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.     

Patterns of plant diversity in Africa south of the Sahara and their implications for conservation management

Plant species richness and range-size rarity in Africa south of the Sahara is concentrated in centres of plant diversity and endemism. Distribution patterns of plants mapped in the Distributiones Plantarum Africanum series and selected taxonomic monographs are analysed using the computer programme WORLDMAP. The plants are divided into four groups: herbaceous geophytes, mesophytic herbs, light-demanding shrubs and woody genera. Each group has peaks of species richness and range-size rarity at locations different to the other groups. Herbaceous geophytes and mesophytic herbs have their peaks of species richness and range-size rarity in the same location, the western Cape for geophytes and the Crystal Mountain for mesophytic herbs, whereas light-demanding shrubs and woody genera have peaks in different places. The results are discussed in relation to possible factors determining species richness and endemism and their likely conservation significance.     

The relative importance of local, regional and historical factors determining the distribution of plants in fragmented riverine forests: an emergent group approach

Aim To investigate how local, regional and historical factors shape the herbaceous plant communities in fragmented riverine forests, and how the community composition and species richness of these fragments is related to the interplay between the environmental factors and specific plant life‐trait combinations. Location Riverine forest fragments in the Grand‐duché de Luxembourg. Methods Forest fragments were surveyed for their abundance in herbaceous plant species. All plant species where clustered into Emergent Groups (EG) by means of a formal classification based on 14 life‐history traits. Within each EG, the local, regional and historical factors were related to the community composition using partial Canonical Correspondence Analyses (pCCA) and to the species richness using Generalized Linear Models (GLMs). The EG colonization ability was characterized by means of logistic regressions. Results We defined and characterized seven EGs, among which three consisted of forest specialist species (barochorous perennials, short geophytes and zoochorous perennials), which exhibited specific life‐trait combinations: large and short‐lived seeds and/or vernal phenology. Differences in EG composition between forest fragments were mainly explained by local environmental factors such as soil productivity and pH. The richness of barochorous perennials and short geophytes was well predicted by the historical and regional factors. The colonization ability appeared very low for barochorous perennials and short geophytes. Main conclusions Local environmental conditions appear to drive the differentiation of the riverine forest plant communities owing to the specific habitat requirements of many forest species. Spatial and temporal forest discontinuities affect the richness of forest specialist species, due to dispersal and/or recruitment limitations. The emergent group approach enhances the understanding of the relative influence of local, regional and historical factors by distinguishing between forest specialists from generalists or ‘matrix’ species, which have a masking effect.     

Composition of grazed and cleared temperate grassy woodlands in eastern Australia: patterns in space and inferences in time

Abstract. A regional vegetation survey of the temperate grassy woodlands (temperate savanna) in Australia was designed to assess the effects of clearing and grazing on the composition of vegetation remnants and the adjacent pasture matrix. Vegetation was sampled across a range of habitats using 77 0.1024‐ha quadrats; the relative abundance of species was recorded. Classification analysis clustered the sites into three main groups that corresponded to intensity of grazing/clearing followed by groups based on underlying lithology (basalt, metasediment, granites). Using Canonical Correspondence Analysis, exogenous disturbance and environmental variables were related to the relative abundance of species; grazing intensity had the highest eigenvalue (0.27) followed by tree canopy cover (0.25), lithology (0.18), altitude (0.17) and slope (0.10). Based on two‐dimensional ordination scores, six species response groups were defined relating to intensity of pastoralism and nutrient status of the landscape. Abundance and dominance of native shrubs, sub‐shrubs, twiners and geophytes were strongly associated with areas of less‐intense pastoralism on low‐nutrient soils. The strongest effects on species richness were grazing followed by canopy cover. Continuously grazed sites had lower native species richness across all growth forms except native grasses. There was no indication that intermediate grazing intensities enhanced forb richness as a result of competitive release. Species richness for all native plants was lowest where trees were absent especially under grazed conditions. Canopy cover in ungrazed sites appeared to promote the co‐existence of shrubs with the herbaceous layer. Predicted declines in forb richness in treeless, ungrazed, sites were not detected. The lack of a disturbance‐mediated enhancement of the herbaceous layer was attributed to habitat heterogeneity at 0.1 ha sampling scale.     

