Delayed germination and dispersal in desert annuals: Escape in space and time

Summary A model is developed to consider the interplay between dispersibility and delayed germination in desert annuals. The model explores the effect of low levels of dispersal, considered realistic for annual plants, on optimal germination fraction. The model also demonstrates the effect of the amount and accuracy of predictive (responsive to the environment) dormancy on the optimal innate germination fraction (not responsive to environmental conditions).Optimal germination fraction is found to be very sensitive to changes in despersibility especially at the limited dispersibilities that are realistic for annual plants. As dispersibility increases, optimal germination fraction increases. If plants make two kinds of seeds with differing despersibility, reproduction is maximized if the low dispersal seeds have delayed germination and the high dispersal seeds have quick germination. If dormancy mechanisms permit seeds to germinate when environmental conditions allow successful maturation, and remain dormant when environmental conditions do not permit successful maturation, what fraction of seeds should remain dormant under predicted good conditions as a hedge against inaccurate prediction of the environment? If environmental cues that break dormancy are uncorrelated with environmental conditions that permit successful maturation, predictive dormancy has little or no effect on the optimal innate germination fraction. When predictive dormancy lowers the probability of germinating when environmental conditions preclude successful maturation, the optimal innate germination fraction increases with increasing germination control by predictive dormancy. With a moderate degree of germination control by predictive dormancy, the optimal innate dormancy is still sensitive to changes in dispersal in the low dispersal ranges characteristic of annual plants.Evidence is presented from plant species that have both dispersal and germination dimorphisms to support the predicted correlation of high germination fractions with high dispersal.     

Using upper boundary constraints to quantify competitive response of desert annuals

Using a target-neighborhood approach with six annual dicot species in the Negev Desert of Israel, we tested whether neighborhood biomass constrained the upper limit of plant performance and if the slope of the upper boundary was correlated to species trait means, such as relative growth rate (RGR) and seed mass. Target individuals were measured in early spring and then collected at the onset of the dry season along with all naturally occurring neighbors within a 5-cm radius of the target. Using a minimum of 50 samples for each of the six target species, we found no significant relationships between target performance and either density or aboveground biomass of neighbors, when including all target individuals in the regressions. However, aboveground neighbor biomass did influence the maximum potential size of targets. When regressions were restricted to the largest target plant within classes of neighbor biomass, significant negative relationships were found for all six species, with neighbor biomass explaining 55 to 94% of the variance in maximum target biomass. The slopes of the regressions were used as an index of competitive response. The correlations between competitive response and estimates of species traits were not significant with the possible exception that species with lower RGRmax may be better response competitors (i.e. were less sensitive to competition). These findings indicate the usefulness of the boundary regression technique for describing competitive interactions among neighbors, particularly in low productivity environments.     

Ecophysiology of two solar tracking desert winter annuals

Summary The seasonal course of water relations in field populations of two leaf solar tracking desert winter annuals was examined. Measurements were made of leaf movements in relation to leaf conductance and water potential. Malvastrum rotundifolium maintained solar tracking movements up to the wilting point of the plant (-4 MPa). Lupinus arizonicus altered its morphology through paraheliotropic leaf movements as leaf water potentials declined to-1.8 MPa. Diurnal patterns of leaf conductance showed marked seasonal trends, with gas exchange activity being restricted to early morning hours as water availability declined. Studies of potted plants showed that L. arizonicus was not able to alter its osmotic potential in response to drought, while M. rotundifolium underwent a 1.86 MPa reduction in osmotic potential. The significance of the two contrasting patterns is discussed in terms of observed plant distribution and origin.     

Density-dependent germination response by seeds of desert annuals

Summary Field observations and experiments indicate that the presence of seedlings at high densities inhibits subsequent germination of desert annuals. Since plants growing at high densities face severe competition for limited resources, this response by seeds is interpreted as an adaptation to avoid an unfavorable competitive climate where growth and survivorship are likely to be low.     

Photosynthetic capacity in relation to leaf position in desert versus old-field annuals

Summary Desert annuals of Death Valley, California have higher average light-saturated photosynthetic capacities and leaf nitrogen contents than do early-successional annuals of Illinois. The leaves of annuals in the light-unlimited Death Valley environment change little in specific weight, nitrogen, or photosynthetic capacity with age. In contrast, these properties decrease markedly with age in the leaves of the Illinois annuals even in leaves not exposed to the usual shading that accompanies canopy development. These results are interpreted in a carbon-gained-per-nitrogen-invested context.This study is dedicated to Professor M. Evenari for his continuing strong contributions to desert ecology     

Phenotypical temperature adaptation of protein turnover in desert annuals

Protein synthesis and protein degradation rates were measured in three desert annual species at four different experimental temperatures. The taxa chosen for this study were the C₃ winter annuals, Bowlesia incana Ruiz & Pavon and Plantago insularis Eastw., and a C₄ summer annual, Atriplex elegans (Moq.) D. Dietr. Peak rates of protein synthesis correlated well with the preferred habitat temperatures of B. incana and A. elegans; optima occurred at 25 and 35°C, respectively. Plants of P. insularis showed an optimum protein synthesis rate at 35°C; however, this optimum rate was considerably lower than for the other two species. Higher activation energies for protein synthesis tended to parallel adaptation to higher temperature habitats. Responses of protein degradation to temperature in A. elegans and B. incana were consistent with their natural thermal regimes, when evaluated for the transition from 25 to 35°C. Again, protein degradation in P. insularis shows an intermediate response to temperature during the 25 to 35°C transition.     

