The influence of Prosopis canopies on understorey vegetation: Effects of landscape position

Abstract. The influence of canopy trees and shrubs on under‐storey plants is complex and context‐dependent. Canopy plants can exert positive, negative or neutral effects on production, composition and diversity of understorey plant communities, depending on local environmental conditions and position in the landscape. We studied the influence of Prosopis velutina (mesquite) on soil moisture and nitrogen availability, and understorey vegetation along a topographic gradient in the Sonoran Desert. We found significant increases in both soil moisture and N along the gradient from desert to riparian zone. In addition, P. velutina canopies had positive effects, relative to open areas, on soil moisture in the desert, and soil N in both desert and intermediate terrace. Biomass of understorey vegetation was highest and species richness was lowest in the riparian zone. Canopies had a positive effect on biomass in both desert and terrace, and a negative effect on species richness in the terrace. The effect of the canopy depended on landscape position, with desert canopies more strongly influencing soil moisture and biomass and terrace canopies more strongly influencing soil N and species richness. Individual species distributions suggested interspecific variation in response to water‐ vs. N‐availability; they strongly influence species composition at both patch and landscape position levels.     

Impact of shrub canopies on understorey vegetation in western Eurasian tundra

Question: How does the composition and species richness of understorey vegetation associate with changing abundance of deciduous shrub canopies? What are the species‐specific associations between shrubs and understorey plants? Location: Tundra habitats along an over 1000‐km long range, spanning from NW Fennoscandia to the Yamal Peninsula in northwest Russia. Methods: The data from 758 vegetation sample plots from 12 sites comprised cover estimates of all plant species, including bryophytes and lichens, and canopy height of deciduous shrubs. The relationships between shrub volume and cover of plant groups and species richness of vegetation were investigated. In addition, species‐specific associations between understorey species and shrub volume were analysed. Results: Shrub abundance was shown to be associated with the composition of understorey vegetation, and the association patterns were consistent across the study sites. Increased forb cover was positively associated with shrub volume, whereas bryophyte, lichen, dwarf shrub and graminoid cover decreased in association with increasing volume of deciduous shrubs. The total species richness of vegetation declined with increasing shrub volume. Conclusions: The results suggest that an increase of shrubs – due to climatic warming or a decrease in grazing pressure – is likely to have strong effects on plant–plant interactions and lead to a decrease in the diversity of understorey vegetation.     

Tree-grass co-existence in savanna: Interactions of rain and fire

The mechanisms permitting the co-existence of tree and grass in savannas have been a source of contention for many years. The two main classes of explanations involve either competition for resources, or differential sensitivity to disturbances. Published models focus principally on one or the other of these mechanisms. Here we introduce a simple ecohydrologic model of savanna vegetation involving both competition for water, and differential sensitivity of trees and grasses to fire disturbances. We show how the co-existence of trees and grasses in savannas can be simultaneously controlled by rainfall and fire, and how the relative importance of the two factors distinguishes between dry and moist savannas. The stability map allows to predict the changes in vegetation structure along gradients of rainfall and fire disturbances realistically, and to clarify the distinction between climate- and disturbance-dependent ecosystems.     

Structure of the woody vegetation in disturbed and undisturbed arid savanna,Botswana

The structure of woody vegetation was studied in little disturbed arid savanna and in adjacent over-grazed vegetation. In the over-grazed areas density and cover of woody plants were higher than in the less disturbed vegetation. The difference was accounted for by one species, Acacia mellifera, which was strongly dominant in the overgrazed vegetation. In the open savanna, the woody species varied in height from small shrubs to trees, while the dense shrub vegetation was of uniformly low stature.It is suggested that, while the differences in total abundance of woody species depend on differences in the amount of soil water available for woody growth, differences in species composition and height distribution are governed by the spatial and temporal distribution of water in the soil profile.     

