Selection of shade trees in forest restoration plantings should not be based on crown tree architecture alone

The planting of tree seedlings is a common restoration technique in the tropics, and using large‐crowned, fast‐growing shade species is recommended to suppress invasive grasses and accelerate forest succession. We analyzed the effectiveness of shade species in shading the forest floor during the rainy and dry seasons at young forest restoration sites, whether shade changes according to site for a given species, and whether crown architecture can predict the shade level. We measured the photosynthetically active radiation (PAR) intercepted by the tree crowns of 14 species in two 3‐year‐old restoration plantings. The ability to predict shade based on crown architecture traits was evaluated using multiple linear regressions. The interception of PAR varied according to species, site, and season for seven species and was generally higher during the rainy season. Low values of tree and first branch height and high values of trunk diameter and mean area of a leaf predicted greater light interception. For the dry season, the ability to predict PAR interception was weaker than that for the rainy season and affected by a shorter tree height and a greater crown area. The crown architecture of shade species did not completely predict their shading ability, and the preselection of shade species for forest restoration purposes based only on crown architecture traits is not effective. Therefore, it is important to consider other factors, such as how long trees retain their leaves throughout the year and the soil and management conditions of the sites undergoing restoration, during the selection of species.     

Regulation of gut function varies with life-history traits in chuckwallas (Sauromalus obesus: Iguanidae)

We examined the effects of hibernation and fasting on intestinal glucose and proline uptake rates of chuckwallas (Sauromalus obesus) and on the size of organs directly or indirectly related to digestion. These lizards show geographic variation in body size and growth rate that parallels an elevational gradient in our study area. At low elevation, food is available only for a short time during the spring; at high elevation, food may also be available during summer and autumn, depending on rainfall conditions in a given year. We hypothesized that low-elevation lizards with a short season of food availability would show more pronounced regulation of gut size and function than high-elevation lizards with prolonged or bimodal food availability. Hibernating lizards from both elevations had significantly lower uptake rates per milligram intestine for both nutrients, and lower small intestine mass, than active lizards. The combination of these two effects resulted in significantly lower total nutrient uptake in hibernating animals compared to active ones. The stomach, large intestine, and cecum showed lower masses in hibernators, but these results were not statistically significant. The heart, kidney, and liver showed no difference in mass between hibernating and nonhibernating animals. Lizards from low elevations with a short growing season also showed a greater increase in both uptake rates and small intestine mass from the hibernating to the active state, compared to those from high elevations with longer growing seasons. Thus, compared to those from long growing season areas, lizards from short growing season areas have equal uptake capacity during hibernation but much higher uptake capacity while active and feeding. This pattern of regulation of gut function may or may not be an adaptive response, but it is consistent with variation in life-history characteristics among populations. In areas with a short season, those lizards that can extract nutrients quickly and then reduce the gut will be favored; in areas where food may be available later in the year, those lizards that maintain an active gut would be favored. While other researchers have found much greater magnitudes of gut regulation when making comparisons among species, we find the different patterns of change in gut function between different populations of chuckwallas particularly intriguing because they occur within a single species.     

Responses of crown architecture in Betula pendula to competition are dependent on the species of neighbouring trees

We measured the growth responses of individual shoots and branches of Betula pendula when growing next to trees of the same species or Pinus sylvestris, Larix sibirica or Alnus glutinosa. We used the three-dimensionally digitized response variables and the size and distance of trees growing within a 5-m radius of the study trees to establish a relationship between tree performance and the effect of competing neighbouring tree species on crown architecture. B. pendula was able to modify its crown architecture and thus alter its strategy to compete with different neighbours. Trees of B. pendula growing beside species counterparts had the highest growth of new long shoots in relation to the already existing branch length [growth vigour (GV)], while GV was the lowest next to L. sibirica. With B. pendula or P. sylvestris as its main neighbour, B. pendula invested in short shoots by growing them rather densely in short branches with limited numbers, whereas with L. sibirica the number, length and angle of the branches were high. The competitive response was also strongly dependent on tree ontogeny and the shoot and branch characteristics were significantly affected by their location inside the crown. B. pendula was able to respond to the challenges posed by its neighbours, which was also reflected in the GV. The ability to maintain steady growth with alternative crown designs in different neighbourhoods reflects plasticity in the crown responses.     

