EFFECTS OF CLIPPING AND FOUR LEVELS OF NITROGEN ON THE GAS EXCHANGE,GROWTH, AND PRODUCTION OF TWO EAST AFRICAN GRAMINOIDS

Kyllinga nervosa and Themeda triandra plants were subjected to different clipping and nitrogen availability regimes. Following an extended period of growth under these conditions, total biomass, gas exchange and several morphological parameters were measured. Kyllinga nervosa showed compensatory growth to moderate levels of clipping whereas any clipping reduced the total biomass of T. triandra. Unclipped plants of either species were unable to respond to increased levels of nitrogen. Clipped plants responded in an ambiguous fashion, with increased allocation to offtake (material removed by clipping) in both species. Total biomass of K. nervosa was highest at 15 mM nitrogen levels which are equivalent to field levels. Both photosynthesis and respiration rates were unaffected by nitrogen treatments. Photosynthesis was significantly reduced by the most severe clipping regime of K. nervosa, but was unaffected by clipping of T. triandra.     

Tradeoff between height and relative growth rate in a dominant grass from the Serengeti ecosystem

We determined the relationship between plant height and whole-plant relative growth rate (g g^-1 day^-1) for ten genotypes of Sporobolus kentrophyllus collected from an intensively grazed site on the Serengeti Plains, Tanzania. Plants were grown for 7 weeks in a greenhouse in Syracuse, N.Y., and harvested weekly. Plants that received simulated bovine urine showed a negative relationship between plant height and growth rate, suggesting a genetic tradeoff between competitive ability if ungrazed (height) and ability to recover from grazing (growth rate). There was no height-growth rate relationship under nitrogen addition rates similar to field mineralization rates. In addition, faster-growing, shorter plants tended to have relatively higher above-ground growth rates than slower-growing, taller plants. These results suggest that natural selection has maintained a gradient of morphologies within this species ranging from short, rapidly growing genotypes adapted to intense grazing conditions to tall, slow-growing, grazer-susceptible genotypes that are superior light competitors in absence of herbivory.     

Urea as a promotive coupler of plant-herbivore interactions

Summary Growth responses of Kyllinga nervosa Steud., a sedge from the Serengeti short-grass plains, were examined in a factorial experiment which included clipped and unclipped plants, and nitrogen supplied as either urea or ammonium nitrate. Results were expressed in relation to three transfer processes: flow to grazers, flow to producers and flow to reproduction. Clipping increased biomass and nitrogen flow to grazers by significantly increasing nitrogen uptake, aboveground nitrogen flow, and the weights of and proportional allocation to green leaf production. This was at the expense of flow to vegetative and sexual reproduction, since the weights and proportional investments in roots, crowns and reproductive structures were reduced. Urea nutrition increased flow to grazers and plant reproduction through increases in green leaf weight, flower weight, allocation to green leaves, flowers and stems, and aboveground: belowground biomass ratios. Stimulation of aboveground productivity by urea was a consequence of increased tillering rates.Interactive responses of clipping and nitrogen source regulated plant growth, thus controlling flow to each transfer process. Combined effects of clipping and urea resulted in compensatory production of both green leaves and flowers, and maximized biomass and nitrogen flow to grazers. Both urea and clipping tightened herbivore-producer recycling by significantly reducing litter nitrogen and carbon masses. In contrast, when plants were unclipped and grown on NH₄NO₃, biomass allocation and weights of roots and crowns were increased at the expense of aboveground tissues, thus increasing flow to primary producers. Plant growth responses to experimental treatment combinations simulating nutritional status of grazed and ungrazed field plants indicate that urea represents a potential importance beyond it nitrogen contribution by introducing a positive feedback to herbivores.     

Fuzzy set ordination and classification of Serengeti short grasslands,Tanzania

Abstract. Species abundance data from 32 representative stands are used to evaluate the vegetation in the heavily grazed Serengeti short grasslands. The abundant species are: Kyllinga nervosa, Sporobolus ioclados, S. kentro-phyllus and Eustachys paspaloides, all occurring in elevated areas, and Hypoestes forskalei, which is found in the drainage areas. Numerical classification and fuzzy set ordination revealed four main grassland communities associated with a topographic gradient. The communities are considered distinct since their niches, as computed in the community and environmental fuzzy system space are not overlapping. Leaching, erosion and animal disturbance effects correlated with the topographic gradient are found to determine species composition and overall community structure in the grasslands.     