The influence of soil texture on the soil water dynamics and vegetation structure of a shortgrass steppe ecosystem

We analyzed soil water data from three sites with different soil textures in the shortgrass steppe of northeastern Colorado, USA. Our objective was to evaluate the relationship between the occurrence of plant functional types and the effect of soil texture on soil water availability. Soil water availability was greatest in the upper soil layers at all three sites, but the loamy sand site had significantly greater soil water availability than the sandy clay loam and sandy clay sites in wetter years at depths below 60 cm. Calculations of proportional water availability by layer using both field data and fifty-year soil water model simulations, showed that the sandy clay loam and sandy clay soils on average had greater water availability in layers 30 cm and above, but that the loamy sand had the greatest water availability in layers beneath this, particularly at 105 cm. This observation can be linked to the occurrence of a fine textured subsoil at this site. The textural pattern in the loamy sand profile effectively creates two water resources: a shallow pool accessible to all plants; and a deep pool accessible only to deep-rooted plants. This is offered as an explanation for the co-dominance of the two main plant functional types at the loamy sand site. At the other two sites, shallow-rooted shortgrass vegetation dominated, being more consistent with the general pattern for the area. Thus the patterns of vegetation structure at the three sites were consistent with the hypothesis. Aboveground net primary production data for the three sites, along with transpiration estimates from the model simulations, indicated that the additional water availability in the coarse textured soil was associated with higher overall plant productivity.Nomenclature: Taxonomic nomenclature follows R. L. McGregor & T. M. Barkley (1986) Flora of the Great Plains. Great Plains Flora Association. University Press of Kansas, Lawrence.     

Nutrient availability constrains the hydraulic architecture and water relations of savannah trees

Leaf and whole plant-level functional traits were studied in five dominant woody savannah species from Central Brazil (Cerrado) to determine whether reduction of nutrient limitations in oligotrophic Cerrado soils affects carbon allocation, water relations and hydraulic architecture. Four treatments were used: control, N additions, P additions and N plus P additions. Fertilizers were applied twice yearly, from October 1998 to March 2004. Sixty-three months after the first nutrient addition, the total leaf area increment was significantly greater across all species in the N- and the N + P-fertilized plots than in the control and in the P-fertilized plots. Nitrogen fertilization significantly altered several components of hydraulic architecture: specific conductivity of terminal stems increased with N additions, whereas leaf-specific conductivity and wood density decreased in most cases. Average daily sap flow per individual was consistently higher with N and N + P additions compared to the control, but its relative increase was not as great as that of leaf area. Long-term additions of N and N + P caused midday PsiL to decline significantly by a mean of 0.6 MPa across all species because N-induced relative reductions in soil-to-leaf hydraulic conductance were greater than those of stomatal conductance and transpiration on a leaf area basis. Phosphorus-fertilized trees did not exhibit significant changes in midday PsiL. Analysis of xylem vulnerability curves indicated that N-fertilized trees were significantly less vulnerable to embolism than trees in control and P-fertilized plots. Thus, N-induced decreases in midday PsiL appeared to be almost entirely compensated by increases in resistance to embolism. Leaf tissue water relations characteristics also changed as a result of N-induced declines in minimum PsiL: osmotic potential at full turgor decreased and symplastic solute content on a dry matter basis increased linearly with declining midday PsiL across species and treatments. Despite being adapted to chronic nutrient limitations, Cerrado woody species apparently have the capacity to exploit increases in nutrient availability by allocating resources to maximize carbon gain and enhance growth. The cost of increased allocation to leaf area relative to water transport capacity involved increased total water loss per plant and a decrease in minimum leaf water potentials. However, the risk of increased embolism and turgor loss was relatively low as xylem vulnerability to embolism and leaf osmotic characteristics changed in parallel with changes in plant water status induced by N fertilization.     

