Bryophytes, lichens and phanerogams in an alvar grassland: relationships at different scales and contributions to plant community pattern

The relationship between bryophytes and lichens versus phanerogams has been investigated in three stands of a limestone grassland community, along three transects of 500 10 X 10 cm plots. Ordination axes resulting from Detrended Correspondence Analysis for bryophytes and lichens versus phanerogams were correlated using Spearman rank correlation coefficient. The effect of grain (sample plot) size on relationships between species along the gradient and on the correlation between the layers formed by bryophytes and lichens (cryptogams) and phanerogams, and with environmental variables has been as well examined. Values for light, moisture, pH and nitrogen have been derived from the vegetation itself with the help of Ellenberg indicator values. The relative position of species in the ordination space is more or less the same until grain size 3 (10 x 40 cm) for cryptogams and phanerogams and until grain size 2 (10 × 20 cm) for all-species together. Therefore, it is suggested that sample plot sizes of 10 × 10 cm to 10×20 cm are appropriate as units for field experiments testing interactions between cryptogams and phanerogams. The respective layers were weakly correlated and the correlation between them increased with increase in grain size. The correlation of DCA axis 1 from the ordination of cryptogams, phanerogams and all-species with the environmental factors was weak and similar in order of magnitude. Only the environmental variables which were strongly correlated with the DCA axis 1 increased in correlation at larger grain sizes. The ordinations of cryptogams, phanerogams and all-species were correlated along DCA axis 1 with pH in all investigated stands and at almost all grain sizes. Multi-species patches have been detected by pattern analysis (PASFRAN) on DCA sample scores from ordinations of cryptogams, phanerogams and combined matrices. Multi-species patterns with sizes between 20-240 cm. composed by bryophytes and lichens and by phanerogams have been found. Complex patterns formed by cryptogams and phanerogams together, which are different in size than those in the separate layers, suggest that bryophytes + lichens and phanerogams may interact with each other.     

On General Aspects of the Antartica Plants and Vegetation and Them Propects in Botanical Research

The flora of the Antarctic mainly consists of lichens , bryophytes and algae. The author analysed the distribution of plants in relation to their environments , floristic compositions, and types and structures of plant communities in both Antarctic and Subantarctic. A common property was recognized in both Antarctic and Subantarctic vegetation,i. e. ,mixed growth, epiphytic growth and replacement in succession were often found among the lichens, bryophytes and algae. Based on the characteristics of plant and vegetation and research condition, the botanical research of Chinese group may focus on the following subjects: (1)The origin of the Antarctica plants . The relationship between Plant and vegetation on the Antarctic and on other continents; (2)The mechanism of plant resistance to cold and to drought; (3) Antarctica terrestrial ecosystem research; (4) Biological diversity and its conservation; (5)Indication of the effects of Antarctic plants in environmental pollution ,climatic changes, chronology of glacier and mineral resources ; (6)Comparative study of alpine, Arctic and Antarctic plant.     

Effects of continuous and repeated dehydration on carbon fixation by bryophytes from the maritime Antarctic

 The effects of dehydration and rehydration on carbon exchange in 14 bryophytes from the maritime Antarctic were investigated using an infra-red gas analysis system. Continuous long-term (1–12 months) and repeated (1–6 one-month cycles) desiccation responses were investigated under controlled conditions. Loss of photosynthetic rate increased with length of dehydration period in all species, although some desiccation tolerance was observed even in those bryophytes from the most hydric habitats. Percentage retention of photosynthetic rate increased from hydric to xeric species, but this pattern was not repeated in terms of absolute rates of carbon fixation due to the high initial rates in the hydric species. Repeated cycles caused a greater loss of photosynthetic rate than continuous dehydration in hydric species, but the opposite situation occurred in mesic and xeric mosses. The latter groups were possibly better able to utilise the short periods of rehydration during cycles. In most bryophytes an increase in the percentage loss of photosynthetic rate following dehydration-rehydration occurred from spring to summer to autumn samples. This pattern was clearest in the hydric species and reduced in the xeric species. These variations were largely due to changes in the initial rates of photosynthesis during the growing season. It is suggested that this increased photosynthetic capacity is stress-sensitive, and is lost during either desiccation or winter freezing; the base photosynthetic capacity, being stress-tolerant, survives either of these events. The results obtained support the hypothesis that water availability is of importance in determining the distribution of bryophytes in the Antarctic. However, only the broad scale of variation in plant communities could be explained by these observations; other factors must be important in determining the finer scale of species distribution and community composition. The results are applicable to attempts to model the productivity of Antarctic bryophytes from known or predicted environmental data. Received: 15 April 1996 / Accepted: 28 September 1996     

