Desert Shrub Larrea Tridentata Research Articles

Non-structural carbohydrate dynamics associated with antecedent stem water potential and air temperature in a dominant desert shrub.

Non-structural carbohydrates (NSCs) are necessary for plant growth and affected by plant water status, but the temporal dynamics of water stress impacts on NSC are not well understood. We evaluated how seasonal NSC concentrations varied with plant water status (predawn xylem water potential, Ψ) and air temperature (T) in the evergreen desert shrub Larrea tridentata. Aboveground sugar and starch concentrations were measured weekly or monthly for ~1.5 years on 6-12 shrubs simultaneously instrumented with automated stem psychrometers; leaf photosynthesis (Anet ) was measured monthly for 1 year. Leaf sugar increased during the dry, premonsoon period, associated with lower Ψ (greater water stress) and high T. Leaf sugar accumulation coincided with declines in leaf starch and stem sugar, suggesting the prioritization of leaf sugar during low photosynthetic uptake. Leaf starch was strongly correlated with Anet and peaked during the spring and monsoon seasons, while stem starch remained relatively constant except for depletion during the monsoon. Recent photosynthate appeared sufficient to support spring growth, while monsoon growth required the remobilization of stem starch reserves. The coordinated responses of different NSC fractions to water status, photosynthesis, and growth demands suggest that NSCs serve multiple functions under extreme environmental conditions, including severe drought.

Antecedent soil water content and vapor pressure deficit interactively control water potential in Larrea tridentata.

Plant water potential Ψ is regulated by stomatal responses to atmospheric moisture demand D and soil water availability W, but the timescales of influence and interactions between these drivers of plant Ψ are poorly understood. Here, we quantify the effects of antecedent D and W on plant Ψ in the desert shrub Larrea tridentata. Repeated measurements of plant baseline water potential ΨB and diurnal water potential ΨD were analyzed in a Bayesian framework to evaluate the influence of antecedent D and W at daily and subdaily timescales. Both ΨB and ΨD exhibited negative, 2- to 4-d lagged responses to daily-scale D; conversely, plant ΨD responded almost instantaneously to subdaily D, though the direction of this response depended on antecedent moisture conditions. Plant ΨB and ΨD responded positively and immediately (no lag) to shallow W, which contrasts the negative, lagged (6-7d) response to deep W. The changing sensitivity of ΨD to subdaily D highlights shifting modes of plant Ψ regulation: D effects on ΨD range from negative to neutral to positive depending on past conditions and time of day. Explicit consideration of antecedent conditions across multiple timescales can reveal important complexities in plant responses.

The Sensitivity of Evapotranspiration to Inter-Specific Plant Neighbor Interactions: Implications for Models

Evapotranspiration (ET) is an important water loss flux in ecosystem water cycles, and quantifying the spatial and temporal variation of ET can improve ecohydrological models in arid ecosystems. Plant neighbor interactions may be a source of spatial and temporal variation in ET due to their effects on the above- and belowground microclimate and increased water demand for transpiration. Over longer timescales (annual to multiple years), adjustments in plant physiological traits may occur in response to neighbor environments, potentially affecting the transpiration (T) component of ET. We used a dynamic soil water model to assess the sensitivity of ET and T estimates to neighbor effects on soil moisture via competition for water, aboveground microclimate effects via canopy shading, and physiological adjustments (specifically, root distribution, stomatal behavior, and canopy leaf area). We focus on a common desert shrub (Larrea tridentata) under different inter-specific neighbor environments and precipitation regimes. Neighbors impacted T of Larrea by as much as 75% at the patch scale (plant and surrounding soil) and 30% at the stand scale. Annual T estimates were highly sensitive to changes in soil moisture associated with competition for water, and the inclusion of physiological adjustments to neighbor environments significantly impacted seasonal T. Plant neighbor interactions can significantly influence ET and soil moisture, and their inclusion in models can help explain spatial and temporal variation in water fluxes in arid ecosystems. Furthermore, physiological adjustments to neighbor environments may be an important source of variation to include in models that operate over seasonal timescales or in studies focused on plant responses to precipitation under climate change.

Paradoxical cellular effects and biological role of the multifaceted compound nordihydroguaiaretic acid.