The interaction of defoliation and nutrient uptake in Sporobolus kentrophyllus,a short-grass species from the serengeti plains

Summary Sporobolus kentrophyllus, a grazing-tolerant C₄ grass from the southeastern Serengeti Plains, was grown in solution culture to examine the effects of clipping on the uptake, preference and subsequent transport of varying nitrogen forms. Clipping reduced offtake mass, crown mass ane root mass, resulting in a 58% decline in plant mass. Proportional biomass allocation to roots decreased with clipping, while tillering rates increased. Clipping also increased the nitrogen concentrations of all tissues, and plant nitrogen uptake (nitrogen accumulated throughout the experiment per gram root). The ¹⁵N concentrations (% atom excess) of all tissues were higher in clipped compared with unclipped plants, and the average ¹⁵N uptake rate of clipped plants was twice that of unclipped plants. The relative ¹⁵N allocation to aboveground mass, a measure of canopy sink strength, was higher in clipped plants. Plants fed ¹⁵N-ammonium or ¹⁵N-nitrate during the ¹⁵N pulse experiment had greater ¹⁵N tissue concentrations compared with urea-fed plants, and ¹⁵N uptake rates were higher in ammonium-fed and nitrate-fed plants, compared with urea-fed plants. The relative magnitudes of these differences were higher when plants were clipped. Clipped plants had higher uptake rates for potassium, phosphorus and sodium, while differences between clipping treatments for calcium, iron, and magnesium were indistinguishable. Rapid uptake rates for species on the southeastern Serengeti plains, particularly during grazing periods, have important implications for nutrient cycling in this system.     

Interspecific competition among grasshoppers and their effect on plant abundance in experimental field environments

Summary We tested whether grasshoppers in experimental field environments, i.e. cages (40×40 cm) placed on existing old field vegetation, (1) were limited in density by plant abundance and/or nitrogen content, (2) exhibited interspecific competition, and (3) altered the relative abundance of different plant species. We examined interactions among a pair of early season grasshopper species (May–June; Arphia conspersa and Pardalophora apiculata) and a late season pair (July–August; Melanoplus femur-rubrum and Melanoplus bivittatus). Each grasshopper species was placed in cages by itself and with another grasshopper species. Grasshoppers generally survived at higher density in fertilized cages and they reduced plant abundance relative to empty cages, suggesting that grasshoppers may be food limited at these densities. In unfertilized plots, early season grasshoppers preferred grasses (Schizachyrium scoparium and Poa pratensis) and favored the growth of forbs (especially Solidago spp.). However, late in summer, Melanoplus spp. preferred Solidago spp. and favored the growth of grasses.The pattern of grasshopper survivorship and plant reduction within these experimental environments provide preliminary support for some of the predictions of resource competition theory. Grasshoppers exhibited interspecific competition only if they significantly reduced plant biomass. If two species competed, a grasshopper species was eliminated only if the superior competitor, when living by itself, could reduce plant biomass to a significantly lower level than the inferior competitor. Competitors persisted only if they did not differ in their ability to reduce plant biomass or reduced the abundance of different plant species.     

Biomass and mineral element responses of a Serengeti short-grass species to nitrogen supply and defoliation: compensation requires a critical [N]

