Showing papers in "Plant Ecology in 2000"

How do depth, duration and frequency of flooding influence the establishment of wetland plant communities?

Abstract: In many temporary wetlands such as those on the Northern Tablelands of New South Wales Australia, the development of plant communities is largely the result of germination and establishment from a long-lived, dormant seed bank, and vegetative propagules that survive drought. In these wetlands the pattern of plant zonation can differ from year to year and season to season, and depth is not always a good indicator of the plant community composition in different zones. In order to determine which aspects of water regime (depth, duration or frequency of flooding) were important in the development of plant communities an experiment using seed bank material from two wetlands was undertaken over a 16 week period in late spring–early summer 1995–1996. Seed bank samples were exposed to 17 different water-level treatments with different depths, durations and frequencies of flooding. Species richness and biomass of the communities that established from the seed bank were assessed at the end of the experiment and the data were examined to determine which aspects of water regime were important in the development of the different communities. It was found that depth, duration and frequency of inundation influenced plant community composition, but depth was least important, and also that the duration of individual flooding events was important in segregating the plant communities. Species were grouped according to their ability to tolerate or respond to fluctuations in flooding and drying. The highest biomass and species richness developed in pots that were never flooded. Least biomass and species richness developed in pots that were continuously flooded. Short frequent floods promoted high species richness and biomass especially of Amphibious fluctuation-tolerator species and Amphibious fluctuation-responder species that have heterophylly. Terrestrial species were able to establish during dry phases between short floods. Depth was important in determining whether Amphibious fluctuation-tolerator or Amphibious fluctuation-responder species had greater biomass. Longer durations of flooding lowered species richness and the biomass of terrestrial species. Experiments of this kind can assist in predicting vegetation response to water-level variation in natural and modified wetlands.

The evolution of vegetative desiccation tolerance in land plants

Abstract: Vegetative desiccation tolerance is a widespread but uncommon occurrence in the plant kingdom generally. The majority of vegetative desiccation-tolerant plants are found in the less complex clades that constitute the algae, lichens and bryophytes. However, within the larger and more complex groups of vascular land plants there are some 60 to 70 species of ferns and fern allies, and approximately 60 species of angiosperms that exhibit some degree of vegetative desiccation tolerance. In this report we analyze the evidence for the differing mechanisms of desiccation tolerance in different plants, including differences in cellular protection and cellular repair, and couple this evidence with a phylogenetic framework to generate a working hypothesis as to the evolution of desiccation tolerance in land plants. We hypothesize that the initial evolution of vegetative desiccation tolerance was a crucial step in the colonization of the land by primitive plants from an origin in fresh water. The primitive mechanism of tolerance probably involved constitutive cellular protection coupled with active cellular repair, similar to that described for modern-day desiccation-tolerant bryophytes. As plant species evolved, vegetative desiccation tolerance was lost as increased growth rates, structural and morphological complexity, and mechanisms that conserve water within the plant and maintain efficient carbon fixation were selected for. Genes that had evolved for cellular protection and repair were, in all likelihood, recruited for different but related processes such as response to water stress and the desiccation tolerance of reproductive propagules. We thus hypothesize that the mechanism of desiccation tolerance exhibited in seeds, a developmentally induced cellular protection system, evolved from the primitive form of vegetative desiccation tolerance. Once established in seeds, this system became available for induction in vegetative tissues by environmental cues related to drying. The more recent, modified vegetative desiccation tolerance mechanism in angiosperms evolved from that programmed into seed development as species spread into very arid environments. Most recently, certain desiccation-tolerant monocots evolved the strategy of poikilochlorophylly to survive and compete in marginal habitats with variability in water availability.

Vegetation and microclimatic edge effects in two mixed-mesophytic forest fragments

Abstract: Forest edges are known to consist of microenvironments that may provide habitat for a different suite of species than forest interiors. Several abiotic attributes of the microenvironment may contribute to this change across the edge to center gradient (e.g., light, air temperature, soil moisture, humidity). Biotic components, such as seed dispersal, may also give rise to changes in species composition from forest edge to interior. We predicted that abiotic and biotic measures would correlate with distance from forest edge and would differ among aspects. To test these predictions, we measured abiotic and biotic variables on twelve 175 m transects in each of two 24 ha forest fragments in east-central Illinois that have remained in continuous isolation for upwards of 100 years. Both univariate and multivariate techniques were used to best describe the complex relationships among abiotic factors and between abiotic and biotic factors. Results indicate that microclimatic variables differ in the degree to and distance over which they show an edge effect. Relative humidity shows the widest edge, while light and soil moisture have the steepest gradients. Aspect influences are evidenced by the existence of more pronounced edge effects on south and west edges, except when these edges are protected by adjacent habitat. Edges bordered by agricultural fields have more extreme changes in microclimate than those bordered by trees. According to PCA results, species richness correlates well with microclimatic variation, especially light and soil moisture; however, in many cases species richness had a different depth of edge influence than either of these variables. The herbaceous plant community is heavily dominated by three species. Distributions of individual species as well as changes in plant community composition, estimated with a similarity index, indicate that competition may be influencing the response of the vegetation to the edge to interior gradient. This study indicates that edge effects must be considered when the size and potential buffering habitat of forest preserves are planned.

Dynamic and static views of succession: Testing the descriptive power of the chronosequence approach

Abstract: Chronosequence and permanent plot studies are the two most common methods for evaluating successional dynamics in plant communities. We combined these two approaches by re-sampling an old-field chronosequence at Cedar Creek Natural History Area (Minnesota, USA) to: (1) measure rates of secondary succession; and (2) to test the ability of the chronosequence approach to predict actual successional dynamics over a 14-year survey interval. For each of 19 chronosequence fields we calculated four complimentary indices of succession rate for community changes that actually occurred within each of these fields between 1983 and 1997. We found that measures of compositional dissimilarity, species turnover, and the change rates of perennial and native species cover over this 14-year period were all negatively correlated with field age, indicating that the rate of successional change in these old-fields generally declines over time. We also found that data collected from the initial static chronosequence survey (1983) accurately predicted many of the observed changes in species abundance that occurred between 1983 and 1997, but was a poor predictor of changes in species richness. In general, chronosequence re-sampling confirmed the validity of using the chronosequence approach to infer basic patterns of successional change.