In situ monitoring of microclimate and metabolic activity in lichens from Antarctic extremes: a comparison between South Shetland Islands and the McMurdo Dry Valleys

Lichens are the dominant organisms in terrestrial Antarctic ecosystems and show a decline in species number, coverage, and growth rate from the maritime Antarctic (62°S) to the McMurdo Dry Valleys (78°S). While Livingston Island (maritime Antarctica) is a hot spot for lichen biodiversity, the McMurdo Dry Valleys (continental Antarctica) are known as one of the most extreme environments for life. Previous studies suggest the biodiversity gradient to be linked to water availability acting through length of active period, but no activity data are available for the Dry Valleys. The work presented here compares metabolic activity of lichens at Livingston Island and the Dry Valleys for 4½ months from continuous monitoring that involves concurrent measurements of chlorophyll fluorescence and microclimate. The latitudinal comparison involves two contrasting habitats for plant physiological activity and microclimate. Two species of the foliose genus Umbilicaria were monitored in both regions plus one sample of the crustose Caloplaca in the Dry Valleys. The results showed a very large difference in the duration of activity over the monitoring period, and this supports the different coverage, species abundance, and growth rates already reported for lichens between both regions. Despite this large difference in activity, and in habitat conditions, analysis of the activity behaviour of the two Umbilicaria species shows interesting common features, while the crustose Caloplaca had additional strategies to improve hydration. This offers one explanation for the abundance of crustose lichens inside the Valleys, indicating better adaptation strategies to a polar desert.     

Summer variability, winter dormancy: lichen activity over 3?years at Botany Bay, 77��S latitude, continental Antarctica

Lichens make up a major component of Antarctic vegetation; they are also poikilohydric and are metabolically active only when hydrated. Logistic constraints have meant that we have little idea of the length, timing or environmental conditions of activity periods of lichens. We present the results of a three-year monitoring of the activity of the lichen Umbilicaria aprina at Botany Bay (77°S latitude) in the Ross Sea region, continental Antarctica. Chlorophyll fluorescence parameters that allowed hydrated metabolic activity to be detected were recorded with a special fluorometer at 2- or 3-h intervals. Air and thallus temperatures and incident PPFD (photosynthetic photon flux density, μmol photon m⁻² s⁻¹) were also recorded at hourly intervals. Activity was extremely variable between months and years and, overall, lichen was active for 7% of the 28-month period. Spring snow cover often delayed the onset of activity. Whereas the period immediately after snow melt was often very productive, the later months, January to March, often showed low or no activity. Mean thallus temperature when active was just above zero degrees and much higher than the annual mean air temperature of −15 to −19°C. Because major snow melts occurred when incident radiation was high, the lichen was also subjected to very high PPFD when active, often more than 2,500 μmol photon m⁻² s⁻¹. The major environmental stress appeared to be high light rather than low temperatures, and the variability of early season snow fall means that prediction of activity will be very difficult.     