Large mammalian herbivores in grassland ecosystems influence plant growth dynamics in many ways, including the removal of plant biomass and the return of nutrients to the soil. A 10-week growth chamber experiment examined the responses of Sporobolus kentrophyllus from the heavily grazed short-grass plains of Serengeti National Park, Tanzania, to simulated grazing and varying nitrogen nutrition. Plants were subjected to two clipping treatments (clipped and unclipped) and five nitrogen levels (weekly applications at levels equivalent to 0, 1, 5, 10, and 40 g N m⁻²), the highest being equivalent to a urine hit. Tiller and stolon production were measured weekly. Total biomass at harvest was partitioned by plant organ and analyzed for nitrogen and mineral element composition. Tiller and stolon production reached a peak at 3–5 weeks in unclipped plants, then declined drastically, but tiller number increased continually in clipped plants; this differential effect was enhanced at higher N levels. Total plant production increased substantially with N supply, was dominated by aboveground production, and was similar in clipped and unclipped plants, except at high nitrogen levels where clipped plants produced more. Much of the standing biomass of unclipped plants was standing dead and stem; most of the standing biomass of clipped plants was live leaf with clipped plants having significantly more leaf than unclipped plants. However, leaf nitrogen was stimulated by clipping only in plants receiving levels of N application above 1 g N m⁻² which corresponded to a tissue concentration of 2.5% N. Leaf N concentration was lower in unclipped plants and increased with level of N. Aboveground N and mineral concentrations were consistently greater than belowground levels and while clipping commonly promoted aboveground concentrations, it generally diminished those belowground. In general, clipped plants exhibited increased leaf elemental concentrations of K, P, and Mg. Concentrations of B, Ca, K, Mg, and Zn increased with the level of N. No evidence was found that the much greater growth associated with higher N levels diminished the concentration of any other nutrient and that clipping coupled with N fertilization increased the total mineral content available in leaf tissue. The results suggest that plants can (1) compensate for leaf removal, but only when N is above a critical point (tissue [N] 2.8%) and (2) grazing coupled with N fertilization can increase the quality and quantity of tissue available for herbivore removal. Received: 25 August 1997 / Accepted: 14 April 1998     

The effects of clipping and soil moisture on leaf and root morphology and root respiration in two temperate and two tropical grasses

Numerous studies have explored the effect of environmental conditions on a number of plant physiological and structural traits, such as photosynthetic rate, shoot versus root biomass allocation, and leaf and root morphology. In contrast, there have been a few investigations of how those conditions may influence root respiration, even though this flux can represent a major component of carbon (C) pathway in plants. In this study, we examined the response of mass-specific root respiration (μmol CO₂ g⁻¹ s⁻¹), shoot and root biomass, and leaf photosynthesis to clipping and variable soil moisture in two C₃ (Festuca idahoensis Elmer., Poa pratensis L.) and two C₄ (Andropogon greenwayi Napper, and Sporobolus kentrophyllus K. Schum.) grass species. The C₃ and C₄ grasses were collected in Yellowstone National Park, USA and the Serengeti ecosystem, Africa, respectively, where they evolved under temporally variable soil moisture conditions and were exposed to frequent, often intense grazing. We also measured the influence of clipping and soil moisture on specific leaf area (SLA), a trait associated with moisture conservation, and specific root length (SRL), a trait associated with efficiency per unit mass of soil resource uptake. Clipping did not influence any plant trait, with the exception that it reduced the root to shoot ratio (R:S) and increased SRL in P. pratensis. In contrast to the null effect of clipping on specific root respiration, reduced soil moisture lowered specific root respiration in all four species. In addition, species differed in how leaf and root structural traits responded to lower available soil moisture. P. pratensis and A. greenwayi increased SLA, by 23% and 33%, respectively, and did not alter SRL. Conversely, S. kentrophyllus increased SRL by 42% and did not alter SLA. F. idahoensis responded to lower available soil moisture by increasing both SLA and SRL by 38% and 33%, respectively. These responses were species-specific strategies that did not coincide with photosynthetic pathway (C₃/C₄) or growth form. Thus, mass-specific root respiration responded uniformly among these four grass species to clipping (no effect) and increased soil moisture stress (decline), whereas the responses of other traits (i.e., R:S ratio, SLA, SRL) to the treatments, especially moisture availability, were species-specific. Consequently, the effects of either clipping or variation in soil moisture on the C budget of these four different grasses species were driven primarily by the plasticity of R:S ratios and the structural leaf and root traits of individual species, rather than variation in the response of mass-specific root respiration.     

The influence of nitrogen,water and competition on the vegetative and reproductive growth of common ragweed (<Emphasis Type="Italic">Ambrosia</Emphasis><Emphasis Type="Italic">artemisiifolia</Emphasis> L.)