Vulnerability of riparian zones to invasion by exotic vascular plants

Abstract: We compared the invasibility of riparian plant communities high on river banks with those on floodplain floors for four South African rivers. Analyses of abundant and significant riparian species showed that the floors have 3.1 times more exotic plants than the banks. The percent exotics ranges from 5% to 11% of total species richness for the banks, and from 20% to 30% for the floors. Species richness and percent exotics are negatively correlated for the banks, but not correlated for the floors. Despite great differences in climate, species richness, and landuse history, the percentages of exotic plants in three rivers in the Pacific Northwest of the USA and one river in southwestern France are similar to those in South Africa (24-30% vs. 20-30%, respectively). Furthermore, the high proportions of exotic species in these riparian plant communities are comparable to those reported for vascular plant communities on islands. We conclude that the macro-channel floor regions of the riparian zones of South African rivers are highly vulnerable to invasion by exotic vascular plants.

Ecological buffering mechanisms in savannas : a unifying theory of long-term tree-grass coexistence

Abstract: Despite the large spatial extent and the obvious importance of the savanna biome, not to mention several decades of savanna research, the origin, age, nature, and dynamics of savannas are not well understood. Basically, the question surrounding the presence or existence of savannas focuses on the long-term coexistence of the dominating life forms – trees and grasses. How do these two very different components coexist, without one of them dominating the other, and what mechanisms determine the proportion of each? Earlier equilibrium concepts have recently been replaced by non-equilibrium concepts, and the current view is that tree-grass interactions in savannas cannot be predicted by a simple model. Instead, many interacting factors operating at various spatial and temporal scales contribute to creating and maintaining savanna physiognomy. In this paper we analyse a number of studies from savannas in different parts of the world and discuss whether a general pattern can be perceived behind the numerous factors influencing the presence of savannas systems. On the basis of this analysis we propose a new unifying concept of savanna existence, i.e., the concept of ecological buffering mechanisms. In contrast to previous approaches to explain tree-grass coexistence in savannas, the concept of buffering mechanisms does not focus on equilibria or non-equilibria, steady states of the system or domains of attraction. Instead, in the concept of ecological buffering mechanisms we suggest that it is much more useful to focus on the boundaries of savanna existence itself and to investigate the mechanisms that allow a savanna to persist in critical situations where this system is driven to its boundaries, e.g., pure grasslands or tropical forests. The concept of ecological buffering mechanisms integrates both earlier concepts of ecological theory and general ideas on savanna dynamics as well as specific studies of savannas in different parts of the world.

A comparison of mechanisms of desiccation tolerance among three angiosperm resurrection plant species

Abstract: The mechanisms of protection against mechanical and oxidative stress were identified and compared in the angiosperm resurrection plants Craterostigma wilmsii, Myrothamnus flabellifoliusand Xerophyta humilis. Dryinginduced ultrastructural changes within mesophyll cells were followed to gain an understanding of the mechanisms of mechanical stabilisation. In all three species, water filled vacuoles present in hydrated cells were replaced by several smaller vacuoles filled with non-aqueous substances. In X. humilis, these occupied a large proportion of the cytoplasm, preventing plasmalemma withdrawal and cell wall collapse. In C. wilmsii, vacuoles were small but extensive cell wall folding occurred to prevent plasmalemma withdrawal. In M. flabellifolius ,s ome degree of vacuolation and wall folding occurred, but neither were sufficient to prevent plasmalemma withdrawal. This membrane was not ruptured, possibly due to membrane repair at plasmodesmata junctions where tearing might have occurred. In addition, the extra-cytoplasmic compartment appeared to contain material (possibly similar to that in vacuoles) which could facilitate stabilisation of dry cells. Photosynthesis and respiration are particularly susceptible to oxidative stress during drying. Photosynthesis ceased at high water contents and it is proposed that a controlled shut down of this metabolism occurred in order to minimise the potential for photo-oxidation. The mechanisms whereby this was achieved varied among the species. In X. humilis, chlorophyll was degraded and thylakoid membranes dismantled during drying. In both C. wilmsii and M. flabellifolius, chlorophyll was retained, but photosynthesis was stopped due to chlorophyll shading from leaf folding and anthocyanin accumulation. Furthermore, in M. flabellifolius thylakoid membranes became unstacked during drying. All species continued respiration during drying to 10% relative water content, which is proposed to be necessary for energy to establish protection mechanisms. Activity of antioxidant enzymes increased during drying and remained high at low water contents in all species, ameliorating free radical damage from both photosynthesis and respiration. The nature and extent of antioxidant upregulation varied among the species. In C. wilmsii, only ascorbate peroxidise activity increased, but in M. flabellifolius and X. humilis ascorbate peroxidise, glutathione reductase and superoxide dismutase activity increased, to various extents, during drying. Anthocyanins accumulated in all species but this was more extensive in the homoiochlorophyllous types, possibly for protection against photo-oxidation.

Granitic and gneissic outcrops (inselbergs) as centers of diversity for desiccation-tolerant vascular plants

Abstract: Although desiccation tolerance is common in non-vascular plants, this adaptive trait is very rare in vascular plants. Desiccation-tolerant vascular plants occur particularly on rock outcrops in the tropics and to a lesser extent in temperate zones. They are found from sea level up to 2800 m. The diversity of desiccation-tolerant species as measured by number of species is highest in East Africa, Madagascar and Brazil, where granitic and gneissic outcrops, or inselbergs, are their main habitat. Inselbergs frequently occur as isolated monoliths characterized by extreme environmental conditions (i.e., edaphic dryness, high degrees of insolation). On tropical inselbergs, desiccation-tolerant monocotyledons (i.e., Cyperaceae and Velloziaceae) dominate in mat-like communities which cover even steep slopes. Mat-forming desiccation-tolerant species may attain considerable age (hundreds of years) and size (several m in height, for pseudostemmed species). Both homoiochlorophyllous and poikilochlorophyllous species occur. In their natural habitats, both groups survive dry periods of several months and regain their photosynthetic activity within a few days after rainfall. Other desiccation-tolerant species colonize shallow depressions, crevices and even temporarily water-filled rock pools on inselbergs. Desiccation-tolerant vascular plants occur in 13 families and are best represented within the monocotyledons and ferns. Only a few desiccation-tolerant dicots exist, in the Gesneriaceae, Myrothamnaceae and Scrophulariaceae. In total, about 330 species of vascular desiccation-tolerant plants are known, of which nearly 90% occur on inselbergs. With regard to morphological adaptations, the mat-forming monocotyledons are particularly remarkable due to the possession of roots with a velamen radicum, which is reported here in the genus Borya for the first time.