Nordihydroguaiaretic acid (NDGA) is a phenolic compound obtained from the leaves of the evergreen desert shrub Larrea tridentata (Creosote bush), which has been used anciently in folk medicine for the treatment of multiple diseases. At the molecular level, NDGA is a potent scavenger of reactive oxygen species. Lipoxygenase inhibition by NDGA has been broadly studied over several cell models; however, NDGA exerts other antioxidant properties and cytoprotective effects in non-tumor cells, which are related with its role as modulator of the nuclear factor erythroid 2-related factor 2 (Nrf2)/antioxidant response element (ARE) antioxidant pathway. In contrast, in tumor cells NDGA exerts pro-apoptotic activity and anti-tumor effects. Different effects of NDGA have been observed in mitochondria, where NDGA prevents mitochondrial damage in non-tumor cells and induces loss of mitochondrial function in tumor cells. Moreover, NDGA exerts beneficial effects in diverse diseases like cancer, renal damage, Huntington's disease, Alzheimer's disease, and other neurodegenerative pathologies. This work represents a critical review about relevant NDGA mechanisms, cellular effects, and signal pathways involved with possible useful effects.

Elevated CO2 increases plant uptake of organic and inorganic N in the desert shrub Larrea tridentata

Resource limitations, such as the availability of soil nitrogen (N), are expected to constrain continued increases in plant productivity under elevated atmospheric carbon dioxide (CO(2)). One potential but under-studied N source for supporting increased plant growth under elevated CO(2) is soil organic N. In arid ecosystems, there have been no studies examining plant organic N uptake to date. To assess the potential effects of elevated atmospheric CO(2) on plant N uptake dynamics, we quantified plant uptake of organic and inorganic N forms in the dominant desert shrub Larrea tridentata under controlled environmental conditions. Seedlings of L. tridentata were grown in the Mojave Desert (NV, USA) soils that had been continuously exposed to ambient or elevated atmospheric CO(2) for 8 years at the Nevada Desert FACE Facility. After 6 months of growth in environmentally controlled chambers under ambient (380 micromol mol(-1)) or elevated (600 micromol mol(-1)) CO(2), pots were injected with stable isotopically labeled sole-N sources ((13)C-[2]-(15)N glycine, (15)NH(4) (+), or (15)NO(3) (-)) and moved back to their respective chambers for the remainder of the study. Plants were destructively harvested at 0, 2, 10, 24, and 49 days. Plant uptake of soil N derived from glycine, NH(4) (+), and NO(3) (-) increased under elevated CO(2) at days 2 and 10. Further, root uptake of organic N as glycine occurred as intact amino acid within the first hour after N treatment, indicated by approximately 1:1 M enrichment ratios of (13)C:(15)N. Plant N uptake responses to elevated CO(2) are often species-specific and could potentially shift competitive interactions between co-occurring species. Thus, physiological changes in root N uptake dynamics coupled with previously observed changes in the availability of soil N resources could impact plant community structure as well as ecosystem nutrient cycling under increasing atmospheric CO(2) levels in the Mojave Desert.

Enhanced monsoon precipitation and nitrogen deposition affect leaf traits and photosynthesis differently in spring and summer in the desert shrub Larrea tridentata

Leaf-level CO2 assimilation (A(area)) can largely be predicted from stomatal conductance (g(s)), leaf morphology (SLA) and nitrogen (N) content (N(area)) in species across biomes and functional groups. The effects of simulated global change scenarios, increased summer monsoon rain (+H2O), N deposition (+N) and the combination (+H2O +N), were hypothesized to affect leaf trait-photosynthesis relationships differently in the short- and long-term for the desert shrub Larrea tridentata. During the spring, +H2O and +H2O +N plants had lower A(area) and g(s), but similar shoot water potential (Psi(shoot)) compared with control and +N plants; differences in A(area) were attributed to lower leaf N(area) and g(s). During the summer, +H2O and +H2O +N plants displayed higher A(area) than control and +N plants, which was attributed to higher Psi(shoot), g(s) and SLA. Throughout the year, A(area) was strongly correlated with g(s) but weakly correlated with leaf N(area) and SLA. We concluded that increased summer monsoon had a stronger effect on the performance of Larrea than increased N deposition. In the short term, the +H2O and +H2O +N treatments were associated with increasing A(area) in summer, but also with low leaf N(area) and lower A(area) in the long term the following spring.