As common ragweed (Ambrosia artemisiifolia L.) spreads across Europe and other regions, it is becoming both a health and an economic threat. To better understand which environmental conditions facilitate the spread of the invasive species, in 2010, a greenhouse experiment was conducted determining the effects of various nitrogen levels (10, 50 and 100 kg N/ha), soil moisture level (low and high) and competition levels (no competition, medium competition and high competition) on the growth parameters of ragweed. Single-grown ragweed responded favourably to the medium nitrogen and water increase, whereas the ragweed growth parameters in competition stands increased only when high levels of nitrogen and water were added. High competition reduced the total dry matter of ragweed by up to 83%, but the ragweed continued to increase its relative growth rate during the full-flowering stage and allocate its dry matter to reproductive parts, producing up to 70 seeds per plant. Ragweed is a poor competitor when there is high resource availability; however, under disturbance and in the shortage of nutrients and water conditions, the intensity of competition decreases and the ragweed performance is minimally affected. The addition of medium levels of nitrogen to promote the growth of competitive species, prevention of disturbance and establishment of plant communities with stress-tolerant species is measures that should help to prevent the further spread of ragweed.     

Sensitivity of the invasive geophyte Oxalis pes-caprae to nutrient availability and competition

BACKGROUND AND AIMS: Invasion by alien plants may be partially related to disturbance-related increases in nutrient availability and decreases of competition with native species, and to superior competitive ability of the invader. Oxalis pes-caprae is an invasive winter geophyte in the Mediterranean Islands that reproduces vegetatively via bulbs. An investigation was made into the relative responses of O. pes-caprae and the native annual grass Lolium rigidum to nutrient availability and to competition with each other in order to understand patterns of invasion in the field. Because Oxalis accumulates oxalic acid in its leaves, which could ameliorate soil phosphorous availability, field observations were made to determine whether the presence of Oxalis alters soil P availability. METHODS: A full-factorial glasshouse experiment was conducted with nutrient availability (high and low) and competition (Lolium alone, Oxalis alone, and Lolium and Oxalis together). Plant performance was assessed by determining (1) above- and below-ground biomass at the time of Oxalis maximum biomass and (2) reproductive output of Oxalis and Lolium at the end of their respective growth cycles. Measurements were also taken for leaf N and P content. Soil samples were taken in the field from paired Oxalis-invaded and non-invaded plots located in Menorca (Balearic Islands) and available P was determined. KEY RESULTS: High nutrient availability increased Oxalis and Lolium vegetative biomass and reproductive output to a similar degree. Competition with Lolium had a much stronger negative effect on Oxalis bulb production than reduced nutrients. Lolium was a superior competitor than Oxalis; the latter did not affect Lolium maximum biomass and spike production. Significantly greater soil-P availability in Oxalis-invaded field soils relative to paired non-invaded soils suggest that Oxalis influences soil P cycling. CONCLUSIONS: Oxalis is a poor competitor. This is consistent with the preferential distribution of Oxalis in disturbed areas such as ruderal habitats, and might explain its low influence on the cover of native species in invaded sites. The results also suggest that certain disturbances (e.g. autumn ploughing) may greatly enhance Oxalis invasion.     

Response of the invader Cortaderia selloana and two coexisting natives to competition and water stress

Alien species’ resistance and adjustment to water stress and plant competition might largely determine the success of invasions in Mediterranean ecosystems because water availability is often limiting biomass production. Two outdoor pot experiments were conducted to test the hypotheses that at the recruitment stage the invader perennial tussock grass Cortaderia selloana is a superior competitor, and that it is more resistant to water stress than the two coexisting native species of the same functional group, Festuca arundinacea and Brachypodium phoenicoides. C. selloana reduced aboveground biomass of target native species, but not more than target native species on each other. Moreover, C. selloana did not resist interspecific competition more than target native species. Under control conditions, C. selloana did not have larger specific leaf area (SLA) and root–shoot ratio (R/S) ratio than target native species, contradicting the general statement that these traits are associated to invasiveness. F. arundinacea was the species which performed best but also the one most affected by water stress. Both C. selloana and B. phoenicoides performed in a similar way under water stress conditions. However, the alien species’ capacity to adjust to water stress, indicated by the increase in the root–shoot ratio under moderate and severe water stress, was slightly better than that of B. phoenicoides. Overall, at early recruitment stages, C. selloana is not a better competitor than the coexisting native species. However, it seems to be more resistant to water stress because as water becomes scarce C. selloana maximizes water uptake and minimizes water losses more than the native species.     