Diversity, composition, and structure of tropical dry forests in Central America

Abstract: Tropical dry forests have been reduced to less than 0.1% of their original expanse on the Pacific side of Central America and are considered by some to be the most endangered ecosystem in the lowland tropics. Plots 1000 m2 were established in seven tropical dry forests in Costa Rica and Nicaragua in order to compare levels of species richness to other Neotropical dry forest sites and to identify environmental variables associated with species richness and abundance. A total of 204 species and 1484 individuals ≥ 2.5 cm were encountered. Santa Rosa National Park was the richest site with the highest family (33), genera (69), and species (75) diversity of all sites. Species richness and forest structure were significantly different between sites. Fabaceae was the dominant tree and shrub family at most sites, but no species was repeatably dominant based on number of stems in all fragments of tropical dry forest. Central American dry forests had similar species richness when compared to other Neotropical forests. There was no correlation between forest cover within reserves, or precipitation and plant species richness. There was a significant correlation between anthropogenic disturbance (intensity and frequency of fire, wood collection, grazing) and total species richness, tree and shrub species richness, and liana abundance. These results suggest controlling levels on anthropogenic disturbance within reserves should be a high priority for resource managers in Central America. Further research in forest fragments which examine individual and a combination of disturbance agents would help clarify the importance of anthropogenic disturbance on species richness and abundance.

Effects of long-term rainfall variability on evapotranspiration and soil water distribution in the Chihuahuan Desert: A modeling analysis

Abstract: We used the patch arid land simulator (PALS-FT) – a simple, mechanistic ecosystem model – to explore long-term variation in evapotranspiration (ET) as a function of variability in rainfall and plant functional type (FT) at a warm desert site in southern New Mexico. PALS-FT predicts soil evaporation and plant transpiration of a canopy composed of five principal plant FTs: annuals, perennial forbs, C4 grasses, sub-shrubs, and evergreen shrubs. For each FT, the fractional contribution to transpiration depends upon phenological activity and cover as well as daily leaf stomatal conductance, which is a function of plant water potential, calculated from root-weighted soil water potential in six soil layers. Simulations of water loss from two plant community types (grass- vs. shrub-dominated) were carried out for the Jornada Basin, New Mexico, using 100 years of daily precipitation data (1891–1990). In order to emphasize variability associated with rainfall and fundamental differences in FT composition between communities, the seasonal patterns cover of perennials were held constant from year to year. Because the relative amount of year to year cover of winter and summer annual species is highly variable in this ecosystem, we examined their influence on model predictions of ET by allowing their cover to be variable, fixed, or absent. Over the entire 100-yr period, total annual ET is highly correlated with total annual rainfall in both community types, although T and E alone are less strongly correlated with rainfall, and variation in transpiration is nearly 3 times greater than evaporation and 2 times greater than variation in rainfall (CV of rainfall = 35%). Water use shows a relatively high similarity between the grass- and shrub-dominated communities, with a 100-yr average T/ET of 34% for both communities. However, based on a year-by-year comparison between communities, T/ET was significantly greater in the grass-dominated community, reflecting the fact that over the long term more than half of the rain occurs in the summer and is used slightly more efficiently (T?E) by the C4-grass community than the shrub community, although we found some rainfall patterns that resulted in much greater T/ET in the shrub community in a given year. Percent of water lost as transpiration (T/ET) suggests that while there is a general trend toward increased T/ET with rainfall in both community types, T/ET is extremely variable over the 100-yr simulation, especially for normal and below normal amounts of rainfall (T/ET values range from 1 to 58% for the grass-dominated site and 6 to 60% for the shrub-dominated site). These predictions suggest that because of the relatively shallow distribution of soil water, there is little opportunity for vertical partitioning of the soil water resource by differential rooting depths of the plant FTs, in contrast to the two-layer hypothesis of Walter (1971). However, functional types may avoid competition by keying on particular `windows' of moisture availability via differences in phenologies. We found very little differences in average, long-term model predictions of T, E, and ET when annual plant cover was variable, fixed, or absent. The results of our simulations help reconcile some of the disparate conclusions drawn from experimental studies about the relative contribution of transpiration vs. evaporation to total evapotranspiration, primarily by revealing the great year-to-year variability that is possible.

Elevational gradients in species richness and endemism of selected plant groups in the central Bolivian Andes.

Abstract: I analyzed the distribution of Acanthaceae, Araceae, Bromeliaceae, Cactaceae, Melastomataceae, and Pteridophyta in 62 vegetation plots of 400 m2 along an elevational transect between 500 m and 2450 m, and at a nearby lowland site in western Santa Cruz department, Bolivia. These groups were selected because they are physiognomically distinctive, have high species numbers, are comparatively easy to identify, adequately reflect overall floristic relationships, include a wide range of life forms, and are small. The transect was located in the Tucumano-Boliviano biogeographic zone and included drought-deciduous ( 1800 m) forests. Elevational patterns of species richness were group-specific and probably related to the ecophysiological properties of each group. Species richness in Pteridophyta and Melastomataceae was correlated with moss cover (i.e., humidity), with elevation (i.e., temperatures) in Acanthaceae and epiphytic Bromeliaceae, with potential evapotranspiration (i.e., ecosystem productivity) in Araceae, and with light availability at ground level in terrestrial Bromeliaceae and Cactaceae. Community endemism generally increased with elevation, but showed a maximum at 1700 m for terrestrial Pteridophyta, and a nonsignificant decline for epiphytic Bromeliaceae and Cactaceae. Endemism was higher for terrestrial than for epiphytic taxa, and was lower among Pteridophyta compared to all other groups, reflecting different dispersal ability among taxonomic and ecological groups. Elevational zonation, tested against a null-model of random distribution of elevational limits, revealed a significant accumulation of upper and lower elevational range boundaries at 900–1050 m and at 1500–1850 m, corresponding to the elevational limits of the main physiognomic vegetation types.