Crown hollowing as a consequence of early shedding of leaves and shoots

Leafing pattern has long been considered as an important element characterizing the growth strategy of tree species; however, the consequences of leafing pattern for tree-crown formation have not been fully understood. To address this issue, the dynamic events (growth, birth, and death) of current-year shoots and leaves were investigated together with their location in saplings of a pioneer tree, Alnus sieboldiana. The leafing pattern was characterized by successive emergence and shedding of short-lived leaves. The combination of successive leafing and within-crown variation in leaf production brought about characteristic outcomes in crown morphology. In the outer crown, because of continuous leaf production, the shoots achieved great extension and enormous daughter shoot production, resulting in rapid expansion of the crown. In contrast, in the inner crown, due to early termination of leaf production, the shoots completely lost their leaves early in the growing season and consequently themselves died and were shed within the season. Such quick shedding of shoots caused “crown hollowing”, i.e., the interior crown consisted of primary branches with little secondary development or foliage. These dynamic features are an effective adaptive strategy in early succession but also may be a disadvantage to maintaining foliage for longer period. Crown maintenance associated with the longevity of structural components is thought to play an important role in survival strategy of tree species.     

Asynchronicity in root and shoot phenology in grasses and woody plants

Phenology is central to understanding vegetation response to climate change, as well as vegetation effects on plant resources, but most temporal production data is based on shoots, especially those of trees. In contrast, most production in temperate and colder regions is belowground, and is frequently dominated by grasses. We report root and shoot phenology in 7‐year old monocultures of 10 dominant species (five woody species, five grasses) in southern Canada. Woody shoot production was greatest about 8 weeks before the peak of root production, whereas grass shoot maxima preceded root maxima by 2–4 weeks. Over the growing season, woody root, and grass root and shoot production increased significantly with soil temperature. In contrast, the timing of woody shoot production was not related to soil temperature (r=0.01). The duration of root production was significantly greater than that of shoot production (grasses: 22%, woody species: 54%). Woody species produced cooler and moister soils than grasses, but growth forms did not affect seasonal patterns of soil conditions. Although woody shoots are the current benchmark for phenology studies, the other three components examined here (woody plant roots, grass shoots and roots) differed greatly in peak production time, as well as production duration. These results highlight that shoot and root phenology is not coincident, and further, that major plant growth forms differ in their timing of above‐ and belowground production. Thus, considering total plant phenology instead of only tree shoot phenology should provide a better understanding of ecosystem response to climate change.     

Hydrostatic constraints on morphological exploitation of light in tall Sequoia sempervirens trees

We studied changes in morphological and physiological characteristics of leaves and shoots along a height gradient in Sequoia sempervirens, the tallest tree species on Earth, to investigate whether morphological and physiological acclimation to the vertical light gradient was constrained by hydrostatic limitation in the upper crown. Bulk leaf water potential (Psi) decreased linearly and light availability increased exponentially with increasing height in the crown. During the wet season, Psi was lower in the outer than inner crown. C isotope composition of leaves (delta(13)C) increased with increasing height indicating greater photosynthetic water use efficiency in the upper crown. Leaf and shoot morphology changed continuously with height. In contrast, their relationships with light availability were discontinuous: morphological characteristics did not correspond to increasing light availability above 55-85 m. Mass-based chlorophyll concentration (chl) decreased with increasing height and increasing light availability. In contrast, area-based chl remained constant or increased with increasing height. Mass-based maximum rate of net photosynthesis (P (max)) decreased with increasing height, whereas area-based P (max) reached maximum at 78.4 m and decreased with increasing height thereafter. Mass-based P (max) increased with increasing shoot mass per area (SMA), whereas area-based P (max) was not correlated with SMA in the upper crown. Our results suggest that hydrostatic limitation of morphological development constrains exploitation of light in the upper crown and contributes to reduced photosynthetic rates and, ultimately, reduced height growth at the tops of tall S. sempervirens trees.     

Common allometric response of open-grown leader shoots to tree height in co-occurring deciduous broadleaved trees

Background and Aims

Morphology of crown shoots changes with tree height. The height of forest trees is usually correlated with the light environment and this makes it difficult to separate the effects of tree size and of light conditions on the morphological plasticity of crown shoots. This paper addresses the tree-height dependence of shoot traits under full-light conditions where a tree crown is not shaded by other crowns.

Methods

Focus is given to relationships between tree height and top-shoot traits, which include the shoot''s leaf-blades and non-leafy mass, its total leaf-blade area and the length and basal diameter of the shoot''s stem. We examine the allometric characteristics of open-grown current-year leader shoots at the tops of forest tree crowns up to 24 m high and quantify their responses to tree height in 13 co-occurring deciduous hardwood species in a cool-temperate forest in northern Japan.