Desert dogma revisited: coupling of stomatal conductance and photosynthesis in the desert shrub, Larrea tridentata

AbstractThe success of the desert shrub Larrea tridentata (creosotebush) has been largely attributed to temperature acclimation and stomatal control of photosynthesis (A) under drought stress. However, there is a paucity of field data on these relationships. To address this void, we conducted a joint field and modelling study that encompassed a diverse set of environmental conditions. At a Larrea‐dominated site in southern New Mexico we manipulated soil moisture during the growing season over a 2‐year period and measured plant pre‐dawn water potential (Ψpd), stomatal conductance (g) and A of individual shrubs. We used these data to develop a semi‐mechanistic photosynthesis model (A–Season) that explicitly couples internal CO2 (Ci) and g. Vapour pressure deficit (VPD) and Ψpd affect instantaneous g in a manner that is consistent with a biophysical model of stomatal regulation of leaf water potential. Ci is modelled as a function of g, derived from a simplification of a typical A–Ci curve. After incorporating the effects of growing temperature on stomatal behaviour, the model was able to capture the large diurnal fluctuations in A, g and Ci and the observed hysteresis in g versus Ci dynamics. Our field data and application of the A–Season model suggest that dogma attributed to Larrea's success is supported with regard to stomatal responses to VPD and Ψpd, but not for mechanisms of temperature acclimation and CO2 demand.

Effects of surface and sub-surface soil horizons on the seasonal performance of Larrea tridentata (creosotebush)

1. In warm arid and semiarid environments, the accumulation of clay minerals produces increasingly well developed soil horizons with the passage of time. Differences in the strength of development of two prominent soil horizons, silt- and clay-rich surface vesicular (Av), and clay-enriched subsurface argillic (Bt), may strongly influence the amount and seasonal continuity of plant-available water and the physiological activity of long-lived desert shrubs. Three sites were selected on an alluvial piedmont (bajada) in the Mojave Desert that varied in surface and subsurface horizon development. The first site, a deep deposit of stabilized dune sand, entirely lacked soil horizon development. The second site had a well developed surface stone pavement and underlying Av horizon, but lacked an argillic horizon in the sandy subsoil. The third had a well developed surface pavement and Av horizon, and a deeper, well developed clay-rich argillic horizon. Seasonal water potential and gas-exchange responses of the evergreen desert shrub Larrea tridentata[DC.] Cov., and volumetric soil water content (θ), were measured monthly in 1996 on these three soils in order to test the hypothesis that desert pavements, Av and subsurface Bt horizons differentially affect the effectiveness and utilization of seasonal precipitation. 2. Predawn and midday water potentials (ψpd and ψmid), net photosynthetic rates (Anet), and stomatal conductances (gs) in L. tridentata were highest in the deep, sandy dune soils lacking horizons that could restrict surface and subsurface infiltration. Plants growing in these soils also showed no physiological response to summer precipitation events. Following a single large precipitation event during the growing season (3·8 cm), the water potentials, Anet and gs in L. tridentata were similar in the first (sand dune) and second (pavement and Av horizon) sites. However, plant performance on these soil surfaces showed marked seasonal declines, and did not respond to a small pulse of summer rainfall. Plants growing at the third site (older soils with strongly developed pavement, Av and Bt horizons) had very low gas-exchange rates and water potentials. However, following convectional summer thunderstorms L. tridentata showed improved water relations and gas exchange in these soils. 3. Midday water potentials were frequently anomalously higher than predawn water potentials, up to +6 MPa late in the growing season, especially on soil surfaces with well developed soil horizons. This anomaly was due to seasonal decreases in ψpd accompanied by invariant midday ψ. In general, ψpd was correlated with θ across the depths measured. 4. Correlations between ψmid with θ at 35 cm increased dramatically with increasing Bt horizon development, suggesting that seasonal ψmid may have been due to the vertical translocation of water (‘hydraulic lift’). 5. Our results show that subsurface and surficial soil horizon development differentially affects the seasonal availability of water for desert plants. During wetter parts of the season, subsurface horizons limit the degree of water availability, while surface soil characteristics have the greatest influence on the effectiveness of summer precipitation. These findings suggest that the effectiveness of climatic precipitation and attendant plant utilization of water resources in warm desert systems may depend on the physical soil condition.