Flora, vegetation, and soil in broadleaved ancient and planted woodland, and scrub on RøsnÆs, Denmark

9 small ancient woodlands (>200 years), 9 planted woodland sites (25 — ca. 100 years), and 6 sites in grazed scrub (ca. 100 years) on the RøsnÆs peninsula, Denmark, showed characteristic differences: in ancient woodland, the tree and shrub layer was fairly rich, the field layer rather poor in species. The dominance of spring flowering geophytes, the abundance of Anemone nemorosa , and the occurrence of Corylus avellana and Polygonatum multiflorum were characteristic. pH of the soil was relatively low, organic matter content high, and light intensity at the forest floor in summer low. In planted woodland there was more light, and the field layer was rich in short-lived species, but poor in spring flowering geophytes. Many woodland species were rare in planted woodland, some did not at all occur there, and none were specific for this type of woodland. The scrub was marked by grazing and a strong relief, hence pH was high and organic matter content low. The field layer was rich both in shortlived species and in spring flowering geophytes. — It is suggested that ancient woodland species (i.e. species restricted to or preferably occurring in woodlands existing prior to the enclosure ca. 200 years ago) is a heterogenous group, consisting of a) species favoured by traditional woodland management; b) species restricted to woodlands where specific environmental (e.g. soil) conditions have had sufficient time to develop; c) species with limited ability to spread or establish; or d) species which in ancient woodland are represented by small and scattered populations.     

Genome size and DNA base composition of geophytes: the mirror of phenology and ecology?

Background and Aims

Genome size is known to affect various plant traits such as stomatal size, seed mass, and flower or shoot phenology. However, these associations are not well understood for species with very large genomes, which are laregly represented by geophytic plants. No detailed associations are known between DNA base composition and genome size or species ecology.

Methods

Genome sizes and GC contents were measured in 219 geophytes together with tentative morpho-anatomical and ecological traits.

Key Results

Increased genome size was associated with earliness of flowering and tendency to grow in humid conditions, and there was a positive correlation between an increase in stomatal size in species with extremely large genomes. Seed mass of geophytes was closely related to their ecology, but not to genomic parameters. Genomic DNA GC content showed a unimodal relationship with genome size but no relationship with species ecology.

Conclusions

Evolution of genome size in geophytes is closely related to their ecology and phenology and is also associated with remarkable changes in DNA base composition. Although geophytism together with producing larger cells appears to be an advantageous strategy for fast development of an organism in seasonal habitats, the drought sensitivity of large stomata may restrict the occurrence of geophytes with very large genomes to regions not subject to water stress.     

古田山自然保护区地下芽植物多样性及其生物量分配策略

地下芽植物能够通过地下储存器官占据生境资源、储存营养物质等策略来获得生态优势,其地下储存器官多样性以及生物量分配策略,对地下芽植物物种组成以及生态系统功能产生重要影响。然而,以往研究多关注草地生态系统的地下芽植物,对森林地下芽植物的了解仍然缺乏。采集了古田山国家级自然保护区不同海拔分布的693个草本植物个体,分析了地下芽植物及其地下储存器官的类型与多样性,比较了地下芽植物与非地下芽植物的地上、地下各器官的绝对、相对生物量。结果显示：（1）地下芽植物的相对丰富度为69.1%,相对多度为88.2%。大多为根状茎植物,主要由禾本科、莎草科、堇菜科和蕨类植物组成。（2）除茎外,地下芽植物各器官的绝对生物量（叶：1.94g,根：0.65g,地上部分：2.0g,地下部分：4.1g）均大于非地下芽植物（叶：0.26g,根：0.13g,地上部分：0.68g,地下部分：0.13g）。（3）地下芽植物叶（0.40）与茎（0.14）的相对生物量小于非地下芽植物（叶：0.48,茎：0.35）,地下部分相对生物量（0.56）大于非地下芽植物（0.17）。本研究表明,以根状茎植物为主的地下芽植物是古田山亚热带森林生态系统草本植物的主要构成者,且个体普遍较大,倾向于将生物量投资于地下器官。这些结果为认识地下芽植物的生态策略与功能以及草本植物群落管理提供了科学依据。     