Living on the edge – plants and global change in continental and maritime Antarctica

Antarctic terrestrial ecosystems experience some of the most extreme growth conditions on Earth and are characterized by extreme aridity and subzero temperatures. Antarctic vegetation is therefore at the physiological limits of survival and, as a consequence, even slight changes to growth conditions are likely to have a large impact, rendering Antarctic terrestrial communities sensitive to climate change. Climate change is predicted to affect the high‐latitude regions first and most severely. In recent decades, the Antarctic has undergone significant environmental change, including the largest increases in ultraviolet‐B (UV‐B; 290–320 nm) radiation levels in the world and, in the maritime region at least, significant temperature increases. This review describes the current evidence for environmental change in Antarctica, and the impacts of this change on the terrestrial vegetation. This is largely restricted to cryptogams, such as bryophytes, lichens and algae; only two vascular plant species occur in the Antarctic, both restricted to the maritime region. We review the range of ecological and physiological consequences of increasing UV‐B radiation levels, and of changes in temperature, water relations and nutrient availability. It is clear that climate change is already affecting the Antarctic terrestrial vegetation, and significant impacts are likely to continue in the future. We conclude that, in order to gain a better understanding of the complex dynamics of this important system, there is a need for more manipulative, long‐term field experiments designed to address the impacts of changes in multiple abiotic factors on the Antarctic flora.     

Leben zwischen Eis und Felsen

Biological soil crusts in Antarctica: Life between ice and rocks Despite its adverse environmental conditions and geographical isolation, Antarctica is home to a rich vegetation of lichens, mosses, algae, fungi and bacteria. In the milder areas of the maritime and continental Antarctic, these pioneer species form widely visible biological soil crusts. In drier areas, they occur mainly within the outer rock and upper soil layers. Among the ecological adaptations that enable these species to survive Antarctic conditions, a good dehydration tolerance stands out. Almost nothing is known about the genetic diversity of most species. While some species probably originated in Antarctica, others are relatively late settlers.     

Global patterns in local number of insect galling species

Abstract. We evaluate a three-part hypothesis explaining why gall-inducing insect species richness is so high in scleromorphic vegetation: (1) persistence of low nutrient status scleromorphic leaves facilitates the galling habit in warm temperate latitudes; (2) favourable colonization sites for gallers result from reduced hygrothermal stress, high phenolics in the outer cortex of the gall, and reduced carnivore and fungal attack in the gall; and (3) in more mesic sites, mortality is high due to carnivore attack and invasion of galls by fungi. Over 280 samples of local species of galling herbivorous insects from fourteen countries on all continents except Antarctica revealed a strong pattern of highest richness in warm temperate latitudes, or their altitudinal equivalents. The peak of galling species richness on the latitudinal gradient from the equator into the Arctic was between 25 to 38° N or S. Galling species were particularly diverse in sclerophyllous vegetation, which commonly had greater than twelve species per local sample. In mesic, non-sclerophyllous vegetation types the number of galling species was lower with twelve or fewer species present. Many sites in sclerophyllous vegetation supported between thirteen and forty-six galling species locally, including campina islands in Amazonia, cerrado savanna in central Brazil, the Sonoran Desert in Arizona and Mexico, shrubland in Israel, fynbos in South Africa and coastal scleromorphic vegetation in Australia. At the same latitude, or its elevational equivalent, galling species richness was significantly higher in relatively xeric sites when compared to riparian or otherwise mesic habitats, even when scleromorphic vegetation dominated the mesic sites. The results were consistent with the hypothesis and extend to a more general level the patterns and predictions on the biogeography of gall-inducing insects.     

Metabolic recovery of continental antarctic cryptogams after winter

The activation of metabolism after the winter period was investigated in several mosses and lichens in continental Antarctica. Thalli that were still in their over-wintering inactive state in early spring were sprayed artificially and the time-dependent activation of photosystem II (PSII), carbon fixation and respiration was determined using gas exchange and chlorophyll a fluorescence techniques. The investigated lichens recovered PSII activity almost completely within the first few minutes and gross photosynthesis was fully reactivated within a few hours. In contrast, photosynthesis took much longer to recover in mosses, which could indicate a general difference between the green-algal symbionts in lichens and moss chloroplasts. Only small and quickly reversible increased rates of respiration were observed for the foliose lichen Umbilicaria aprina from a more xeric habitat. In contrast, species occurring near persistent meltwater, such as the moss Bryum subrotundifolium and the lichen Physcia caesia, had highly increased respiration rates that were maintained for several days after activation. Calculation of the carbon balances indicated that the activation pattern strongly dictated the length of time before a carbon gain was achieved. It appears that the differences in recovery reflect the water relations of the main growth period in summer.     