Effects of surface and sub-surface soil horizons on the seasonal performance of Larrea tridentata (creosotebush)

1. In warm arid and semiarid environments, the accumulation of clay minerals produces increasingly well developed soil horizons with the passage of time. Differences in the strength of development of two prominent soil horizons, silt- and clay-rich surface vesicular (Av), and clay-enriched subsurface argillic (Bt), may strongly influence the amount and seasonal continuity of plant-available water and the physiological activity of long-lived desert shrubs. Three sites were selected on an alluvial piedmont (bajada) in the Mojave Desert that varied in surface and subsurface horizon development. The first site, a deep deposit of stabilized dune sand, entirely lacked soil horizon development. The second site had a well developed surface stone pavement and underlying Av horizon, but lacked an argillic horizon in the sandy subsoil. The third had a well developed surface pavement and Av horizon, and a deeper, well developed clay-rich argillic horizon. Seasonal water potential and gas-exchange responses of the evergreen desert shrub Larrea tridentata[DC.] Cov., and volumetric soil water content (θ), were measured monthly in 1996 on these three soils in order to test the hypothesis that desert pavements, Av and subsurface Bt horizons differentially affect the effectiveness and utilization of seasonal precipitation. 2. Predawn and midday water potentials (ψpd and ψmid), net photosynthetic rates (Anet), and stomatal conductances (gs) in L. tridentata were highest in the deep, sandy dune soils lacking horizons that could restrict surface and subsurface infiltration. Plants growing in these soils also showed no physiological response to summer precipitation events. Following a single large precipitation event during the growing season (3·8 cm), the water potentials, Anet and gs in L. tridentata were similar in the first (sand dune) and second (pavement and Av horizon) sites. However, plant performance on these soil surfaces showed marked seasonal declines, and did not respond to a small pulse of summer rainfall. Plants growing at the third site (older soils with strongly developed pavement, Av and Bt horizons) had very low gas-exchange rates and water potentials. However, following convectional summer thunderstorms L. tridentata showed improved water relations and gas exchange in these soils. 3. Midday water potentials were frequently anomalously higher than predawn water potentials, up to +6 MPa late in the growing season, especially on soil surfaces with well developed soil horizons. This anomaly was due to seasonal decreases in ψpd accompanied by invariant midday ψ. In general, ψpd was correlated with θ across the depths measured. 4. Correlations between ψmid with θ at 35 cm increased dramatically with increasing Bt horizon development, suggesting that seasonal ψmid may have been due to the vertical translocation of water (‘hydraulic lift’). 5. Our results show that subsurface and surficial soil horizon development differentially affects the seasonal availability of water for desert plants. During wetter parts of the season, subsurface horizons limit the degree of water availability, while surface soil characteristics have the greatest influence on the effectiveness of summer precipitation. These findings suggest that the effectiveness of climatic precipitation and attendant plant utilization of water resources in warm desert systems may depend on the physical soil condition.

Plant regulation of soil nutrient distribution in the northern Chihuahuan Desert

Vegetation throughout the southwestern United States has changed from perennial grassland to woody shrubland over the past century. Previous studies on the development of ‘islands of fertility’ focused primarily on only the most limiting, plant-essential element, soil nitrogen (N). The research presented here addressed the question of whether other plant-essential elements, namely phosphorus (P) and potassium (K), showed similar concentration gradients under the desert shrub Larrea tridentata, creosotebush. It also examined whether the spatial distribution of N, P, and K differed from that of essential, but non-limiting nutrients, namely calcium (Ca), magnesium (Mg), and sulfur (S), and non-essential elements, namely sodium (Na), chloride (Cl), and fluoride (F). Within adjacent grassland and shrubland plots, surface soils were collected under and between vegetation and analyzed for a suite of soil nutrients. Soil nutrient distribution followed a uniform pattern that mirrored the spatial homogeneity of bunchgrasses in the grassland, but followed a patchy distribution that mirrored the spatial heterogeneity of individual shrubs in the shrubland. The main differences were that in the grassland, all elements were uniformly distributed, but in the shrubland the plant-essential elements, nitrogen, phosphorus, and potassium, were concentrated under the shrub canopy, and the non-limiting and non-essential elements were either concentrated in the intershrub spaces or were equally concentrated under shrubs and in the interspaces. Our results show how vegetation shifts from grassland to shrubland contribute to long-term, widespread change in the structure and function of desert ecosystems.

Freezing-induced xylem cavitation and the northern limit of Larrea tridentata.