Biogeography of prostrate-leaved geophytes in semi-arid South Africa: hypotheses on functionality

Nowhere is the species diversity of geophytes greater than in the five mediterranean-climate ecosystems of the world. Of these, the Cape mediterranean zone of South Africa is the most speciose. While the relative diversity and importance of geophytes of all of the other four mediterranean regions of the world drops off sharply as one moves into adjacent winter-rainfall desert regions, geophytes in the semi-arid to arid Succulent Karoo (including Namaqualand) remain a very important component of the flora, both in terms of abundance and diversity (comprising 13 to 29% of the regional floras in this region). Apart from species richness, there are also a number of interesting geophyte growth forms in this region. One unusual growth form is geophytes with flattened leaves that lie prostrate on the soil surface. At least eight families (Amaryllidaceae, Colchicaceae, Eriospermaceae, Geraniaceae, Hyacinthaceae, Iridaceae, Orchidaceae and Oxalidaceae) exhibit this growth form. While this growth form is relatively common in many geophyte lineages in the Succulent Karoo biome and the Cape mediterranean zone (Fynbos biome), and occurs infrequently through the summer-rainfall temperate regions of Africa, it is virtually absent in other regions worldwide. A null hypothesis is that the prostrate leaved trait is a neutral characteristic, however biogeographical data do not support this. A neutral trait would be unlikely to show such a clear pattern of distribution. Several alternative hypotheses on the adaptive significance of this growth form are discussed. These include: avoidance of herbivory, reduction in competition from neighbors, creation of a CO₂ enriched environment below the leaves, reduction of water loss around the roots, reduction of water loss through transpiration, precipitation of dew on the leaves and maintenance of optimal leaf temperatures for growth.     

Grazing in remnant woodland vegetation: changes in species composition and life form groups

Abstract. Grazing by domestic livestock in native woodlands can have major effects on ecosystem functioning by the removal of plant species that form important functional groups. This paper documents the changes in floristics in a large group of remnants of native woodland left after agricultural clearing in southwestern Australia. Species richness and diversity were significantly reduced in remnants and the proportion of exotic species increased. Detrended Correspondence Analysis (DCA) was used to identify floristic and environmental patterns among plots and identified two distinct groups based on grazing intensity. This indicated that the significance of the relationship between grazing effects and DCA floristic axes was greater than edaphic characteristics that normally influence floristic patterns. Floristic characteristics of sites that were influencing the position of plots on the ordination diagram included proportion of exotic species and proportion of native perennial shrubs and herbs. Numbers of species of native shrubs and perennial herbs were significantly reduced in grazed plots and numbers of exotic annual grasses and herbs were significantly higher. Other life form groups such as native perennial grasses and geophytes were not significantly affected by grazing. Reproductive strategies of perennial species showed a significant decrease in numbers of resprouters and a significant increase in numbers of facultative seeder/sprouters. Exclosure plots showed increases in number and cover of perennial shrubs and herbs after three years whereas number and cover of exotic species did not change. Time series DCA showed that the floristic composition of exclosure plots in grazed sites became closer to that of the ungrazed sites.     

Nutrient reserves may allow for genome size increase: evidence from comparison of geophytes and their sister non-geophytic relatives

Background and Aims

The genome size of an organism is determined by its capacity to tolerate genome expansion, given the species'' life strategy and the limits of a particular environment, and the ability for retrotransposon suppression and/or removal. In some giant-genomed bulb geophytes, this tolerance is explained by their ability to pre-divide cells in the dormant stages or by the selective advantage of larger cells in the rapid growth of their fleshy body. In this study, a test shows that the tendency for genome size expansion is a more universal feature of geophytes, and is a subject in need of more general consideration.