Accidental transfer of non-native soil organisms into Antarctica on construction vehicles

Antarctic terrestrial ecosystems currently include very few non-native species, due to the continent’s extreme isolation from other landmasses. However, the indigenous biota is vulnerable to human-mediated introductions of non-native species. In December 2005, four construction vehicles were imported by contractors to the British Antarctic Survey’s (BAS) Rothera Research Station (Antarctic Peninsula) from the Falkland Islands and South Georgia (South Atlantic) on board RRS James Clark Ross. The vehicles were contaminated with >132 kg of non-Antarctic soil that contained viable non-native angiosperms, bryophytes, micro-invertebrates, nematodes, fungi, bacteria, and c. 40,000 seeds and numerous moss propagules. The incident was a significant contravention of BAS operating procedures, the UK Antarctic Act (1994) and the Protocol on Environmental Protection to the Antarctic Treaty (1998), which all prohibit the introduction of non-native species to Antarctica without an appropriate permit. The introduction of this diverse range of species poses a significant threat to local biodiversity should any of the species become established, particularly as the biota of sub-Antarctic South Georgia is likely to include many species with appropriate pre-adaptations facilitating the colonisation of more extreme Antarctic environments. Once the incident was discovered, the imported soil was removed immediately from Antarctica and destroyed. Vehicle cleaning and transportation guidelines have been revised to enhance the biosecurity of BAS operations, and to minimise the risk of similar incidents occurring.     

Photosynthetic symbionts in Antarctic terrestrial ecosystems: the physiological response of lichen photobionts to drought and cold

Lichens form an important part of the biodiversity in terrestrial ecosystems of Antarctica where they represent the dominant vegetation. Previous studies on the genetic diversity of photobionts of lichens have indicated that clade S Trebouxia photobionts are the most widespread in continental Antarctica, predominantly in macrolichens. For the first time, a comparative study of the physiology of a variety of isolated Antarctic lichen photobionts (genus Trebouxia) was performed. Photosynthetic activity was examined by chlorophyll a fluorescence and correlated with freezing and desiccation under laboratory conditions and photosynthetic pigments were quantified in response to desiccation. Data were obtained from photobionts collected from the Antarctic regions of North Victoria Land, Coal Nunatak and Rothera Point, as well as from a European site (Gotland, Sweden). While the isolated algae reacted individually to stress treatments, they were highly susceptible to desiccation stress but could rapidly recover from freezing. Photobiont-specific physiological adaptations are considered to explain the dominance of clade S Trebouxia photobionts.     

UAV-based classification of maritime Antarctic vegetation types using GEOBIA and random forest

Development of vegetation communities in areas of Antarctica without permanent ice cover emphasizes the need for effective remote sensing techniques for proper monitoring of local environmental changes. Detection and mapping of vegetation by image classification remains limited in the Antarctic environment due to the complexity of its surface cover, and the spatial heterogeneity and spectral homogeneity of cryptogamic vegetation. As ultra-high resolution aerial images allow a comprehensive analysis of vegetation, this study aims to identify different types of vegetation cover (i.e., algae, mosses, and lichens) in an ice-free area of  Hope Bay, on the northern tip of the Antarctic Peninsula. Using the geographic object-based image analysis (GEOBIA) approach, remote sensing data sets are tested in the random forest classifier in order to distinguish vegetation classes within vegetated areas. Because species of algae, mosses, and lichens may have similar spectral characteristics, subclasses are established. The results show that when only the mean values of green, red, and NIR bands are considered, the subclasses have low separability. Variations in accuracy and visual changes are identified according to the set of features used in the classification. Accuracy improves when multilayer information is used. A combination of spectral and morphometric products and by-products provides the best result for the detection and delineation of different types of vegetation, with an overall accuracy of 0.966 and a Kappa coefficient of 0.946. The method allowed for the identification of units primarily composed of algae, mosses, and lichens as well as differences in communities. This study demonstrates that ultra-high spatial resolution data can provide the necessary properties for the classification of vegetation in Maritime Antarctica, even in images obtained by sensors with low spectral resolution.     