Effects of nutrient availability and neighbours on shoot growth,resprouting and flowering of Erica multiflora

Abstract. To test if low soil fertility and competition limit the performance of Mediterranean shrubs, and if the effects of competition on plant performance were modified by soil fertility, we subjected shrubs of Erica multiflora to a factorial field experiment of fertilization and removal of neighbours around target plants. After 18 months of treatment, fertilization had stimulated the growth of pre-existent sprouts and biomass allocation to stems into new sprouts, but decreased the frequency of sprout flowering. Removal of neighbours increased the number and biomass of new sprouts, the probability of sprout flowering and the biomass of flowers. Fertilization slightly enhanced sprout recruitment and the probability of sprout flowering when neighbours were removed, but did not modify the other parameters of plant performance. According to our results, both low soil fertility and competition limited plant performance. Competition was slightly more intense in fertilized plants, but only in determining sprout and flowering bud stimulation.     

Spatial partitioning of the soil water resource between grass and shrub components in a West African humid savanna

Most savanna water balance models assume water partitioning between grasses and shrubs in a two-layer hypothesis, but this hypothesis has not been tested for humid savanna environments. Spatial partitioning of soil water between grasses and shrubs was investigated in a West African humid savanna by comparing the isotopic composition (oxygen-18 and deuterium) of soil water and plant stem water during rainy and dry conditions. Both grass and shrub species acquire most of their water from the top soil layer during both rainy and dry periods. A shift of water uptake pattern towards deeper horizons was observed only at the end of the dry season after shrub defoliation. The mean depth of water uptake, as determined by the isotopic signature of stem water, was consistent with grass and shrub root profiles and with changes in soil water content profiles as surveyed by a neutron probe. This provides evidence for potentially strong competition between shrubs and grasses for soil water in these humid savannas. Limited nutrient availability may explain these competitive interactions. These results enhance our understanding of shrub-grass interactions, and will contribute to models of ecosystem functioning in humid savannas.     

The effect of increased nutrient availability on vegetation dynamics in wet heathlands

A three year fertilization experiment was conducted in which nitrogen (N series: 20 g N m^–2 yr^–1), phosphorus (P series: 4 g P m^–2 yr^–1) and potassium (K series: 20 g K m^–2 yr^–1) were added to a mixed vegetation of Erica tetralix and Molinia caerulea. At the end of each growing season the percentage cover of each species was determined. At the end of the experiment percentage cover of each species was found to be positively correlated with the harvested biomass. In the unfertilized control series the cover of Erica and Molinia did not change significantly during the experiment. In all fertilized series however, especially in the P series, cover of Erica decreased significantly. The cover of Molinia increased significantly in the P series only.In the fertilized series the biomass of Erica and total biomass per plot did not change significantly compared with the control series. In the P series the biomass of Molinia increased significantly.It is suggested that with increasing phosphorus or nitrogen availability Molinia outcompetes Erica because the former invests more biomass in leaves which in turn permits more carbon to be allocated to the root system, which thereupon leads to a higher nutrient uptake.     

Tree proximity,soil pathways and common mycorrhizal networks: their influence on the utilization of redistributed water by understory seedlings

Hydraulic redistribution (HR) is a process by which water moves through plant roots from moist to dry soils. An experiment was conducted to quantify the influence of common mycorrhizal networks (CMNs) and proximity to mature HR-source trees on the water relations of surrounding seedlings. Douglas-fir (Pseudotsuga menziesii var glauca (Mirb.) Franco) seedlings were planted at four distances (0.5, 1, 2.5, and 5 m) from six mature Douglas-fir trees, either directly into soil (soil plus CMN pathway) or inside 0.5 μm mesh bags (soil-only pathway). Deuterated water was used to irrigate soil beside mature trees in order to identify different HR water pathways to surrounding seedlings. This was followed by measurements of seedling deuterium enrichment, seedling water potential, soil water potential_, gravimetric soil water content, and tree root density surrounding the seedlings. There was no significantly detectable difference in the quantity of HR water transferred to seedlings having access to soil and CMN pathways or soil-only pathways of water movement. Water from the irrigation plot contributed up to 1.4% of the water of Douglas-fir seedlings. Based on the assumption that the only pathway through which seedlings could access irrigation water was through the mature trees, we estimate that as much as 21.6% of the seedling water was supplied by the nearby tree. Seedling water potential was not significantly affected either by proximity to mature trees or pathway, suggesting HR may have compensated for increasing tree competitive effects with proximity. It is also possible that the lack of difference was due to a relatively moist summer. Our results suggest that residual mature trees are potentially important for hydraulic redistribution to regenerating seedlings in harvested dry interior Douglas-fir forests.     