Spatial patterns of desert annuals in relation to shrub effects on soil moisture

Questions: What are the effects of a shrub (Haloxylon ammodendron) on spatial patterns of soil moisture in different seasons? How does productivity of understorey annuals respond to these effects? Are such effects always positive for annuals under shrubs? Location: South Gurbantunggut Desert, northwest China. Methods: Using geostatistics, we explored seasonal patterns of topsoil moisture in a 12 × 9‐m plot over the growing season. To determine spatial patterns of understorey annuals in response to H. ammodendron presence, biomass of annuals was recorded in four 0.2 × 5.0‐m transects from the centre of a shrub to the space between shrubs (interspace). We also investigated vertical distribution of root biomass for annuals and soil moisture dynamics across soil profiles in shrub‐canopied areas and interspaces. Results: Topsoil moisture changed from autocorrelation in the wet spring to random structure in the dry season, while soil moisture below 20 cm was higher in shrub‐canopied areas. Across all microhabitats, soil moisture in upper soil layers was higher than in deeper soil layers during the spring wet season, but lower during summer drought. Topsoil was close to air‐dry during the dry season and developed a ‘dry sand layer’ that reduced evaporative loss of soil water from deeper layers recharged by snowmelt in spring. Aboveground biomass of understorey annuals was lowest adjacent to shrub stems and peaked at the shrub margin, forming a ‘ring’ of high herbaceous productivity surrounding individual shrubs. To acclimate to drier conditions, annuals in interspaces invested more root biomass in deeper soil with a root/shoot ratio (R/S) twice that in canopied areas. Conclusions: Positive and negative effects of shrubs on understorey plants in arid ecosystems are commonly related to nature of the environmental stress and tested species. Our results suggest there is also microhabitat‐dependence in the Gurbantunggut Desert. Soil water under H. ammodendron is seasonally enriched in topsoil and deeper layers. Understorey annuals respond to the effect of shrubs on soil water availability with lower R/S and less root biomass in deeper soil layers and develop a ‘ring’ of high productivity at the shrub patch margin where positive and negative effects of shrubs are balanced.     

Growth and abundance of desert annuals in an arid woodland in Oman

Given the role of Scaevola plumieri as a major pioneer species in the dune environment, as well as the need forindigenous sand stabilisers for South Africa’s coastal dunes, the reproductive phenology was unravelled as a firststep to establish the potential usefulness of the species in dune stabilisation programmes. In the highly variablecoastal environment the plants must cope with many problems for successful reproduction to take place. Theplants of S. plumieri overcome these difficulties through vegetative reproduction, but sexual reproduction is byno means insignificant. Numerous peduncles, each carrying multiple buds, were produced per stem, but only fewdeveloped into ripe seeds. Several of the flowers were unfertilised and of the seeds produced many were infectedand/or aborted. There was a strong effect of dune position on the reproductive phenology of S. plumieri. Thestems situated at the landward face of the foredunes showed an overall higher reproductive performance numberof buds, flowers, unripe seeds, ripe seeds. The phenogram showed that the reproduction of stems at the landwardside of the foredunes started on average 34 days earlier and showed a tendency for a longer reproductionperiod. For the duration of the different stages in the reproduction sequence, the bud stage’ took the shortesttime and the ‘ripe seed stage’ longest time. Between the different sample years, the production of the number ofpeduncles, buds, flowers and seeds was the similar; only differences in reproduction time (duration) of the differentstages were observed for the bud, flower an unripe seeds stage between the sample years.     

Effect of protection on the nutrient concentration and uptake of some Mediterranean desert annuals

The present study includes an evaluation of the effect of protection of a grazing ecosystem on the nutrient concentration and uptake in the organs of four annual species of non-saline depressions in the western Mediterranean regionof Egypt.The concentration of macro- and micronutrients in different organs became lower due to the increase in biomass of individuals with protection. The maximum uptake of nutrients occurred during the early vegetative stage, except for a few cases in which the uptake in some species reached two peaks, one in the early vegetative stage and the other in the reproductive stage.The efficiency of the selected annual species in uptaking and accumulating nutrients was compared with rates of uptake of nutrients and amounts of nutrients accumulated in tissues of two species representing other life forms. The annuals and perennial herbs (ephemeroids) are in general more efficient in uptaking and accumulating nutrients in their tissues than perennial shrubs.     