Alien lichens unintentionally transported to the “Arctowski” station (South Shetlands, Antarctica)

The recent climate changes combined with intensified human activity in Antarctica are promoting the synanthropization process and increasing the likelihood of alien species establishing in the native communities. Cargo items, expedition members’ equipment and food destined for the 32nd Polish Antarctic Expedition to the “Arctowski” station in the 2007/2008 season were inspected to determine their potential as vectors for alien lichen species. Within the cargo, packaging and foodstuffs scanned, a total of 45 lichen specimens (24 species) were identified. Most of them had been accidentally transported with various timbers. Cargo containers and fresh food were also found to harbour for single specimens. The majority of lichens detected are alien to the Antarctic biota and had never been observed in the region. This paper estimates the potential risk of these lichens establishing themselves in remote southern latitudes. The results emphasize the threat of accidental introduction of alien organisms into Antarctica and the need for taking every precaution to prevent the importation of non-native species to this unique environment.     

Measured and modelled leaf and stand‐scale productivity across a soil moisture gradient and a severe drought

Environmental controls on carbon dynamics operate at a range of interacting scales from the leaf to landscape. The key questions of this study addressed the influence of water and nitrogen (N) availability on Pinus palustris (Mill.) physiology and primary productivity across leaf and canopy scales, linking the soil‐plant‐atmosphere (SPA) model to leaf and stand‐scale flux and leaf trait/canopy data. We present previously unreported ecophysiological parameters (e.g. V_cmax and J_max) for P. palustris and the first modelled estimates of its annual gross primary productivity (GPP) across xeric and mesic sites and under extreme drought. Annual mesic site P. palustris GPP was ～23% greater than at the xeric site. However, at the leaf level, xeric trees had higher net photosynthetic rates, and water and light use efficiency. At the canopy scale, GPP was limited by light interception (canopy level), but co‐limited by nitrogen and water at the leaf level. Contrary to expectations, the impacts of an intense growing season drought were greater at the mesic site. Modelling indicated a 10% greater decrease in mesic GPP compared with the xeric site. Xeric P. palustris trees exhibited drought‐tolerant behaviour that contrasted with mesic trees' drought‐avoidance behaviour.     

Photosynthesis and dark respiration in Antarctic mosses—an initial comparative study

Rates of dark respiration (DR), gross photosynthesis (GPS) and net photosynthesis (NPS) were investigated for 14 species of moss from a maritime Antarctic locality. The rates found were similar to those reported in studies of temperate, alpine and Arctic species, indicating no physiological specialisation to the Antarctic environment. There was no relationship between the habitat occupied by a species (hydric, mesic, xeric) and physiological measures. There was, however, a loose correlation between NPS and a species' ecology or reproductive behaviour in the maritime Antarctic — species with high NPS being either colonists or those that show high and regular investment in sporophyte production.     

Species richness of vascular plants, bryophytes and lichens in dry grasslands: The effects of environment, landscape structure and competition

We studied the relative importance of local habitat conditions and landscape structure for species richness of vascular plants, bryophytes and lichens in dry grasslands on the Baltic island of Öland (Sweden). In addition, we tested whether relationships between species richness and vegetation cover indicate that competition within and between the studied taxonomic groups is important. We recorded species numbers of vascular plants, bryophytes and lichens in 4 m² plots (n=452), distributed over dry grassland patches differing in size and degree of isolation. Structural and environmental data were collected for each plot. We tested effects of local environmental conditions, landscape structure and vegetation cover on species richness using generalized linear mixed models. Different environmental variables explained species richness of vascular plants, bryophytes and lichens. Environmental effects, particularly soil pH, were more important than landscape structure. Interaction effects of soil pH with other environmental variables were significant in vascular plants. Plot heterogeneity enhanced species richness. Size and degree of isolation of dry grassland patches significantly affected bryophyte and lichen species richness, but not that of vascular plants. We observed negative relationships between bryophyte and lichen species richness and the cover of vascular plants. To conclude, effects of single environmental variables on species richness depend both on the taxonomic group and on the combination of environmental factors on a whole. Dispersal limitation in bryophytes and lichens confined to dry grasslands may be more common than is often assumed. Our study further suggests that competition between vascular plants and cryptogams is rather asymmetric.     