Multiscale soil and vegetation patchiness along a gradient of herbivore impact in a semi-arid grazing system in West Africa

Abstract: We studied the degree and scale of patchiness of vegetation and selected soil variables along a gradient of herbivore impact. The gradient consisted of a radial pattern of ‘high’, ‘intermediate’ and ‘low’ herbivore impact around a watering point in a semi-arid environment in Burkina Faso (West Africa). We hypothesised that, at a certain range of herbivore impact, vegetated patches alternating with patches of bare soil would occur as a consequence of plant-soil feedbacks and run-off-run-on patterns. Indeed, our transect data collected along the gradient showed that vegetated patches with a scale of about 5‐10 m, alternating with bare soil, occurred at intermediate herbivore impact. When analysing the data from the experimental sites along the gradient, however, we also found a high degree of patchiness of vegetation and soil variables in case of low and high herbivore impact. For low herbivore impact, most variation was spatially explained, up to 100% for vegetation biomass and soil temperature, with a patch scale of about 0.50 m. This was due to the presence of perennial grass tufts of Cymbopogon schoenanthus . Patterns of soil organic matter and NH4-N were highly correlated with these patterns of biomass and soil temperature, up to r D 0: 7( P< 0:05) for the in situ correlation between biomass and NH4-N. For high herbivore impact, we also found that most variation was spatially explained, up to 100% for biomass and soil temperature, and 84% for soil moisture, with three distinct scales of patchiness (about 0.50 m, 1.80 m and 2.80 m). Here, microrelief had a corresponding patchy structure. For intermediate herbivore impact, again most variation was spatially explained, up to 100% for biomass and soil temperature, and 84% for soil moisture, with a patch scale of about 0.95 m. Here, we found evidence that vegetated patches positively affected soil moisture through less run-off and higher infiltration of rainwater that could not infiltrate into the bare soil elsewhere, which was not due to microrelief. Thus, we conclude that our findings are in line with our initial hypothesis that, at intermediate herbivore impact, vegetated patches alternating with patches of bare soil persist in time due to positive plant-soil feedbacks.

The discovery, scope, and puzzle of desiccation tolerance in plants

Abstract: The modern scientific study of desiccation tolerance began in 1702 when Anthony von Leeuwenhoek discovered that rotifers could survive without water for months By 1860, the controversy over whether organisms could dry up without dying had reached such a pitch that a special French commission was convened to adjudicate the dispute In 2000, we know that a few groups of animals and a wide variety of plants can tolerate desiccation in the active, adult stages of their life cycles Among plants, this includes many lichens and bryophytes, a few ferns, and a very few flowering plants, but no gymnosperms nor trees Some desiccation-tolerant species can survive without water for over ten years, recover from desiccation to unmeasurably low water potentials, and, when plants are desiccated, endure temperature extremes from −272 to 100 °C Desiccation-tolerant plants occur on all continents but mainly in xeric habitats or microhabitats where the cover of desiccation-sensitive species is low Two main puzzles arise from these patterns: What are the mechanisms by which plants tolerate desiccation? and Why are desiccation-tolerant plants not more ecologically widespread? Recent molecular and biochemical studies suggest that there are multiple mechanisms of tolerance, many of which involve protection from oxidants and from the loss of configuration of macromolecules during dehydration Hypotheses to explain the restricted ecological range of desiccation-tolerance plants include inability to maintain a cumulative positive carbon balance during repeated cycles of wetting and drying and inherent trade offs between desiccation tolerance and growth rate

The bryophyte paradox: tolerance of desiccation, evasion of drought

Abstract: Vascular plants represent one strategy of adaptation to the uneven and erratic supply of water on land. Desiccation-tolerant (DT) bryophytes represent an alternative, photosynthesising and growing when water is freely available, and suspending metabolism when it is not. By contrast with vascular plants, DT bryophytes are typically ectohydric, carrying external capillary water which can vary widely in quantity without affecting the water status of the cells. External water is important in water conduction, and results in bryophyte leaf cells functioning for most of the time at full turgor; water stress is a relatively brief transient phase before full desiccation. All bryophytes are C3 plants, and their cells are essentially mesophytic in important physiological respects. Their carbohydrate content shows parallels with that of maturing embryos of DT seeds. Initial recovery from moderate periods of desiccation is very rapid, and substantial elements of it appear to be independent of protein synthesis. Desiccation tolerance in effect acts as a device that evades the problems of drought, and in various adaptive features DT bryophytes are more comparable with (mesic) desert ephemerals or temperate winter annuals (but on a shorter time scale, with DT vegetative tissues substituting for DT seeds) than with drought-tolerant vascular plants.

Effects of extreme high temperature, drought and elevated CO2 on photosynthesis of the Mojave Desert evergreen shrub, Larrea tridentata