We investigated the occurrence of freezing-induced cavitation in the evergreen desert shrub Larrea tridentata and compared it to co-occurring, winter-deciduous Prosopis velutina. Field measurements indicated that xylem sap in L. tridentata froze at temperatures below c. -5°C, and that this caused no measurable cavitation for minimum temperatures above -7°C. During the same period P. velutina cavitated almost completely. In the laboratory, we cooled stems of L. tridentata to temperatures ranging from -5 to -20°C, held them at temperature for 1 or 12 h, thawed the stems at a constant rate and measured cavitation by the decrease in hydraulic conductivity of stem segments. As observed in the field, freezing exotherms occurred at temperatures between -6.5 and -9°C and as long as temperatures were held above -11°C there was no change in hydraulic conductivity after thawing. However, when stems were cooled to between -11°C and -20°C, stem hydraulic conductivity decreased linearly with minimum temperature. Minimum temperatures between -16 and -20°C were sufficient to completely eliminate hydraulic conductance. Record (>20 year) minimum isotherms in this same range of temperatures corresponded closely with the northern limit of L. tridentata in the Mojave and Sonoran deserts.

Effects of plant size and water relations on gas exchange and growth of the desert shrub Larrea tridentata.

Larrea tridentata is a xerophytic evergreen shrub, dominant in the arid regions of the southwestern United States. We examined relationships between gasexchange characteristics, plant and soil water relations, and growth responses of large versus small shrubs of L. tridentata over the course of a summer growing season in the Chihuahuan Desert of southern New Mexico, USA. The soil wetting front did not reach 0.6 m, and soils at depths of 0.6 and 0.9 m remained dry throughout the summer, suggesting that L. tridentata extracts water largely from soil near the surface. Surface soil layers (<0.3 m) were drier under large plants, but predawn xylem water potentials were similar for both plant sizes suggesting some access to deeper soil moisture reserves by large plants. Stem elongation rates were about 40% less in large, reproductively active shrubs than in small, reproductively inactive shrubs. Maximal net photosynthetic rates (Pmax) occurred in early summer (21.3 μ mol m-2 s-1), when pre-dawn xylem water potential (XWP) reached ca. -1 MPa. Although both shrub sizes exhibited similar responses to environmental factors, small shrubs recovered faster from short-term drought, when pre-dawn XWP reached about -4.5 MPa and Pmax decreased to only ca. 20% of unstressed levels. Gas exchange measurements yielded a strong relationship between stomatal conductance and photosynthesis, and the relationship between leaf-to-air vapor pressure deficit and stomatal conductance was found to be influenced by pre-dawn XWP. Our results indicate that stomatal responses to water stress and vapor pressure deficit are important in determining rates of carbon gain and water loss in L. tridentata.

Effects of manipulation of water and nitrogen regime on the water relations of the desert shrub Larrea tridentata.

Water and nitrogen regimes of Larrea tridentata shrubs growing in the field were manipulated during an annual cycle. Patterns of leaf water status, leaf water relations characteristics, and stomatal behavior were followed concurrently. Large variations in leaf water status in both irrigated and nonirrigated individuals were observed. Predawn and midday leaf water potentials of nonirrigated shrubs were lowest except when measurements had been preceded by significant rainfall. Despite the large seasonal variation in leaf water status, reasonably constant, high levels of turgor were maintained. Pressure-volume curve analysis suggested that changes in the bulk leaf osmotic potential at full turgor were small and that nearly all of the turgor adjustment was due to tissue elastic adjustment. The increase in tissue elasticity with increasing water deficit manifested itself as a decrease in the relative water content at zero turgor and as a decrease in the tissue bulk elastic modulus. Because of large hydration-induced displacement in the osmotic potential and relative water content at zero turgor, it was necessary to use shoots in their natural state of hydration for pressure-volume curve determinations. Large diurnal and seasonal differences in maximum stomatal conductance were observed, but could not easily be attributed to variations in leaf water potential or leaf water relations characteristics such as the turgor loss point. The single factor which seemed to account for most of the diurnal and seasonal differences in maximum stomatal conductance between individual shrubs was an index of soil/root/ shoot hydraulic resistance. Daily maximum stomatal conductance was found to decrease with increasing soil/root/ shoot hydraulic resistance. This pattern was most consistent if the hydraulic resistance calculation was based on an estimate of total canopy transpiration rather than the more commonly used transpiration per unit leaf area. The reasons for this are discussed. It is suggested that while stomatal aperture necessarily represents a major physical resistance controlling transpiration, plant hydraulic resistance may represent the functional resistance through its effects on stomatal aperture.

Nutrient reabsorption efficiency and the response to phosphorus fertilization in the desert shrubLarrea tridentata (DC.) Cov.