Effects of a nutrient pulse supply on nutrient status of the Mediterranean trees Quercus ilex subsp. ballota and Pinus halepensis on different soils and under different competitive pressure

Nutrient availability is a key factor in Mediterranean ecosystems that affects the primary productivity and the community structure. The great variability of its natural availability is now increasing due to frequent fires, pollution events and changes in rainfall regime associated to climate change. Quercus ilex ssp. ballota and Pinus halepensis are the most abundant tree species in the NW Mediterranean basin. They frequently compete in the early and middle successional stages. We investigated the effects of N and P pulse supplies on nutrient uptake capacity in these two species in an after-fire field area and in nursery conditions on different soil types and competing conditions. In the field, N fertilisation had weak effects on nutrient concentration and mineralomass likely as a consequence of this nutrient not being limiting in this field site whereas P fertilisation increased the P mineralomass and the Mg, S, Fe, K and Ca concentrations and mineralomass in the different biomass fractions of both species 1 and 3 years after fertilisation application. In the nursery experiments, P fertilisation increased the mineralomass and concentrations of P, Mg, S, Fe, K and Ca in all biomass fractions including the roots in both species and in different soils and competition conditions. The increment of nutrient mineralomass was due to both the increase of growth and of nutrient concentrations. Both species were able to absorb significant amounts of the P applied by fertilisation (between 5 and 20%) in short time (18 months). Competing vegetation decreased the positive effects of P fertilisation, and in many cases the negative effect of competing vegetation on nutrient mineralomass was stronger when P availability was increased by fertilisation. Q. ilex subsp ballota showed a greater competitive ability for P than the more pioneer species Pinus halepensis in the field but not in the nursery conditions. Pinus halepensis had greater nutrient mineralomass in calcareous than in siliceous soils. Q. ilex subsp. ballota had a higher root biomass allocation and root nutrient allocation than P. halepensis, but both species showed a high capacity to increase their nutrient uptake when its availability increased by fertilisation, thus assuring a great nutrient reserve for future growth periods and contributing to retain nutrients in the ecosystem.     

Drought avoidance and the effect of local topography on trees in the understorey of Bornean lowland rain forest

The water relations of two tree species in the Euphorbiaceae werecompared to test in part a hypothesis that the forest understorey plays anintegral role in drought response. At Danum, Sabah, the relatively commonspecies Dimorphocalyx muricatus is associated with ridgeswhilst another species, Mallotus wrayi, occurs widely bothon ridges and lower slopes. Sets of subplots within two 4 -hapermanent plots in this lowland dipterocarp rain forest, were positioned onridges and lower slopes. Soil water potentials were recorded in1995–1997,and leaf water potentials were measured on six occasions. Soil water potentialson the ridges (–0.047 MPa) were significantly lower than onthe lower slopes (–0.012 MPa), but during the driest periodin May 1997 they fell to similarly low levels on both sites (–0.53MPa). A weighted 40-day accumulated rainfall index was developedtomodel the soil water potentials. At dry times, D.muricatus(ridge) had significantly higher pre-dawn (–0.21 v.–0.57 MPa) and mid-day (–0.59 v.–1.77 MPa) leaf water potentials than M.wrayi (mean of ridge and lower slope). Leaf osmotic potentials ofM. wrayi on the ridges were lower (–1.63MPa) than on lower slopes (–1.09 MPa), withthose for D. muricatus being intermediate (–1.29MPa): both species adjusted osmotically between wet and dry times.D. muricatus trees were more deeply rooted thanM. wrayi trees (97 v. 70cm). M. wrayi trees had greaterlateral root cross-sectional areas than D. muricatus treesalthough a greater proportion of this sectional area for D.muricatus was further down the soil profile. D.muricatus appeared to maintain relatively high water potentialsduring dry periods because of its access to deeper water supplies and thus itlargely avoided drought effects, but M. wrayi seemed to bemore affected yet tolerant of drought and was more plastic in its response. Theinteraction between water availability and topography determines these species'distributions and provides insights into how rain forests can withstandoccasional strong droughts.     