Drought response of five conifer species under contrasting water availability suggests high vulnerability of Norway spruce and European larch

The ability of tree species to cope with anticipated decrease in water availability is still poorly understood. We evaluated the potential of Norway spruce, Scots pine, European larch, black pine, and Douglas‐fir to withstand drought in a drier future climate by analyzing their past growth and physiological responses at a xeric and a mesic site in Central Europe using dendroecological methods. Earlywood, latewood, and total ring width, as well as the δ¹³C and δ¹⁸O in early‐ and latewood were measured and statistically related to a multiscalar soil water deficit index from 1961 to 2009. At the xeric site, δ¹³C values of all species were strongly linked to water deficits that lasted longer than 11 months, indicating a long‐term cumulative effect on the carbon pool. Trees at the xeric site were particularly sensitive to soil water recharge in the preceding autumn and early spring. The native species European larch and Norway spruce, growing close to their dry distribution limit at the xeric site, were found to be the most vulnerable species to soil water deficits. At the mesic site, summer water availability was critical for all species, whereas water availability prior to the growing season was less important. Trees at the mesic were more vulnerable to water deficits of shorter duration than the xeric site. We conclude that if summers become drier, trees growing on mesic sites will undergo significant growth reductions, whereas at their dry distribution limit in the Alps, tree growth of the highly sensitive spruce and larch may collapse, likely inducing dieback and compromising the provision of ecosystem services. However, the magnitude of these changes will be mediated strongly by soil water recharge in winter and thus water availability at the beginning of the growing season.     

Phenotypic plasticity and population differentiation in seeds and seedlings of the grass Anthoxanthum odoratum

Summary Seeds of Anthoxanthum odoratum were transplanted reciprocally between a xeric and a mesic field population that were genetically differentiated in adult traits. In one experiment seeds were reciprocally buried in bags in the soil, in a second experiment seeds were reciprocally sown in small plots. For most traits, site effects were much larger than seed-source effects. Germination, emergence, mortality of buried seed and recruitment were significantly higher at the xeric site than at the mesic site, irrespective of population of origin. Seed dormancy, was significantly higher for seed originating from the mesic than from the xeric population. Seedling recruits originating from the xeric population tended to be larger at both sites. Fecundity of seedling recruits depended on the environment; fecundities of plants growing in the xeric site had more than double the fecundity of plants growing in the mesic site. Phenotypic plasticity rather than population differences determined variation in performance in the seed and seedling stages.     

Functional and spatial pressures on terrestrial vegetation in Antarctica forced by global warming

There is growing interest in what controls the present distribution of terrestrial vegetation in Antarctica because of the potential use of biodiversity as an indicator or predictor of the effects of climate change. Recent advances in knowledge of distribution and ecophysiological performance of terrestrial vegetation means that an initial analysis of the potential influence of temperature is now possible. Regressions of species numbers of lichens, mosses and hepatics on latitude and mean annual temperature (standard macroclimatic data) were carried out, and the terrestrial vegetation in Antarctica could be divided into two zones. The microenvironmental zone lies south of around 72°S, and biodiversity (richness and location) is uncoupled from the macroenvironment and is, instead, determined by the occasional coincidences of warmth, water, light and shelter. The macroenvironmental zone lies north of about 72°S, and biodiversity (richness, cover and growth) is strongly positively linked to mean annual temperature; species numbers increase at about 9–10% per K (24.0, 9.3 and 1.8 species for lichens, mosses and hepatics, respectively) probably due to improved water availability through increased precipitation and longer active period (monthly degree-days also reach zero at about 72°S) allowing greater productivity, completion of metabolic processes and a switch from survival to growth strategies. Cyanobacterial lichens appear to be a special case and may be expanding after being forced into northerly refugia. Warming will cause a southward movement of the boundary between the two zones but distribution in the microenvironmental zone will remain determined by local coincidences of environment and resources.