Methods

Differences in monoploid genome sizes were compared using standardized phylogenetically independent contrasts in 47 sister pairs of geophytic and non-geophytic taxa sampled across all the angiosperms. The genome sizes of 96 species were adopted from the literature and 53 species were newly measured using flow cytometry with propidium iodide staining.

Key Results

The geophytes showed increased genome sizes compared with their non-geophytic relatives, regardless of the storage organ type and regardless of whether or not vernal geophytes, polyploids or annuals were included in the analyses.

Conclusions

The universal tendency of geophytes to possess a higher genome size suggests the presence of a universal mechanism allowing for genome expansion. It is assumed that this is primarily due to the nutrient and energetic independence of geophytes perhaps allowing continuous synthesis of DNA, which is known to proceed in the extreme cases of vernal geophytes even in dormant stages. This independence may also be assumed as a reason for allowing large genomes in some parasitic plants, as well as the nutrient limitation of small genomes of carnivorous plants.     

Relationships between environmental variables and vegetation across mountain wetland sites,N. Iran

The mountain wetlands studied represent a unique habitat on the southern slopes of the Alborz mountain range, the second largest range in Iran. In comparison with other parts of this range the western section is ecologically and botanically unknown. Floristic and vegetation variation were assessed using diverse environmental variables along a broad altitudinal span (350 m to 3200 m a.s.l.). Using both statistical and ordination analyses floristic variation was assessed on three defined altitudinal belts which were delimited based on Alborz macro-climatic boundaries. The distribution of individual wetland plant species, of phytogeographic elements and of life-forms all differ among altitudinal belts. This result is also shown in both direct and indirect analyses of ordinations. The proportion of geophytes significantly increases with altitude and geophytes are very well represented in the upper altitudinal belt. The number of species of a narrow phytogeographical distribution (e.g. endemics) increases with altitude, soil pH and EC declined with altitude. The first axis of DCA ordination with passively projected environmental variables indicates that, organic matter and concentration of Fe²⁺ are increased toward higher altitude. The second axis of ordination is related to both soil texture and slope inclination. The distribution of species in the CCA species plot is also close to the distribution of those in the DCA ordination. This study indicates that altitude and slope together with other dependent environmental variables (pH, EC, Ca²⁺ and soil texture) are the main ecological factors controlling species distribution across the Western Alborz wetland sites.     

Decoupling of reproduction and growth: an unusual pattern in the life cycle of the Mediterranean geophyte Narcissus serotinus

The unpredictable seasonality of the Mediterranean climate imposes on the development of specific life‐cycle phenologies such as those seen in geophyte species that have a period of dormancy to avoid stressful abiotic conditions. To date, two life‐cycle strategies are recognized in Mediterranean geophytes. In some species, growth and reproduction occur in different seasons (hysteranthous geophytes), whereas in others these two processes occur at the same time (synanthous geophytes). A study of the life cycle of the autumnal‐flowering Narcissus serotinus in three populations over two consecutive years confirmed that neither the hysteranthy nor the synanthy strategy could be applied to this geophyte. Rather, reproduction and growth occur simultaneously, but in different plants. A strong correlation was found between bulb weight and the presence of reproductive or vegetative plants.     