The role of light availability and herbivory on algal responses to nutrient enrichment in a riparian wetland,Alaska

We investigated how the relative availability of solar radiation in the presence or absence of grazing alters the ability of benthic algae to respond to nutrient enrichment in an Alaskan marsh. We used a factorial mesocosm experiment that included nutrient enrichment (enriched or control), grazing (grazed or ungrazed), and light (unshaded or shaded) to simulate shading by macrophytes early and late in the growing season, respectively. We found stronger effects of grazers and nutrients compared to light on benthic algal biomass and taxonomic composition. Algal biomass increased in nutrient‐enriched treatments and was reduced by grazing. Shading did not have an effect on algal biomass or taxonomic composition, but the concentration of chl a per algal biovolume increased with shading, demonstrating the ability of algae to compensate for changes in light availability. Algal taxonomic composition was more affected by grazer presence than nutrients or light. Grazer‐resistant taxa (basal filaments of Stigeoclonium) were replaced by diatoms (Nitzschia) and filamentous green algae (Ulothrix) when herbivores were removed. The interacting and opposing influences of nutrients and grazing indicate that the algal community is under dual control from the bottom‐up (nutrient limitation) and from the top‐down (consumption by herbivores), although grazers had a stronger influence on algal biomass and taxonomic composition than nutrient enrichment. Our results suggest that low light availability will not inhibit the algal response to elevated nutrient concentrations expected with ongoing climate change, but grazers rapidly consume algae following enrichment, masking the effects of elevated nutrients on algal production.     

Facilitation, competition, and vegetation patchiness: from scale free distribution to patterns

A new technique for the modeling of perennial vegetation patchiness in the arid/semiarid climatic zone is suggested. Incorporating the stochasticity that affects life history of seedlings and the deterministic dynamics of soil moisture and biomass, this model is flexible enough to yield qualitatively different forms of spatial organization. In the facilitation-dominated regime, scale free distribution of patch sizes is observed, in correspondence with recent field studies. In the competition controlled case, on the other hand, power-law statistics is valid up to a cutoff, and an intrinsic length scale appears.     

The influence of light and nutrient addition upon the sediment chemistry of iron in an arctic lake

The geochemical response of sediments to increased nutrient input to an Alaskan, arctic lake was examined using direct measurements of sediment-water chemical fluxes. An unexpected increase in Fe flux occurred when sediments were exposed to high incident radiation and nutrient concentrations. Correlation between light and acid-soluble Fe concentrations suggests that photoreduction of Fe(III) oxides may occur, but nutrient addition enhanced the effect indicating that primary productivity was also important. The processes controlling the flux of Fe from sediments in this lake were complex and included the redox potential (dissolved oxygen concentration) of the water, quality of organic matter present in the sediment, light, and nutrients supplied from the sediments and/or water column. These four factors together with the possibility of direct uptake of Fe by phytoplankton and the possible release of algal reductants may contribute to Fe cycling in this lake.     

Resource limitation,habitat segregation,and species interactions of british tree-hole mosquitoes in nature