Water relations in competitive interactions of Mediterranean grasses and shrubs

In Mediterranean ecosystems, competition between opportunistic grasses and slower-growing woody species may affect the speed and path of ecosystem recovery and the success of restoration plantings after natural or human-induced disturbance. In this experiment, competitive interactions between Mediterranean annual and perennial grass species (Avena fatua and Brachypodium retusum, respectively) and an important Mediterranean shrub (Rosmarinus offlcinalis) were examined under semi-controlled conditions simulating wet and dry Mediterranean rainfall regimes. The identity of the grass competitor and the level of water availability in the plots interacted to produce differing rates of R. offlcinalis growth but similar levels of mortality. In particular, competition with the perennial grass resulted in very low rates of R. offlcinalis growth at both irrigation levels. Measurements of soil water content showed that both grasses reduced soil moisture to low levels, though this effect was temporary in the case of the winter annual grass. Resistance to hydraulic flow in roots was highest in the perennial grass, smaller but of similar magnitude in the shrub, and much lower in the annual grass. Transpirational response to decreasing leaf water potential was a quick, sharp drop in conductance in R. offlcinalis, in contrast to a moderated decline from much lower initial transpiration rates in B. retusum. The annual grass largely maintained both leaf water potential and transpiration through leaf-tip senescence and death. Quantification of the rate of hydric recuperation of leaves after irrigation of drought-stressed plants showed that the perennial grass recovered at a rate four times that of R. offlcinalis, suggesting a strategy for making quick use of rare summer rains that may contribute to its competitive success. The appropriateness of planting or suppressing grasses in restoration of disturbed sites in Mediterranean Spain is discussed.     

不同水分条件下混种密度对芦苇和反枝苋种间竞争的影响

与本地植物的种间竞争是影响外来植物能否成功入侵的关键因素之一,该研究通过受控模拟试验研究了本地植物芦苇(Phragmites australis)和外来入侵植物反枝苋(Amaranthus retroflexus)在淹水和干旱两种水分条件下混种密度(6∶2、4∶4和2∶6)对其种间竞争的影响。结果表明:(1)芦苇和反枝苋的相对产量与相对产量总和均小于1,即两种植物存在种间竞争。(2)种间竞争使芦苇和反枝苋的生长均受到了不同程度的抑制,表现在两者的株高和生物量均随着竞争者密度的增加而降低。(3)植株地上部分和地下部分的氮浓度表现出与株高和生物量相同的趋势,且在不同水分条件下存在差异。(4)芦苇和反枝苋分别在淹水和干旱环境下具有较强竞争力,但在各自较高混种密度下亦具有较强竞争力。可见,芦苇和反枝苋的种间竞争受到了水分和混种密度的影响。因此,在有反枝苋分布的湿地中,植物生长初期可通过增加土壤水分和/或增加芦苇等本地植物的种群密度以降低反枝苋的种群密度来限制其竞争能力,防止反枝苋在湿地中生长建群和扩散入侵。     

Mycorrhizal feedback is not associated with the outcome of competition in old‐field perennial plants

The symbiosis between plants and arbuscular mycorrhizal (AM) fungi is hypothesized to be an important contributor to plant–soil feedbacks, which can influence the outcome of inter‐specific competition. Mycorrhizal feedbacks can be conspecific, which affects individuals of the same species, or heterospecific, which affects individuals of a different species. When heterospecific feedbacks are more positive than conspecific feedbacks, heterospecific individuals are expected to outcompete conspecific individuals. To test this hypothesis, we quantified conspecific mycorrhizal feedback for Plantago lanceolata as a focal species, and heterospecific mycorrhizal feedbacks for 21 competitor old‐field species using mycorrhizae cultured with P. lanceolata. We quantified inter‐specific competition against the focal species by growing the 21 old‐field species with and without P. lanceolata in the presence of mycorrhizae cultured with P. lanceolata. Heterospecific and conspecific feedbacks were both positive, and average heterospecific feedbacks exceeded conspecific feedback by 75%. Competition suppressed P. lanceolata biomass by 14% and average competitor biomass was reduced by 44% in the presence of P. lanceolata, and these effects varied with competitor species identity. Contrary to predictions, the magnitude of heterospecific feedbacks did not predict the ability of competitor species to either suppress or resist suppression by P. lanceolata. Instead, the outcome of competition was significantly and positively correlated with intrinsic growth rate, measured as biomass of competitor species five weeks after germination in non‐inoculated conditions. Our findings suggest that species experiencing more positive mycorrhizal feedbacks than a competitor do not necessarily have a competitive advantage. Mycorrhizal mediated soil feedbacks may be less important than intrinsic differences in growth rate in determining competitive outcomes.     