Key Results

Dry mass allocated to leaf blades in a leader shoot increased with tree height in all 13 species. Specific leaf area decreased with tree height. Stem basal area was almost proportional to total leaf area in a leader shoot, where the proportionality constant did not depend on tree height, irrespective of species. Stem length for a given stem diameter decreased with tree height.

Conclusions

In the 13 species observed, height-dependent changes in allometry of leader shoots were convergent. This finding suggests that there is a common functional constraint in tree-height development. Under full-light conditions, leader shoots of tall trees naturally experience more severe water stress than those of short trees. We hypothesize that the height dependence of shoot allometry detected reflects an integrated response to height-associated water stress, which contributes to successful crown expansion and height gain.     

Positive Effects of Non-Native Grasses on the Growth of a Native Annual in a Southern California Ecosystem

Fire disturbance is considered a major factor in the promotion of non-native plant species. Non-native grasses are adapted to fire and can alter environmental conditions and reduce resource availability in native coastal sage scrub and chaparral communities of southern California. In these communities persistence of non-native grasses following fire can inhibit establishment and growth of woody species. This may allow certain native herbaceous species to colonize and persist beneath gaps in the canopy. A field manipulative experiment with control, litter, and bare ground treatments was used to examine the impact of non-native grasses on growth and establishment of a native herbaceous species, Cryptantha muricata. C. muricata seedling survival, growth, and reproduction were greatest in the control treatment where non-native grasses were present. C. muricata plants growing in the presence of non-native grasses produced more than twice the number of flowers and more than twice the reproductive biomass of plants growing in the treatments where non-native grasses were removed. Total biomass and number of fruits were also greater in the plants growing in the presence of non-native grasses. Total biomass and reproductive biomass was also greater in late germinants than early germinants growing in the presence of non-native grasses. This study suggests a potential positive effect of non-native grasses on the performance of a particular native annual in a southern California ecosystem.     

Fast and furious: Early differences in growth rate drive short‐term plant dominance and exclusion under eutrophication

1. The reduction of plant diversity following eutrophication threatens many ecosystems worldwide. Yet, the mechanisms by which species are lost following nutrient enrichment are still not completely understood, nor are the details of when such mechanisms act during the growing season, which hampers understanding and the development of mitigation strategies.
2. Using a common garden competition experiment, we found that early‐season differences in growth rates among five perennial grass species measured in monoculture predicted short‐term competitive dominance in pairwise combinations and that the proportion of variance explained was particularly greater under a fertilization treatment.
3. We also examined the role of early‐season growth rate in determining the outcome of competition along an experimental nutrient gradient in an alpine meadow. Early differences in growth rate between species predicted short‐term competitive dominance under both ambient and fertilized conditions and competitive exclusion under fertilized conditions.
4. The results of these two studies suggest that plant species growing faster during the early stage of the growing season gain a competitive advantage over species that initially grow more slowly, and that this advantage is magnified under fertilization. This finding is consistent with the theory of asymmetric competition for light in which fast‐growing species can intercept incident light and hence outcompete and exclude slower‐growing (and hence shorter) species. We predict that the current chronic nutrient inputs into many terrestrial ecosystems worldwide will reduce plant diversity and maintain a low biodiversity state by continuously favoring fast‐growing species. Biodiversity management strategies should focus on controlling nutrient inputs and reducing the growth of fast‐growing species early in the season.

     

How season of grazing and herbivore selectivity influence monsoon tall-grass communities of northern Australia

Abstract. The hypothesis that season of defoliation and herbivore selectivity may be as important as level of use in determining plant community response to grazing was tested in a monsoon grassland in northern Australia. Plots, dominated by the tussock grasses Themeda triandra and Chrysopogon fallax, were grazed by cattle at low, medium and high rates of utilization in either the early wet, late wet or dry seasons. Effects of grazing on species composition were greatest in the early wet season when high rates of utilization significantly reduced the proportion and occurrence of Themeda and increased the proportion of forbs. Grazing in the dry season had no significant effect on composition. At medium and high levels of utilization in the early wet season, the pasture responded negatively to defoliation, only partially compensating for plant tissue lost to herbivory. The negative response to defoliation carried over to the next wet season when these same medium and high-grazing treatments produced only 80 % and 60 % growth, respectively, of that in treatments grazed at low levels of utilization or those grazed during the dry season. The frequency of Themeda was still lower, and that of annual grasses and non-leguminous forbs higher, in plots that had been grazed at a high rate of utilization for just eight weeks in the early wet season two years previously. Species richness and diversity were also significantly affected by this grazing disturbance. If species composition is to be maintained in these grasslands then stocking rates must be set at low levels to cope with the combined effect of undercompensation in response to defoliation in the wet season and strong dietary preferences for grazing sensitive species.     