Using niche‐based GIS modeling to test geographic predictions of competitive exclusion and competitive release in South American pocket mice

Geographic studies addressing the role of competition in determining species’ macrodistributions have been limited by only simple or subjective means of identifying regions of suitable habitat. Now, ecological‐niche models of species’ potential distributions present a possible approach to testing for the geographic patterns predicted under competitive exclusion and competitive release. Previously, we modeled the potential distributions of two spiny pocket mice (Heteromys australis and H. anomalus) in northwestern South America using specimen localities, environmental data, and the Genetic Algorithm for Rule‐Set Prediction (GARP). Here we superimpose the models to examine known distributional records in areas of potential sympatry between the two species, thus testing the geographic predictions of competitive exclusion. In addition, we examine environmental characteristics of known localities, testing for data consistent with competitive release. Areas of potential sympatry are minimal, lying in regions of intermediate water balance. Only records of H. australis are known from areas of potential sympatry in regions where the species’ ranges meet, consistent with exclusion of H. anomalus by H. australis. Heteromys anomalus inhabits areas ecologically suitable for both species only in the isolated Sierra Nevada de Santa Marta, in which H. australis is not present (most likely for historical reasons). Furthermore, environmental characteristics of localities of H. anomalus in biogeographic regions where H. australis is absent fit the pattern predicted under competitive release. In contrast, localities of H. australis show no indication of competitive release. Although the results of present analyses do not conclusively demonstrate competitive exclusion or release, they provide directional hypotheses that can now be tested in experimental field and laboratory studies.     

Foraging in the subterranean social highveld mole-rat (Cryptomys hottentotus pretoriae): an investigation into mass-dependent geophyte use and foraging patterns

The foraging behaviour of captive colonies of the highveld mole-rat Cryptomys hottentotus pretoriae was investigated in an artificial soil-filled burrow system provided with four trays (patches) that varied in geophyte density and mass. An initial trial involving empty trays (only soil) revealed that there was no preference for any specific tray. There were no statistically significant preferences for excavating in any of the patches of different geophyte density. No preferences were evident for excavation in patches containing geophytes of different mass classes. Empty patches seemed to be preferred over patches containing geophytes when combinations of geophyte density/mass were investigated. The duration of handling and the rate of consumption of geophytes were recorded for 23 individuals of two mass classes. Handling time of geophytes was not related to mole-rat sex, but was strongly linked to mole-rat mass class. Handling time of geophytes was related to geophyte mass class. Small geophytes were less profitable to consume. These findings are considered in light of optimal foraging theory and the situation in the field. It was concluded that the mole-rats generally followed the qualitative predictions of optimal foraging theory, although falling short of being energy maximizers.     

Water as a resource,stress and disturbance shaping tundra vegetation

Water is crucial for plant productivity and survival as a fundamental resource, but water conditions can also cause physiological stress and mechanical disturbance to vegetation. However, these different influences of water on vegetation patterns have not been evaluated simultaneously. Here, we demonstrate the importance of three water aspects (spatial and temporal variation of soil moisture and fluvial disturbance) for three ecologically and evolutionary distinct taxonomical groups (vascular plants, mosses and lichens) in Fennoscandian mountain tundra. Fine‐scale plant occurrence data for 271 species were collected from 378 × 1 m² plots sampled over broad environmental gradients (water, temperature, radiation, soil pH, cryogenic processes and the dominant allelopathic plant species). While controlling all other key environmental variables, water in its different aspects proved to be a crucial environmental driver, acting on individual species and on community characteristics. The inclusion of the water variables significantly improved our models. In this high‐latitude system, the importance of spatial variability of water exceeds the importance of temperature for the fine‐scale distribution of species from the three taxonomical groups. We found differing responses to the three water variables between and within the taxonomical groups. Water as a resource was the most important water‐related variable in species distribution models across all taxonomical groups. Both water resource and disturbance were strongly related to vascular plant species richness, whereas for moss species richness, water resources had the highest influence. For lichen species richness, water disturbance was the most influential water‐related variable. These findings demonstrate that water variables are not only independent properties of tundra hydrology, but also that water is truly a multifaceted driver of vegetation patterns at high‐latitudes.     