Abstract: The interaction of extreme temperature events with future atmospheric CO2 concentrations may have strong impacts on physiological performance of desert shrub seedlings, which during the critical establishment phase often endure temperature extremes in conjunction with pronounced drought. To evaluate the interaction of drought and CO2 on photosynthesis during heat stress, one-year-old Larrea tridentata[DC] Cov. seedlings were exposed to nine days of heat with midday air temperature maxima reaching 53 °C under three atmospheric CO2 concentrations (360, 550 and 700 μmol mol−1) and two water regimes (well-watered and droughted). Photosynthetic gas exchange, chlorophyll fluorescence and water potential responses were measured prior to, during and one week following the high temperature stress event. Heat stress markedly decreased net photosynthetic rate (A net), stomatal conductance (g s), and the photochemical efficiency of photosystem II (F v/F m) in all plants except for well-watered L. tridentata grown in 700 μmol mol−1 CO2. A net and g s remained similar to pre-stress levels in these plants. In droughted L. tridentata, A net was ca. 2× (in 550 μmol mol−1 CO2) to 3× (in 700 μmol mol−1 CO2) higher than in ambient-CO2-grown plants, while g s and F v/F m were similar and low in all CO2 treatments. Following heat stress, g s in all well-watered plants rose dramatically, exceeding pre-stress levels by up to 100%. In droughted plants, g s and A net rose only in plants grown at elevated CO2 following release from heat. This recovery response was strongest at 700 μmol mol−1 CO2, which returned to A net and g s values similar to pre-heat following several days of recovery. Extreme heat diminished the photosynthetic down-regulation response to growth at elevated CO2 under well-watered conditions, similar to the action of drought. Ambient-CO2-grown L. tridentata did not show significant recovery of photosynthetic capacity (A \max and CE) after alleviation of temperature stress, especially when exposed to drought, while plants exposed to elevated CO2 appeared to be unaffected. These findings suggest that elevated CO2 could promote photosynthetic activity during critical periods of seedling establishment, and enhance the potential for L. tridentata to survive extreme high temperature events.

Hydrochory, seed banks, and regeneration dynamics along the landscape boundaries of a forested wetland

Abstract: Following the environmental sieve concept, the setting in which the recruitment of Taxodium distichum occurs in, becomes increasingly restrictive from the seed to seedling stage in an impounded forested wetland. Although a wide elevational band of dispersing seed moves across the boundary of a swamp-field in the water sheet, the zone of germination is relegated to that portion of the forested wetland that draws down during the growing season. Seedling recruitment is further restricted to the uppermost zone of the winter water sheet. These patterns are likely applicable to other species of dominant swamp species, e.g., Cephalanthus occidentalis crossed the boundary of a forested wetland and abandonded field in winter flooding (November–December and November–March, respectively) in Buttonland Swamp. The elevation of the boundary was 101.3 m NGVD. While the seeds of at least 40 swamp species were dispersed across the boundary, few viable seeds were dispersed after the winter season. Kriged maps showed seeds of T. distichum and C. occidentalis dispersed in patches in the water depending on the position of the water sheet. Most species of both water- and gravity-dispersed species had a localized pattern of seed distribution (either spherical or exponential) and this indicated that seeds may not be dispersed for great distances in the swamp. Water-dispersed T. distichum and C. occidentalis had larger dispersal ranges (A0=225 and 195 m, respectively) than Bidens frondosa and B. discoidea (A0=14 and 16 m, respectively). Seed dispersal varied with season depending on the availability of seeds. In Buttonland Swamp, viable seeds typically were dispersed for T. distichum in November–June, and for C. occidentalis in November-July. Low water occurred in August 1993 and high in February 1994 (99.8 and 101.6 m NGVD, respectively). The seed banks along the landscape boundary varied in species composition according to elevation (r2 = 0.996). While the similarity of species richness between water-dispersed seeds and the seed bank at elevations that flooded (during June 1993 through May 1995) was high (10–17%), it was low between water-dispersed seeds and the seed bank at elevations that did not flood (5%). T. distichum seeds had a short germination window in that seeds germinated within a year following their production in zones that were flooded in the winter followed by drawdown during the next growing season. After 1 year, less than 5% of the T. distichum seeds remained viable on the surface of the soil. Germination of T. distichum was confined to specific elevations (above 99.3 but below 101.6 m NGVD) during this study with 4.1% of the seedlings surviving for more than 2 years at a mean of 101.4 m NGVD. All seedlings below this elevation died. To maximize natural regeneration along the boundaries of swamps in abandoned farm fields targeted for restoration, this study suggests a flood pulse regime consisting of high water in the winter to maximize dispersal of live seeds followed by low water in the summer to facilitate seed germination and seedling recruitment. Hydrologic restoration could assist in the natural recovery of damaged wetlands if a seed source exists nearby.

Seed germination response to cold stratification period and thermal regime in Phacelia secunda (Hydrophyllaceae) – Altitudinal variation in the mediterranean Andes of central Chile

Abstract: The ability to germinate under a variety of environmental conditions is essential for plant species inhabiting a wide range of altitudes and latitudes. Phacelia secunda J. F. Gmel. (Hydrophyllaceae) is a perennial herb with wide latitudinal and altitudinal distributional ranges. In the central Chilean Andes (33 °S) P. secunda can be found from 1600 m sealevel up to the vegetation limit at 3400 m. It has been suggested that seeds from populations encountering long periods with snow cover and adverse winter conditions would require longer periods of cold stratification for germination than those from populations exposed to milder winters. Given that the snow-free period decreases with elevation, seeds from high elevation populations could require longer period of cold stratification to germinate. Moreover, it has been shown that seeds from arctic and higher elevations environments are adapted to germinate better under high temperature conditions. Germination response with increasing periods of cold stratification (0–6 mo.) and under two contrasting thermoperiods (20 °/1O °C; 10 °/5 °C; 12 h day/night), were studied for 4 populations of P. secunda located at 1600, 2100, 2900 and 3400 m a.s.l. Initiation of germination required increasingly longer periods of stratification with elevation, and proportionately fewer seeds germinated for any one stratification treatment at the higher elevations. Seeds from higher elevations germinated to a higher percentage under the high than the low temperature thermoperiods. These results illustrates a significant variation in germination characteristics over a spatially short environmental gradient.