大针茅草原地上生物量形成的规律与特点

大针茅草原能进行光合作用的时间为160—170天。地上生物量的季节生长曲线呈单峰型,适宜的收获期在8月份。地上生物量的增长与群落的高度增长呈明显相关(R=0.959)。立枯量于6月份开始出现,其增长规律与绿色量呈相反的趋势。刈割后的再生草量以春季(5月份)刈割后的产量最高。仲夏(7月份)刈割对草场生产力的威胁最大。群落产量结构的研究表明：5己于人cm以上可供牲畜采食的部分约占总产量的70—80％。     

C3 shrub expansion in a C4 grassland: positive post-fire responses in resources and shoot growth

Changes in land management and reductions in fire frequency have enabled woody species to increase in grasslands worldwide. Nevertheless, fire is rarely eliminated from grasslands, and for shrubs to survive, they must be able cope with fire and replace aboveground structures. Because new shoots may have more available solar radiation, greater root?:?shoot ratios, and thus more resources available belowground after fire compared to undisturbed shrub communities, we hypothesized that carbon, nutrient, and water relations may be enhanced in stems compared to those in an undisturbed grassland. However, this same post-fire resource pulse stimulates the grasses and may intensify competitive interactions between shrubs and grasses. To test these predictions, we measured seasonal patterns in net photosynthesis (A), predawn xylem pressure potentials (XPP), leaf nitrogen (N) content, and productivity of Cornus drummondii shoots from shrub patches (islands) of different sizes in mesic grasslands burned annually, burned infrequently, and protected from fire. Seasonal average A was 20% higher (P = 0.016) in burned than in unburned shrubs, regardless of island size. Shrubs in burned sites also produced shoots with higher leaf N than unburned shrubs, and N content was higher in leaves from small islands compared to large islands (P < 0.0001). Burning caused a decrease in late summer predawn XPP in small islands (-3.1 MPa), whereas burned large islands did not differ from unburned shrubs. Post-fire productivity of new shoots was significantly greater compared to shoots in unburned sites. These results indicate that a transient period of high resource availability after fire allows for increased growth and rapid recovery of grassland shrubs. Thus, although fire has a negative effect on aboveground biomass of shrubs, the post-fire increases in resource availability, which enhance growth in the dominant grasses, are also important for recovery of woody species.     

PLANT SIZE AND FORM IN THE UNDERSTORY PALM GENUS GEONOMA: ARE SPECIES VARIATIONS ON A THEME?

This study addressed the hypothesis that phylogenetic changes in plant size at reproductive maturity may have facilitated adaptive radiation of Geonoma species within rain forest understory habitats. Leaf size, leaf form, plant size, and growth form were compared within and among 23 species of Geonoma from lowland and montane rain forest areas of Costa Rica and Colombia. Leaf size was significantly correlated with crown height in 18 of the 21 species examined, and with stem diameter in 17 of the species. In species characterized by a gradual ontogenetic transition from bifid to dissected leaves, shoots with bifid leaves were significantly smaller than shoots with dissected leaves with respect to rachis length, number of plications, and stem diameter. Among species, stem diameter below the crown explained 74% of the variation in leaf size (rachis length). Crown height and stem diameter were positively correlated among clustered species, but not among solitary species or all species combined. Leaf dissection was correlated with crown height among the 17 species with dissected leaves; species with bifid leaves were significantly smaller than species with dissected leaves with respect to leaf size and stem diameter. Solitary species had larger leaves and larger stem diameters than clustered species at the same crown heights. Morphological patterns among species generally followed within-species trends. These patterns suggest that Geonoma species are variants on a generic theme:within and among species, leaf size and complexity of form increase with stem diameter and crown height. Solitary and clustered growth forms appear to be morphologically convergent; within each of these architectural groups, the generic theme still applies. Evolutionary changes in leaf size, leaf form, and plant size, however, have clearly involved other factors in addition to variation in plant size.     