A field fertilization experiment demonstrated that growth ofLarrea tridentata was not limited by phosphorus, even though soils contained high levels of pedogenic carbonates that can potentially fix high amounts of phosphorus. Nutrient reabsorption efficiencies in unfertilized shrubs ranged from 72–86% for P, making nutrient reabsorption a very effective nutrient conservation mechanism. Absolute amounts of N and P reabsorption increased with N and P concentrations in leaves, with reabsorption being greater during drought stress than during rapid leaf growth. However, only N reabsorption efficiency increased with increasing plant N status. A model was developed to explain patterns of nutrient reabsorption efficiencies over large gradients in nutrient availability.

A conceptual model for primary productivity, decomposition and nitrogen cycling in the Chihuahuan creosotebush desert.

The conceptual framework for a simulation model of primary productivity, decomposition and nitrogen cycling in a shrub-dominated desert ecosystem in southern New Mexico is presented. This model is based on our previous attempt to simulate carbon allocation patterns in the desert shrub Larrea tridentata Cov., which demonstrated that moisture patterns alone are insufficient to predict desert productivity. These results, as well as others, suggest that mineral nutrients, especially nitrogen, may also be an important determinant of productivity in arid environments. Our current research in the Chihuahuan desert is directed towards elucidating the numerous biotic and abiotic interactions that determine the rates and directions of carbon, nitrogen and water fluxes in this ecosystem. The development of this working model will serve as a tool to accomplish three major objectives: (1) to synthesize the large amount of existing data on decomposition and nitrogen cycling in deserts, (2) to quantify our present state of knowledge about the structure and function of ecosystem components important in carbon and nitrogen dynamics in deserts, and (3) to address hypotheses concerning the complex mechanisms of interactions and feedbacks among the organisms involved in carbon and nitrogen exchanges in deserts.

Are Regular Patterns in Desert Shrubs Artefacts of Sampling?

Ebert & McMaster (1981) show that, if shrubs which are within 4.2 m of one another are mistaken for single individuals, a random pattern of dots, representing the desert shrub Larrea tridentata (DC.) Coville at a density of 250 individuals ha-1, could be mistaken for a significantly regular pattern by the variance: mean ratio method. They suggest on this basis that the regular patterns detected in Larrea tridentata by Woodell, Mooney & Hill (1969) do not exist and argue that there is no convincing evidence for a regular pattern in any desert shrub. We comment as follows. (i) In the study of Woodell, Mooney & Hill (1969) individual plants of Larrea tridentata were easily distinguished in the field. Many of the plants were multi-stemmed. In the absence of conflicting evidence, however, Woodell and his co-workers assumed that the stems in each discrete group were, or had been, connected underground. Unless the connections were visible, they assumed that each separate clump of stems constituted an individual. Ebert & McMaster (1981) state that during early growth in Ambrosia (= Franseria) dumosa and Encelia farinosa, clumps are formed by the simultaneous germination of different genets. Evidence for this phenomenon in large clumps of Larrea tridentata is lacking. (ii) The main problem in studies of the distribution of Larrea tridentata individuals is its clonal growth pattern, which at some sites may make the definition of an individual arbitrary. Sternberg (1976) described it: 'The centre of a creosote bush dies and is replaced by new stems which eventually form new crowns peripheral to the original central crown. Over long periods of time with repeated cycles in which old stems die and new ones are formed peripherally, the creosote bush growth pattern results in a ring of satellite clumps around a circular to elliptic sterile centre.' The impact of this morphological pattern on pattern analyses is uncertain, especially at sites where the plants are not even-aged. When the variance: mean ratio method is used to detect non-random patterns, however, the clonal spread of L. tridentata seems likely to bias the ratio away from regularity. This is because when two groups of stems begin to appear distinct they will be close together. Larrea tridentata grows so slowly that most plants at most sites will never reach an age where the groups are widely separated. (iii) At the four sites with significant regular patterns of L. tridentata, the sizes and densities of the plants differed considerably (Table 1). At Inyokern, shrubs with their centres only 0. 5 m apart were easily distinguished. At Yuma, shrubs with their centres 2 m apart were distinguishable. (iv) Ebert & McMaster's (1971) simulation of the pattern analysis has two main deficiencies. First, plants as far apart as 4.2 m would, in some cases, be considered the same individual although the median maximum diameter of the shrubs ranged from 1 1 m to 2.0 m only at the four sites (Table 1). Secondly, Woodell, Mooney & Hill (1969) found a shrub density of 110 ha-1 at Yuma. Ebert & McMaster assumed that the shrub density at all sites was 250 ha-'. At lower densities a smaller proportion of shrubs will be confounded.