Seed arrival under different genera of trees in a neotropical pasture

Abstract: Trees in pastures attract seed dispersers, leading to increased seed arrival under their canopies and more rapid regrowth around them The characteristics that make some trees better `recruitment foci' than others, however, are poorly understood In a neotropical pasture, we examined the arrival of seeds to open areas and underneath four genera of trees that varied in canopy architecture and type of fruit produced: Ficus trees had dense canopies and fleshy fruits, Pentaclethra trees had dense canopies and dry fruits, Cecropia trees had sparse canopies and fleshy fruits, and Cordia trees had sparse canopies and dry fruits We found that all trees received more seeds than open pasture, probably because trees provided seed dispersers with better perches, protection from predators, nesting sites, etc Among the tree genera, more seeds arrived under trees that produced fleshy fruits than trees that did not This occured even during periods when trees were not fruiting (ie, non-fruiting Ficus and Cecropia trees received more seeds than Cordia or Pentaclethra trees) Seed dispersers may periodically check Ficus and Cecropia trees for fruits, or they may become familiar with these trees while feeding and thereafter use them for other reasons Height of trees had a slight positive effect on seed arrival, possibly because taller trees offered more protection from predators Canopy architecture and distance to forest edge did not significantly affect seed arrival This study demonstrates that trees in general are potentially important recruitment foci, but that different types of trees vary in the kind of recruitment that they foster in pastures

Effect of microbiotic soil surface crusts on emergence of vascular plants

Abstract: Microbiotic crusts are a common and widespread feature of arid and semi-arid landscapes. Their effect on vascular plant success has been discussed controversially. However, only very few field studies have yet tried to experimentally investigate the effect of microbiotic crusts on vascular plant establishment. In this study we investigate the influence of cyanobacteria dominated microbiotic soil surface crust on vascular plants in a desert sand dune area by using a series of manipulative experiments. Crusts were disturbed and removed and density response to these treatments was monitored during the two following years. Emergence densities of vascular plants were considerably higher when crusts were removed and destroyed. This effect was particularly pronounced when the disturbances were applied before the main seed dispersal period. By experimentally preventing seed dispersal into disturbed areas we could show that undisturbed crusts reduce the probability for seeds to come to rest. This indicates that soil surface roughness is a major determinant for the establishment of vascular plants in the study area. We conclude that small-scale and well-timed disturbances of microbiotic crusts in arid lands may increase vascular plant establishment probabilities. A potential feedback process of long-term vegetation dynamics is suggested: disturbance of microbiotic crusts results in higher numbers of emerging plants, which in turn represent a rough seed-trapping element leading to a further increase in densities in consecutive years.

Bryophytes as experimental models for the study of environmental stress tolerance: Tortula ruralis and desiccation-tolerance in mosses

Abstract: The development of a complete understanding of how plants interact with the environment at the cellular level is a crucial step in advancing our ability to unravel the complexities of plant ecology particularly with regard to the role that many of the less complex plants (i.e., algae, lichens, and bryophytes) play in plant communities and in establishing areas for colonization by their more complex brothers. One of the main barriers to the advancement of this area of plant biology has been the paucity of simple and appropriate experimental models that would enable the researcher to biochemically and genetically dissect the response of less complex plants to environmental stress. A number of bryophytes model systems have been developed and they have been powerful experimental tools for the elucidation of complex biological processes in plants. Recently there has been a resurgent interest in bryophytes as models systems due to the discovery and development of homologous recombination technologies in the moss Physcomitrella patens (Hedw.) Brach & Schimp. In this report we introduce the desiccation-tolerant moss Tortula ruralis (Hedw.) Gaert., Meyer, and Scherb, as a model for stress tolerance mechanisms that offers a great deal of promise for advancing our efforts to understand how plants respond to and survive the severest of stressful environments. T. ruralis, a species native to Northern and Western North America, has been the most intensely studied of all bryophytes with respect to its physiological, biochemical, and cellular responses, to the severest of water stresses, desiccation. It is our hope that the research conducted using this bryophyte will lay the foundationfor not only the ecology of bryophytes and other less complex plants but also for the role of desiccation-tolerance in the evolution of land plants and the determination of mechanisms by which plant cells can withstand environmental insults. We will focus the discussion on the research we and others have conducted in an effort to understand the ability of T. ruralis to withstand the complete loss of free water from the protoplasm of its cells.

Gap-phase regeneration in a tropical montane forest: the effects of gap structure and bamboo species

Abstract: To study the influence of gap structure and bamboo species on the regrowth of montane Atlantic forest, colonization by plants was characterized in 30 treefall gaps (30.3–500.5 m2). The study was conducted at Santa Virginia (45°30′ W, 23°17′ S), a 4970-ha reserve of Atlantic montane forest in southeastern Brazil. Area covered by bamboos ranged from 0% to 100% of gap area. Average height of surrounding canopies ranged from 12 to 30 m. As gap are covered by bamboo and average height of surrounding canopies increased, both density and richness of pioneer woody species decreased. Density and richness of shade-tolerant species were negatively influenced by gap area. Low-light-demanding species of Miconia, Leandra and Rapanea accounted for the majority of both pioneer species and individuals sampled, whereas high-light demanding pioneers of Cecropia, Alchornea and Tibouchina were poorly represented. We suggest that in the Atlantic montane forest bamboo species compete for gaps, excluding other light-demanding pioneers. This results in an overall reduction of pioneer species richness in the Atlantic forest.

Inhibitory effects of Artemisia herba-alba on the germination of the gypsophyte Helianthemum squamatum

Abstract: The potential allelopathic role of Artemisia herba-alba has been evaluated in order to explain the community pattern of the gypsum semiarid environments of central Spain. This pattern shows a sharp ecotone between a gypsophile sparse shrubby community dominated by Helianthemum squamatum, which grows on slopes with gypsum surface crusts, and a nitrohalophilous community on the gypsum alluvial soils of piedmont dominated by Artemisia herba-alba. In order to explain this pattern, resource limitation was discarded because no significant differences in several soil parametrers, but fine earth fraction and organic matter content, had been detected in a previous study. Results confirm the inhibitory effect of aqueous extracts on the final germination percentage of scarified seeds of Helianthemum squamatum and also on the shape of the germination curves, which indicate delay of germination for the aqueous extracts. Assays with soil of the Artemisiacommunity (three types) showed that germination was strongly inhibited in soils obtained below the canopy of mature plants (litter maintained) and retarded in the other treatments. We suggest that the spatial community pattern detected in this gypsum environment and characterised by a sharp ecotone could be at least partially controlled by interference through allelopathy. This determines the excliusion of Helianthemum squamatum plants from alluvial soils.