Changes in crown development patterns and current-year shoot structure with light environment and tree height in Fagus crenata (Fagaceae)

The relative effects of light and tree height on the architecture of leader crowns (i.e., the leading section of the main trunk, 100 cm in length) and current-year shoots for a canopy species, Fagus crenata, occupying both the ridge top and the valley bottom in a cool-temperate forest in Japan were investigated. For leader crowns, the number of current-year shoots and leaves increased with increasing tree height, whereas the mean length of current-year shoots increased with increasing relative photon flux density (PFD). The leader crown area decreased, and the depth and leaf area index of leader crowns increased, with increasing relative PFD. The mass of current-year shoots increased with relative PFD. However, this total mass was allocated differently between stems and leaves depending on tree height, such that the relative allocation to stems increased with increasing tree height. Furthermore, stem structures within current-year shoots also changed with height, such that taller trees produced thicker and shorter stems of the same volume. In contrast, leaf structure and leaf biomass allocations changed with relative PFD. Specific leaf area decreased with increasing relative PFD. In addition, leaf number increased more rapidly with increasing individual leaf mass for trees exposed to greater relative PFD. Consequently, the total leaf area supported by a stem of a given diameter decreased with increasing tree height and relative PFD. Thus, the architecture of leader crowns and current-year shoots were related differently to light and tree height, which are considered important for efficient light capture and the growth of small and tall trees in different environments.     

Effect of tall-grass invasion on the flowering-related functional pattern of submediterranean hay-meadows

Several studies demonstrated that abandonment changes the functional composition of grasslands; nevertheless, little is known about the effects of grassland abandonment on the flowering-related functional pattern. We hypothesized that invasion by tall grasses affects this pattern. We counted the number of flowering shoots per species at five times during the growing season, in 80 plots placed in mown and in abandoned grasslands (central Apennines), and assessed the differences in the trait composition of flowering species between the two treatments. The selected traits were linked to resource acquisition and stress tolerance strategies. Our results indicated that abiotic environmental control is prevalent in determining the phenological pattern in both conditions and in accordance with the phenological “mid-domain hypothesis”. We demonstrated that when the dominant species is a tall grass with competitive behaviour, the magnitude of this phenomenon is amplified due to the abiotic changes yielded by the tall grass invasion. Indeed, in the central and late phases of the growing season (when invasive tall grasses are growing and blooming), abandoned grasslands were marked by a set of traits devoted to stress tolerance or underlying a long reproductive cycle or linked to competition for light.     

Testing a new hypothesis: plant vigor and phylloxera distribution on wild grape in Arizona

Summary Longer, meaning more vigorous, shoots of a wild grape clone (Vitis arizonica) were more susceptible to attack by second and third generations of leaf-galling grape phylloxera,Daktulopsphaira vitifoliae, as the growing season progressed. Although there was no significant difference in mean shoot length between attacked and unattacked shoots within a clone at the beginning of shoot elongation, attacked shoots were significantly longer than unattacked shoots when elongation had ceased (P<0.01). Also, long attacked shoots had a significantly greater population of phylloxera galls than short attacked shoots (P<0.01) as the season progressed. The phylloxera population on long shoots increased rapidly while the population on short shoots remained the same. Longer shoots also produced significantly more axillary shoots than shorter shoots as the season progressed (P<0.001), and the number of axillary shoots accounted for 66 percent of the variance in number of attacked leaves on a shoot. Experimental evidence showed that there was a significantly greater percentage of available leaves attacked on long shoots than on short shoots (P<0.05) and the leaves on long shoots generally had a greater number of galls per leaf. The relationship between shoot length and probability of attack was also tested by comparing shoots lengths of 10 attacked clones and 10 unattacked clones at a second location. Mean shoot lengths of attacked clones were significantly longer than mean shoot lengths of unattacked clones (P<0.05), and mean shoot lengths of attacked shoots within a clone were significantly longer than unattacked shoots (P<0.001). Longer shoot length accounted for 81 percent of the variance in probability of attack. The reason for this pattern of attack was that long shoots produced newly expanding leaves over a longer time during the growing season and multivoltine phylloxera require undifferentiated tissue to initiate gall formation. Patterns of attack within a shoot were characterized by an uneven distribution of galls among leaves. This was due to development time between generations and the current availability of undifferentiated tissue at times of colonization. This study supports the hypothesis that some herbivore species are favored more by vigorous plants than by stressed plants.     