Theophrastus on geophytes

NEGBI, M., 1989. Theophrastus on geophytes . The nature of the geophilous organs of the geophytes described by Theophrastus (371-287 B.C.) in Historia Planiarum and De Causis Planlarum is presented and discussed. Most of the geophytes and the hydrogeophytes mentioned in these books (excluding species of Allium , biennial vegetables, and a few others) are critically reviewed. Certain lost agricultural practices mentioned by Theophrastus are strengthened and made explicable by comparisons from Pliny's Natural History , and Hebrew literature of the first centuries A.D.: Mishnah and Talmud. Theophrastus's interest in plants is shown to be of a pure nature although most of his examples are cultivated or otherwise useful plants. The possible sources of his knowledge of Egyptian plants are discussed.     

Changes in the dominant assembly mechanism drive species loss caused by declining resources

The species–energy hypothesis predicts that more productive areas support higher species richness. Conversely, when resources are reduced, species richness is reduced. Empirical tests of whether extinctions are predominantly caused by environmental constraints or competitive exclusion are lacking. We experimentally reduced dead wood to c. 15% of the initial amount after a major windstorm and examined changes in assembly mechanisms by combining trait‐based and evolutionary species dissimilarities of eight taxonomic groups, differing in their dependence on dead wood (saproxylic/non‐saproxylic). Species richness and assembly mechanisms of non‐saproxylic taxa remained largely unaffected by removal of dead wood. By contrast, extinctions of saproxylic species were caused by reversing the predominant assembly mechanisms (e.g. increasing importance of competitive exclusion for communities assembled through environmental filtering or vice versa). We found no evidence for an intensification of the predominant assembly mechanism (e.g. competitive exclusion becoming stronger in a competitively structured community).     

The seasonal response of diatom communities to variable water quality in some English urban lakes

SUMMARY. 1. Five diatom assemblages and their corresponding site groups were associated with the urban lakes. When treated as a single dataset, DC A and PCA grouped these diatom assemblages along an environmental gradient from circum natural. dilute waters to strongly alkaline, concentrated, nutrient-rich waters.
2. When treated as individual datasets, 75% of the known variance was explained by the first axis of DCA in all five diatom assemblages; this indicated the existence of a strong primary environmental gradient. The PCA results showed that the identity of this primary environmental gradient can be different for each assemblage.
3. The statistical analyses showed that the physicochemical gradients were continuous and not discrete. Nevertheless, fine-scale analysis can identify smaller, more precise species and site groups.
4. Two types of species response could be identified within most sub-assemblages in the urban lakes: the first was a quasi-seasonal response and was characterized by a gradual rise to a peak relative abundance of one dominant or several co-dominant species which could be sustained for a period of weeks before a gradual decline. A quasi-seasonal response was sometimes interrupted by a storm response. The storm response was characterized by a sudden increase in the relative abundance of one or more diatom species following a storm, after which the species declined to pre-storm abundances within 7 days.
5. The effects of urban runoff on the water quality of urban lakes can override the effects of true seasonal factors such as water temperature and day length on the relative abundance of species.     

Ecological evidence concerning the adaptive significance of the C4 dicarboxylic acid pathway of photosynthesis

Summary Ecological data were analyzed to determine the environmental conditions associated with the occurrence of plants possessing C₃ and C₄ photosynthetic pathways. Non-parametric analysis of variance, multiple regression analysis and discriminant analysis techniques were applied to information derived from separate studies on plant species in California and central Europe. All the analyses revealed significant differences in ecology between the C₃ and C₄ groups. The occurrence of C₄ species was greater where summer or winter temperatures were relatively high and moisture availability relatively low. Using discriminant analysis, more than 80% of the species considered were classified into the correct photosynthetic category on the basis of their distribution with respect to temperature. No significant differences between the C₃ and C₄ groups were found for other environmental factors, including light, soil nitrogen, soil salinity and continentality of climate. These results support the suggestion from previous physiological studies that C₄ species may possess competitive advantages under conditions of high temperature and intermittent water stress. The C₄ species were relatively restricted in the range of environmental conditions where they occurred, and as a group C₄ species may be ecologically more specialized than C₃ species.