The influence of tidal channels on the distribution of salt marsh plant species in Petaluma Marsh, CA, USA

Abstract: Tidal channels influence the distribution and composition of salt marsh vegetation in a San Francisco Bay salt marsh. Two channel networks in the Petaluma Marsh, Sonoma County, CA, were mapped and characterized using global positioning and geographic information systems. Plant species abundance was sampled on transects placed perpendicular to and extending away from the channel banks. The vegetation showed significant increases in species richness along channel banks and larger areas of effect which increased approximately linearly with channel size. Composition of species assemblages varies with distance from the channel bank and channel size. These results demonstrate that salt marsh plant assemblages, composed of both major and minor species, are distributed with respect to the channel network in Petaluma Marsh.

Population structure and spatial patterns for trees in a temperate old-growth evergreen broad-leaved forest in Japan

Abstract: The population structure and spatial pattern of major tree species in a warm-temperate old-growth evergreen broad-leaved forest in the Tatera Forest Reserve of Japan were investigated. All stems ≥ 5 cm in diameter at breast height (DBH) were mapped on a 4 ha plot and analyses were made of population structure and the spatial distribution and spatial association of stems in different vertical layers for nine species. This was done in the context of scale dependency. The plot was located on a very gentle slope and 17.1% of its canopy layer was in gaps. It contained 45 woody plant species and 4570 living stems with a basal area of 63.9 m2 ha−1. Castanopsis cuspidata var. sieboldii, the most dominant species for the basal area, had the maximum DBH among the species present, fewer smaller stems and a lower coefficient of statistical skewness of the DBH distribution. The second most dominant species, Dystilium racemosum, had the highest stem density (410 ha−1), more abundant smaller stems and a relatively higher coefficient of skewness. Most stems in different vertical layers showed a weakly aggregated distribution with loose colonies as basic units. Gap dependency for the occurrence of stems under the canopy layer was weak. Maximum slope degree of the plot also weakly affected the occurrence of stems. Spatial associations varied among intra- and interspecific cohorts in the different layers and spatial scales examined, and positive associations among cohorts were found more frequently as the scales examined became larger. This tendency suggests that key factors forming observed spatial associations might vary with the spatial scales.

Ecophysiological differences between male and female plants of Pistacia lentiscus L.

Abstract: Previous studies in spatial distribution of male and female shrubs of Pistacia lentiscus have demonstrated that less perturbed areas, older communities with a well developed cover, have male-biased sex ratios, whereas in abandoned old agricultural areas there are no significant differences between the number of male and female plants. In this study, we analyse both sexes in terms of their photosynthetic features that could provide a physiological basis for habitat partitioning between sexes. Rates of light-saturated assimilation and stomatal conductance were studied in male and female plants during summer. Assimilation rates were higher in the morning than in the afternoon and mean daily maximum assimilation rates reached 10.9 and 6.6 μmol m−2 s−1, for male and female plants, respectively. In the absence of drought stress (laboratory conditions), the measured photosynthetic characteristics of leaves of male and female plants, provided by fluorescence studies and light and CO2 response curves, were similar. Under natural stress conditions however, lower CO2 assimilation rates and stomatal conductances were recorded in female plants. The differences in the light response curve of effective quantum yield (ΦII) recorded under stress conditions showed also higher quantum yield for male plants under low irradiances. From this study we suggest that the differences observed between male and females are largely due to different degrees of stomatal control rather than to differences in photosynthetic activity, leading to higher water use efficiency (WUE) in female plants. However, despite the higher leaf control of water loss by females, they reduce the water potential to the same values as male plants, probably due to specific characteristics of the root system or of the conducting xylem. These results suggest that the ecological advantage of male plants in older communities is due to a higher competition for water uptake, while in the youngest open areas is the higher WUE in female plants that confer an ecological advantage.

Effects of Hurricane Andrew on stands of slash pine (Pinus elliottii var. densa) in the everglades region of south Florida (USA)

Abstract: Few hurricanes affect intact stands of subtropical pines. We examined effects of winds in the eyewalls of Hurricane Andrew, where wind speeds were>200 km h 1 , on all remaining large mainland stands of Pinus elliottii var. densa (south Florida slash pine) on limestone outcroppings (rocklands) in the everglades region of southern Florida. We measured densities and sizes of trees and assessed damage and mortality in plots in old-growth stands in the Lostman’s Pines (LOP) region of Big Cypress National Preserve and in second-growth stands in the Pines West (PIW) and Long Pine Key (LPK) regions of Everglades National Park. We also examined age-size relationships using sections from trees killed by the hurricane in LOP and LPK. We used the data to predict effects of recurrent hurricanes on the structure and dynamics of the old-growth stand and to compare effects of hurricanes on old- and second-growth stands. Slash pine was resistant to hurricane winds. Most trees in stands (68‐76%) were not severely damaged; mortality in the three regions averaged 17‐25% shortly after the hurricane and 3‐7% during the following year. Mortality was positively associated with tree size; mean tree sizes decreased and size-selective thinning occurred in all stands. Nonetheless, local mortality ranged from 3‐4% to 50‐60% among plots in all stands. Such local variation in mortality resulted from clustering of large trees, especially in old-growth stands, and from microbursts during the hurricane, which affected all stands. Recurrent, intense hurricanes are predicted to kill larger trees, slowly opening new patches and increasing sizes of extant patches, thus resulting in almost continual presence of openings suitable for recruitment in old-growth stands. Age-size relationships also indicated that large trees in old-growth stands may survive 2‐3 centuries. The combination of frequent openings and wind resistance of large trees is predicted to result in old-growth stands that are highly uneven aged, with trees locally distributed in similar-aged patches. The extent to which such stands deviate from demographic equilibrium, as well as turnover rates within stands, are likely to increase as the frequency of recurrent, intense hurricanes increases. Damage and mortality differed in old- and second-growth stands. Large trees were more, but small trees less likely to be damaged in old- than second-growth stands. In contrast, mortality was significantly lower in old- (LOP: 16.9% 3.1 [mean s.e.]) than second-growth stands (PIW: 22.5% 2.0; LPK: 25.2% 2.7). Total hurricanerelated mortality was 30‐60% higher in second- than old-growth stands. Size class structure, more uneven in oldthan second growth stands prior to the hurricane, diverged even more afterwards. Hurricane Andrew removed

From snapshot information to long-term population dynamics of acacias by a simulation model.