Shoot Growth and Mortality Patterns of Urtica dioica, a Clonal Forb

The growth and mortality patterns of the clonal forb Urticadioica were investigated at the level of the individual shootin two growing seasons, 1991 and 1992, in a natural stand. Shootheight and diameter at ground level of each shoot tagged inspring were measured repeatedly five times during the growingseason. Dry weights of these repeatedly measured shoots wereestimated using an allometric relationship between dry weight,height and diameter of harvested shoots. A large decrease inshoot density occurred with stand development from the beginningof the growing season in both the years: (1) shoot survivalrate was about 30% at the end of the growing season; (2) shootmortality rate per 10 x 10 cm subplot between censuses was positivelydependent on shoot density per subplot; (3) the mortality rateof individual shoots was negatively dependent on shoot size(height, diameter and weight) at each growing stage, suggestingone-sided competition between living and dying shoots; (4) shootsize (height, diameter and weight) variability in terms of thecoefficient of variation and skewness decreased in accordancewith shoot mortality. Symmetric competition between living shootswas detected by regression analysis based on a model for individualshoot growth considering the degree of competitive asymmetry.However, the competitive effect on individual shoot growth wasvery small (nearly absent). The mortality pattern of Urticadioica indicates that shoot self-thinning occurred from theearly growing stage as in non-clonal crowded monospecific stands,and contrasts with many clonal plants where shoot self-thinningrarely occurs or, if any, is confined only to a short periodof the later growing stage. The pattern of growth and competitionbetween living shoots of Urtica dioica contrasts with non-clonalcrowded plants undergoing intense competition (usually asymmetric)between individuals, but is a common feature of many clonalplants where shoot competition is supposed to be reduced by'physiological integration' between shoots. These form a newpattern not reported yet for clonal plants. It is pointed outthat clonal plants show a wider spectrum of the growth, competitionand mortality patterns of shoots than non-clonals. Some possiblemechanisms for the pattern of Urtica dioica are discussed.Copyright1995, 1999 Academic Press Shoot competition, diffusion model, individual shoot growth, shoot self-thinning, shoot size variability, Urtica dioica L     

The effect of crown position and date of sampling on biomass,nutrient concentrations and contents of needles and shoots in European larch

 The nutrient concentrations and contents of needles and shoots of 22-year-old European larch (Larix decidua Mill.) were evaluated with respect to crown position, age of tissues and sampling date during a complete growing season. Concentrations of N, P, K, Ca, Mg and Zn in the needles and of N, P and K in the shoots differed significantly among the dates of sampling. The concentrations of N and Mn in the needles and all nutrients in the shoots (except Mg) also differed significantly with crown position. Maximum needle biomass was observed in the middle crown position (55% of the total) and maximum shoot biomass, in the lower crown position (52% of the total). Maximum needle and shoot nutrient contents were observed in the middle position of the living crown for long shoot, short shoot-1, short shoot-2, short shoot-3 and, short shoot-4 age classes while highest contents for short shoot-5 and short shoot-6 age classes were observed in the lower crown position. Biases up to 42% for Mg in the needles and 200% for K in the shoots were obtained when only long shoot tissues are used for content evaluation. For needles and shoots, Mg and K are more difficult nutrients to evaluate. A sampling methodology is proposed for evaluating nutrient contents of the living crown. Accepted: 10 August 1995     

Plant feedbacks increase the temporal heterogeneity of soil moisture

Plant feedbacks on resource levels are well-known, but feedbacks on resource variability have received little attention. Semi-arid grasslands have greater temporal heterogeneity of rainfall than mesic forests, leading to the possibility that grasses further enhance this variability as a mechanism for excluding woody plants originating in habitats with less heterogeneity. Here we test the hypothesis that grasses create greater levels of temporal heterogeneity of soil resources than do woody plants. We used monocultures of five replicate species of both growth forms. Daily soil moisture measurements taken 10 and 30 cm beneath monocultures over a growing season showed that temporal heterogeneity was significantly greater under grasses than under woody plants. This occurred during a dry period when plants are most likely to compete for moisture. Differences in temporal heterogeneity between growth forms were related to differences in their abilities to reduce soil moisture: during the dry period, the net effect of vegetation on moisture 10 cm deep was greatest under grasses. Although the rate of change of soil moisture was higher under grasses, the growth forms exploited different depths of soil moisture: soils 10 cm deep were driest under grasses, but soils 30 cm deep were driest under woody species. In summary, grasses increased within-season resource variability in a habitat already characterized by high among-year variability.