Summary The insect fauna of water-filled tree holes in southern Britain consists primarily of the mosquitoes Aedes geniculatus, Anopheles plumbeus, Culex torrentium, and a benthic detritivorous fauna that includes primarily the scirtid beetle Prionocyphon serricornis and the chironomid midge Metriocnemus martinii. Culex torrentium has been documented only relatively recently in tree holes but all three species of mosquitoes partition the resource in space and time. When mosquito larvae were forced to coexist in natural tree holes at limiting densities and at higher than natural levels of interspecific encounter, there was no evidence that Aedes geniculatus or Anopheles plumbeus affected pupation success, pupal weight, or development time of the other or that either Aedes geniculatus or C. torrentium affected the survivorship, pupation success, pupal weight, and biomass yield of the other. When A. geniculatus at limiting densities were forced in natural tree holes to live without or to coexist with natural or twice natural densities of P. serricornis and M. martinii, the presence, absence, or superabundance of the benthic insects did not affect pupation success or pupal weight of A. geniculatus; development time of A. geniculatus was faster when a superabundance of the benthic fauna was present. Effects of the benthic fauna on A. geniculatus are slight and the only significant interaction is facilitative, not competitive. The pattern of habitat segregation among treehole mosquitoes in southern Britain is characteristic of their respective genera and we propose that this pattern is more likely (but not certain) to have arisen through a process of independent evolution than through competitively driven niche shifts among already coexisting species.     

Effects of light and soil water availability on leaf photosynthesis and growth of Arisaema heterophyllum,a riparian forest understorey plant

The effects of soil-water availability on leaf light acclimation and whole-plant carbon gain were examined in Arisaema heterophyllum Blume, a riparian deciduous forest understorey plant. Photosynthesis, above-ground morphology and ramet biomass accumulation (relative growth rate: RGR of a corm for a full leaf life-span) were measured on plants raised under three light treatments combined with two soil water conditions. The two higher light treatments during growth (high: max. 550 μmol photons m^–2 s^–1; medium: 150 μmol photons m^–2 s^–1) resulted in a twofold increase in RGRs, 30% higher photosynthetic capacities and 20% less photosynthetic low-light use efficiency than those under a low light condition (50 μmol photons m^–2 s^–1). Leaf area was the smallest and leaf mass area ratio was the largest under the high light treatment. Water stress decreased both photosynthetic rate and leaf area and, hence, RGR in all the light regimes. However, water stress did not alter the general patterns of physiological and morphological responses to different light regimes. We estimated that higher photosynthetic low-light use efficiency and larger leaf area in the low light leaf would lead to a threefold carbon gain as compared with the high light leaf under simulated low light conditions. Both experimental and simulation results suggest that the physiological and morphological acclimations tend to be beneficial to carbon gain when light availability is low, whereas they favor increased water use efficiency when light availability is sufficiently high. Electronic Publication     

Soil type, microsite, and herbivory influence growth and survival of Schinus molle (Peruvian pepper tree) invading semi-arid African savanna

Naturalization of Schinus molle (Anacardiaceae) has been observed in semi arid savanna of the Northern Cape Province of South Africa. However, with high dispersal ability, the species is expected to achieve greater densities and invade more widely. The study involved a field manipulation experiment over 14 months using a factorial block design to examine transplanted seedlings in different savanna environments. The experiments examine the effects of soil type (sandy and clay), microsite, and herbivores on seedling performance (establishment, growth and survival). Seedlings were grown in a greenhouse and individually transplanted into four treatment groups: in open grassland, under tree canopies, and with and without cages to exclude large herbivores (cattle and game). The same experiment was repeated in two different soil types: coarse sand and fine-textured clay soil. Results suggest that protection provided by canopies of large indigenous Acacia trees facilitates S. molle invasion into semi-arid savanna. In the field, S. molle seedlings performed considerably better beneath canopies of indigenous Acacia trees than in open areas regardless of soil type. Whether exposed or protected from large herbivores, no seedlings planted in open grassland survived the first winter. Although, seedlings grew better and had higher survival rates beneath tree canopies than in the open sites, exposure to large herbivores significantly decreased heights and canopy areas of seedlings compared with those protected from large herbivores. The effect was greater on clay soil than on sandy soil. The results suggest that low temperature (frost), and possibly inter-specific competition with grasses, may limit S. molle seedling establishment, survival and growth away from tree canopies in semi arid savannas. Low soil nutrient status and browsing may also delay growth and development of this species. The invasive potential of S. molle is thus greatest on fertile soils where sub-canopy microsites are present and browsing mammals are absent.