Simulated effects of herb competition on planted Quercus faginea seedlings in Mediterranean abandoned cropland

Abstract. We tested simulated effects of herb competition on the performance of planted seedlings of Quercus faginea ssp. faginea in Mediterranean abandoned cropland. We produced three types of environment with respect to herb competition: absence of competition (AC), below‐ground competition (BGC), and total competition (TC). We assessed the performance of Q. faginea seedlings in each treatment in five ways: (1) seedling mortality, (2) leaf length and total plant leaf area, (3) water potential, (4) total biomass and biomass allocation, and (5) non‐structural carbohydrate storage in different plant organs. We also measured (6) soil moisture at different depths and (7) biomass production of herbs. The TC treatment reduced water availability more than the BGC treatment, in agreement with the most pronounced water stress in seedlings under TC conditions. BGC and TC treatments showed a high and similar seedling mortality, which was one order of magnitude higher than that in the AC treatment. Competition treatments affected glucose concentration in both shoots and roots, and followed the rank TC > BGC > AC. Q. faginea seedlings might compensate a lower water availability through glucose accumulation in leaves to reduce the osmotic potential. There was a maximum starch concentration in the BGC treatment that hints that a moderate resource limitation would limit tissue growth but not carbon assimilation. We conclude that the negative effects of herbs on Q. faginea seedlings are mostly a result of competition for water, and that this competition is noticeable since the earliest stages of the establishment. Complete weed removal is a technique that would strongly improve seedling survivorship.     

Size-uneven competition and resource availability: A factorial experiment on seedling establishment of three Mediterranean species

Abstract

A factorial experiment was conducted in order to test whether the ranking of competitive ability at seedling stage is affected by different resource availability and competition asymmetry levels. Seedlings of three Mediterranean species (Rhamnus alaternus, Ampelodesmos mauritanicus and Cistus incanus) were grown at two competition intensities with different water and nutrient levels. Plants responses were expressed as mortality, biomass production and plant architecture. Data were also analysed by an index of competitive response intensity. A clear ranking of competitive response was observed, with Rhamnus > Ampelodesmos > Cistus. Competition was affected both by water and nutrient availability, but none of these factors modified competitive ranking. Seedling competition behaviour appeared to be related to species successional roles.     

不同氮素水平下入侵种豚草与本地种黄花蒿、蒙古蒿的竞争关系

已有研究表明,土壤氮素增加可提高外来植物的入侵性,降低本地植物的竞争力.为揭示全球氮沉降对入侵种与本地种之间竞争关系的影响,我们于2010年5-8月在中国科学院北京森林生态系统定位研究站温室内,采用取代系列实验方法(standard replacement experiment),研究了3个氮素水平下入侵种豚草(Ambrosia artemisiifolia)与本地种黄花蒿(Artemisia annua)、蒙古蒿(Artemisia mongolica)的生长特征及种内、种间竞争关系的变化.实验采用双因素-随机区组设计,设置了低氮、中氮和高氮3个氮素水平,每一氮素水平分别设置豚草和黄花蒿、豚草和蒙古蒿组成的竞争实验,生长90 d后测量株高和生物量.结果表明:单栽情况下,随氮素水平的增加3个物种的株高均增加,而生物量均无显著变化;混栽情况下,3个物种株高和生物量随氮素水平的增加变化各异,豚草呈极显著增加趋势,而黄花蒿无明显变化,蒙古蒿则先增加后减少.豚草的快速生长使其在竞争中处于优势地位,对本地种黄花蒿和蒙古蒿产生明显的竞争效应.但不同氮素水平下,豚草对本地种的竞争力不同:低氮素水平下,豚草＜两个本地种;中氮素水平下,黄花蒿＜豚草＜蒙古蒿;高氮素水平下,豚草＞两个本地种.氮素添加显著提高了豚草的种间竞争力,改变了豚草与本地种之间的竞争关系,使竞争有利于入侵种.据此推测,在全球变化的背景下,氮沉降的增加将会促进外来种豚草的入侵,增加本地群落的可入侵性.     