Abstract: The African Acacia species A. raddiana is believed to be endangered in the Negev desert of Israel. The ecology of this species is not well understood. The main idea of our study is to learn more about the long-term population dynamics of these trees using snapshot information in the form of size frequency distributions. These distributions are highly condensed indices of population dynamics acting over many years. In this paper, we analyse field data on recruitment, growth, and mortality and use an existing simulation model of the population dynamics of A. raddiana (SAM) to produce contrasting scenarios of these live history processes that are based on the analysed field evidence. The main properties of simulated as well as observed tree size frequency distributions are characterised with Simpson's index of dominance and a new permutation index. Finally, by running the SAM model under the different scenarios, we study the effect of these different processes on simulated size frequency distributions (pattern) and we compare them to size distributions observed in the field, in order to identify the processes acting in the field. Our study confirms rare recruitment events as a major factor shaping tree size frequency distributions and shows that the paucity of recruitment has been a normal feature of A. raddiana in the Negev over many years. Irregular growth, e.g., due to episodic rainfall, showed a moderate influence on size distributions. Finally, the size frequency distributions observed in the Negev reveal the information that, in this harsh environment, mortality of adult A. raddiana is independent of tree size (age).

Competition and coexistence of rhizomatous perennial plants along a nutrient gradient

Abstract: I studied competition and coexistence of three tall clonal perennial plant species, Calamagrostis epigejos (L.) Roth, Solidago canadensis L., and Tanacetum vulgare L. along a gradient of soil productivity over five years. A replacement series field experiment was conducted with high, moderate and low fertility levels in 1m×1m plots. There were significant effects of soil type on ramet density (P<0.001), mean height (P<0.01), and total biomass (P<0.01). Ramet density, mean height, and total biomass increased with increasing soil fertility. There were also significant effects of mixture on ramet density (P<0.01), but not on mean height and total biomass for all species. Significant neighbor effects on ramet density and total biomass (P<0.01) were found for Solidago, showing that it is important whether Tanacetum or Calamagrostis is its neighbor within mixtures. During the five years there was only one case of competitive exclusion: Calamagrostis excluded Solidago on the most fertile substrate in the fifth growing season. In most cases species coexisted over the five years. Each of the three species was able to dominate in at least one combination of substrate type and mixture. The experiment showed that asymmetric competition for light on substrates of high fertility, symmetric competition for nutrients on nutrient-poor soil and positive interactions especially on substrates of intermediate fertility played a role. A founder effect was evident in aggregated mixtures of Calamagrostis and Solidago on the nutrient-rich substrate. A conceptual model of the relative importance of root competition for soil nutrients, shoot competition for light, and positive interactions along the fertility gradient is presented. The model emphasizes that positive interactions play an important role over a broad range of the productivity scale with a peak at intermediate levels of fertility. On the substrate of high productivity shoot competition for light is more important than positive interactions and root competition for soil nutrients as well. The competitive superiority of Calamagrostis on the most productive substrate was evident only in the long run. Rare events like extreme summer drought or selective herbivore pressure caused a switch in dominance in mixtures with Solidago, respectively Tanacetum. The guerrilla growth strategy of Calamagrostis and interference competition through a dense cover of aboveground biomass and litter could further cause competitive exclusion.

Topographic influences on radial growth of Scots pine (Pinus sylvestris L.) at small spatial scales.

Abstract: Dendroecological and numerical methods were used to study the influence of topographic position on radial growth of Scots pine (Pinus sylvestris) stands exposed to soil dryness. The correlation structure of total tree-ring width and latewood width of eight scattered populations representing various topographic habitats (steep south-facing slopes, plateaus and hollows) within a rock-slide area (750 m a.s.l.) of about 1 km2 was investigated by principal component analysis. Scatter plots of component loadings indicated that (i) total ring width and latewood width are influenced by various climatic factors, (ii) stands growing at similar topographic position show a high agreement in year-to-year variability of radial growth, and (iii) distinct effects of topographic features (slope aspect, slope magnitude) on tree growth are modified by local disturbances (erosion, grazing) and the age structure of stands. Furthermore, both the time series of component scores and non-metric multidimensional scaling of chronologies indicated years where extremely limiting or favorable climate conditions prevailed throughout the study area (pointer years). The influence of climate on tree growth in various topographic habitats was mediated through the influence of climatically stressful years. Because stands are located at sites with different levels of water stress, growth differences between chronologies are considered to be caused by site-specific susceptibility of tree growth to soil dryness. Significant correlations between precipitation in April to June and ring-width confirm that water availability is the primary growth-limiting factor within the study area. These small-scale variations in growth-climate relationships have significant implications for dendroclimatological studies. So paleoclimatic reconstructions based on tree rings will have to assure that an unbiased data set is used, which compensates for local growth-variabilities due to site related environmental stresses.

Patterns of abundance of a narrow endemic species in a tropical and infertile montane habitat

Abstract: We examined the entire spatial distribution of a narrow endemic shrub (Coccoloba cereifera, Polygonaceae) in Serra do Cipo, Brazil. We tested the hypothesis that a narrow endemic species would show a gradual decline in either size and density towards the edges of its distribution. The contribution of soil specificity and post-fire growth to C. cereifera abundance and distribution were also investigated. C. cereifera showed multimodal and highly aggregated distribution pattern at several scales, from 25 m2 to 3000 m2 (blocked quadrat variance analyses). This pattern seems to be strongly related to the predominance of clonal recruitment and to the close association of the species to sandfields, which have discrete distribution between gallery forests and rocky outcrops. Population density did not decline towards the edge of the species distribution. Plants near the distribution boundaries had slightly more leaves and more inflorescences per plant (p<0.005), but there was no significant change in the mean number of ramets per clone. The absence of large plants in some populations at the center of the species distribution may be related to the higher frequency of fire in this region, killing aerial plant parts. Nearly all aggregations had inverse-J shaped size-distribution, suggesting effective recruitment of ramets, most frequently via asexual reproduction. Similar patterns of plant abundance may be common in fire-prone habitats characterized by infertile, and well-drained soils since these areas generally have high numbers of endemic plants, with strong soil specificity. Possible mechanisms for the observed pattern are discussed considering current models concerning distribution of abundance of species.