containing the richest fish assemblage, intermediate in the two-species lake, and worst in the mudminnow-only lake. Mudminnow stocks similarly performed significantly better in the four-species lake than the mudminnow-only lake. These results suggest that the lakes' environmental conditions contribute to the patterns of presence and abundance of perch, but that interspecific interactions override a similar contribution for the mudminnow, which is regarded as a fugitive species. Perch performances were also sensitive to fish densities within enclosures, declining significantly when stocks were doubled, either by adding more perch or equal numbers of mudminnows.     

Ecogeography of the grass genusHelictotrichon (Poaceae: Aveneae) in the Mediterranean and adjacent regions

The geographical distribution of the grass genusHelictotrichon in the Mediterranean Region is analysed and mapped for 33 taxa. Based on only a single life form (perennial herb), a variety of edaphically, climatically and altitudinally differently adapted species complexes has evolved in the area. Most of these complexes show west-east disjunctions and contain geographically, sometimes even edaphically vicarious taxa with complementary distribution. A transition from mesomorphic to xeromorphic habit occurred independently in different species groups and led to the establishment of the modern Mediterranean taxa which are in part highly polyploid derivatives of more mesophilic diploids. The significance of polyploidy, patterns of parapatric and sympatric distribution, biogeographical borders, and centres of species diversity are discussed in context with the history of the Mediterranean vegetation. New combinations are:Helictotrichon setaceum subsp.petzense, H. pratense subsp.lusitanicum, H. praetutianum.