Variation in the establishment of a non-native annual grass influences competitive interactions with Mojave Desert perennials

Competition between native and non-native species can change the composition and structure of plant communities, but in deserts, the highly variable timing of resource availability also influences non-native plant establishment, thus modulating their impacts on native species. In a field experiment, we varied densities of the non-native annual grass Bromus madritensis ssp. rubens around individuals of three native Mojave Desert perennials—Larrea tridentata, Achnatherum hymenoides, and Pleuraphis rigida—in either winter or spring. For comparison, additional plots were prepared for the same perennial species and seasons, but with a mixture of native annual species as neighbors. Growth of perennials declined when Bromus was established in winter because Bromus stands had 2–3 months of growth and high water use before perennial growth began. However, water potentials for the perennials were not significantly reduced, suggesting that direct competition for water may not be the major mechanism driving reduced perennial growth. The impact of Bromus on Larrea was lower than for the two perennial grasses, likely because Larrea maintains low growth rates throughout the year, even after Bromus has completed its life cycle. This result contrasts with the perennial grasses, whose phenology completely overlaps with (Achnatherum) or closely follows (Pleuraphis) that of Bromus. In comparison, Bromus plants established in spring were smaller than those established in winter and thus did not effectively reduce growth of the perennials. Growth of perennials with mixed annuals as neighbors also did not differ from those with Bromus neighbors of equivalent biomass, but stands of these native annuals did not achieve the high biomass of Bromus stands that were necessary to reduce perennial growth. Seed dormancy and narrow requirements for seedling survivorship of native annuals produce densities and biomass lower than those achieved by Bromus; thus, impacts of native Mojave Desert annuals on perennials are expected to be lower than those of Bromus.     

Competition between Salicornia europaea and Atriplex prostrata (Chenopodiaceae) along an experimental salinity gradient

Salicornia europaea L. is a halophyte that often occupies the lowestand most saline (>3.5% total salt) areas of salt marshes. Atriplexprostrata Boucher is less salt tolerant than S. europaea and oftengrows in a less saline (<2.0% total salts) zone adjacent to S. europaea. The purpose of this experiment was to determine thecompetitive outcome when these two species are grown at differentsalinities to ascertain the extent salinity and competition affect plantzonation. Plants were grown in a de Wit replacement series at 85, 170,and 340 mM NaCl in half-strength Hoagland's no. 2 nutrient solution fortwo months. There was a significant effect of salt concentration,competition, and their interaction on biomass production of S. europaea plants. However, only salt concentration significantly affectedbiomass production of A. prostrata plants. Results of thisexperiment confirmed the results of other studies that demonstrated thatthe more salt tolerant species were less competitive at lower salinities. Atriplex prostrata was the better competitor at 85 mM NaCl, whereasS. europaea was the better competitor at 340 mM NaClbecause growth of A. prostrata was inhibited. At 170 mMNaCl, A. prostrata biomass production decreased more than S. europaea biomass in mixed culture.     

The role of microsite sunlight environment on growth,architecture, and resource allocation in dominant Acacia tree seedlings,in Serengeti,East Africa

Seedling establishment is a critical life history stage for savanna tree recruitment due to variability in resource availability. While tree–grass competition for water is recognized as an important driver of tree seedling mortality, the importance of sunlight exposure on tree seedling performance has received little attention in savanna ecosystems despite variable seedling light environments caused by heterogeneity in biomass of the grass canopy. We studied the seasonal sunlight micro-environment for two dominant East African tree species (Acacia?=?Vachellia) robusta (Burch) and A. tortilis (Forssk) under natural field conditions. In the Serengeti National Park, Tanzania, A. robusta trees occur in tall grasslands of the north (shady) and A. tortilis in the southern short grasslands (less shaded). We also designed a greenhouse experiment to quantify sunlight effects on seedling growth, architecture, and resource allocation traits. In the field, A. robusta seedlings were associated with lower understorey sunlight during the wet season compared to A. tortilis, with this trend switching during the dry season. In the greenhouse experiments, under low sunlight (25% radiation), A. robusta gained height faster than A. tortilis and self-shading among canopy leaves was evident in A. tortilis but not A. robusta. Biomass allocation to leaves, stems, and roots differed between species under different light environments suggesting phenotypic plasticity in response to variable light availability. Our study suggests that microsite light variability should be incorporated in models of the spatial and temporal variability of savanna tree recruitment.