Growth Rates and Root: Shoot Ratios in Seedlings of the Desert Shrub Larrea tridentata

ABSTRACr.-Sonoran creosotebush seedlings are larger than plants of the Mojave, which are larger than Chihuahuan plants. However, total biomass, shoot biomass, and root biomass from each of three deserts increased at the same relative growth rate of 3% per day. The root:shoot biomass was found to be similar among plants of the three desert groups, but to vary between age classes. The ratio increased from 0.33 at 15 days to 0.49 at 75 days, and then decreased to 0.42 at 150 days. The roots were found to contain significantly less energy (4.1 cal * mg-') than shoots (4.7 cal * mg-'). Relative resource allocation to the root and shoot is no doubt of great importance in determining success of seedlings. This is especially true of desert xerophytes since germination and establishment must occur during brief periods of moisture availability. Went (1948) and Daubenmire (1959) asserted that xerophytic seedlings must allocate the major proportion of their structural carbon resources to the development of a large root system to keep ahead of progressively deeper soil desiccation. Yet remarkably little data relating to this assertion are availabe in the literature. Waisel (1962) demonstrated that the roots of xerophytes may elongate faster than the roots of mesophytes. However, Sankary (1971) and Birand (1961) have shown that rapid root elongation in seedlings does not necessarily imply a high root:shoot biomass ratio. Zohary (1961) showed that deep root systems are not always advantageous to xerophytes. Indeed, ecotypes growing on shallow rocky soil had shallower root systems than ecotypes growing on deep sandy soil. Barbour (1973), therefore, challenged the generalizations of Daubenmire and Went as being too simplistic. Creosotebush (Larrea tridentata) is the most successful perennial species of the hot deserts of North America as demonstrated by its widespread distribution and ubiquity in the deserts of the Southwest (Runyon, 1934), but with three distinct chromosomal races: diploids in the Chihuahuan Desert, tetraploids in the Sonoran Desert, and hexaploids in the Mojave Desert (Yang, 1967, 1978, 1970; Yang and Lowe, 1968: Barbour, 1969). If Went and Daubenmire are correct, creosotebush seedlings should show large increases in root:shoot biomass ratios with time, and possibly even a gradient of increasing root:shoot ratios accompanying the increase in aridity of the deserts. We studied growth rates and root:shoot biomass and energy ratios of populations from the three hot deserts during the first five months of growth to see if this was so.

Validation of a primary production model of the desert shrub Larrea tridentata using soil-moisture augmentation experiments.

In previous papers we have described and verified a primary production model of the desert shrub Larrea tridentata. Here we address the validation phase of the evaluation of this model. Two versions of the model which differ in the priority scheme used for allocating carbon to reproductive or vegetative organs were compared on the basis of their usefulness and reliability over a range of soil-moisture conditions. Over an entire growing season when soil-moisture conditions were near "normal" both versions of the model were adequate predictors of total above-ground vegetative growth and one was an adequate predictor of reproductive growth as well. A more detailed analysis revealed that the versions varied in the range of soil-moisture conditions over which they were adequate and that neither was adequate when soil-moisture had remained high for extended periods. The validation process has revealed some likely areas for model improvement to increase adequacy.

Energy Relationships of the Mammals of a Desert Shrub (Larrea tridentata) Community

A year's study was made of 13 species of mammals in a desert community in southeastern Arizona. Mammal density averaged 17.4/ha: 66% Dipodomys merriami and 10.5% Onychomys torridus. Average biomass was 1130 g/ha: 40% D. merriami and 40% Lepus californicus. Annual energy flow of mammals was 105,950 kcal/ha: 55% by a granivore (D. merriami), 22% by a browser (L. californicus) and 6.5% by an insectivore (O. torridus). 94.6% of the energy flow was spent in maintenance and 5.4% in growth. The secondary productivity of the dominant D. merriami was 1.2% of its energy flow; that of Peromyscus eremicus, the resident species with the lowest and least stable density, was 1.7%. A dominant species may be more important in the cycling of matter in the community, while nondominants may be more important in stabilizing the community and sustaining the higher trophic levels. Since the mammals dissipated only 1.95% of the net annual above ground plant production, their importance must be in their controlling actions on the...