Nurse‐plant effects on the seed biology and germination of desert annuals

Nurse‐plants generally have positive effects on understorey species by creating more suitable conditions for stress‐intolerant plants relative to open micro‐habitats. However, long‐term effects of this plant–plant facilitation system have been rarely examined. Seeds of five desert annual species from Atiquipa coastal desert in Southern Peru were used to examine whether different microenvironmental conditions under the nurse‐plants Caesalpinia spinosa Molina (Kuntze) lead to differences in seed biology and germinability of annual plants relative to open, canopy‐free conditions. Seeds collected from plants associated with nurse‐plants were predicted to be (i) larger due to more favourable growing conditions, (ii) more viable and with greater germination rates, (iii) less variable in size and viability due to reduced environmental heterogeneity, and (iv) to germinate faster to avoid apparent competition with other annuals. Seed attribute measurements and germination trials in growth chambers were used to test these predictions. Although the plant abundance of only 2 of 5 species was strongly facilitated by the nurse‐plant, no significant differences were found in seed mass, viability or relative variability between understorey and open micro‐habitats for any of the species. Contrary to our predictions, final seed germination rates of seeds from open micro‐habitats were higher, and the open micro‐habitat treatment was more favourable for germination of seeds from both open and understorey environments. Taken together, these results suggest that plant–plant facilitation does not necessarily affect seed biology traits. Further studies addressing larger distribution ranges and/or density gradients of understorey species will illuminate the potential evolutionary effects of nurse‐plants.     

Modelling the multidimensional niche by linking functional traits to competitive performance

Linking competitive outcomes to environmental conditions is necessary for understanding species'' distributions and responses to environmental change. Despite this importance, generalizable approaches for predicting competitive outcomes across abiotic gradients are lacking, driven largely by the highly complex and context-dependent nature of biotic interactions. Here, we present and empirically test a novel niche model that uses functional traits to model the niche space of organisms and predict competitive outcomes of co-occurring populations across multiple resource gradients. The model makes no assumptions about the underlying mode of competition and instead applies to those settings where relative competitive ability across environments correlates with a quantifiable performance metric. To test the model, a series of controlled microcosm experiments were conducted using genetically related strains of a widespread microbe. The model identified trait microevolution and performance differences among strains, with the predicted competitive ability of each organism mapped across a two-dimensional carbon and nitrogen resource space. Areas of coexistence and competitive dominance between strains were identified, and the predicted competitive outcomes were validated in approximately 95% of the pairings. By linking trait variation to competitive ability, our work demonstrates a generalizable approach for predicting and modelling competitive outcomes across changing environmental contexts.     

Spatial pattern analysis in Namaqualand desert plant communities: evidence for general positive interactions

This paper investigates of the spatial arrangement of individual plants in mapped plots in two desert communities in the winter rainfall region of South Africa. In both communities there was a very strong tendency towards clumped patterns when all plants were considered together. There was also a predominance of clumped patterns when the most abundant species in both communities were considered individually. When the arrangement of the most abundant species was considered relative to the arrangement of all other individuals at the within-clump scale, there was a high frequency of positive associations in both communities (62% and 65%). We speculate that these patterns represent a combination of seed dispersal strategies that favour clumped patterns and a predominance of positive interactions between plants in both of the communities. When specific pairwise associations between the most abundant species were considered at the within-clump scale, differences were apparent between the two communities. In the short strandveld community neutral associations predominated, while in the medium strandveld, neutral and positive association accounted for equal proportions of the associations. This between-plot difference was also apparent when the volumes of plants were related to an index of neighbourhood competitiveness. In the short strandveld there were no significant relationships while in the medium strandveld there were some weak (but significant) relationships. These differences were not altogether unexpected. If we assume that plants in the medium strandveld are generally longer-lived, then interactions between plants are likely to develop over a longer time and, therefore, are likely to be stronger.     

Suppression of annuals by Tribulus terrestris in an abandoned field in the sandy desert of Kuwait

In an abandoned field in the sandy desert of Kuwait annual plants were less numerous in stands dominated by Tribulus terrestris than in adjacent stands dominated by other species. In the Tribulus stands, annuals were smaller in phytomass and stature. Possible physical and biotic factors were investigated in both types of stands in order to determine the causative agent(s) for the lesser density and development of annuals in the Tribulus stands. Physical conditions of moisture, light and soil characteristics did not appear to be limiting to annuals in the Tribulus stands. Experimental evidence is presented for the leaching from T. terrestris shoots of water-soluble substances which strongly inhibit the germination and radicle elongation of most of the associated annual species. The effect of these substances is fairly specific. Preliminary analysis indicates the presence of phenolic compounds in the leachate of T. terrestris, which are believed to play a significant role in the growth inhibition. The possible role of chemical inhibition on certain annuals in stands of T. terrestris is discussed.     

柴达木盆地东部荒漠植物生态位特征

以柴达木盆地东部27个典型样地的植物群落为研究对象,通过生态位宽度和生态位重叠指数计算,分析该区13种优势植物在土壤含水量、容重、有机质、pH、全盐、全氮维上的生态位特征,以期为柴达木盆地植被恢复重建和荒漠系统生态保护提供理论参考。结果表明:①13种植物在6个土壤因子上平均生态位宽度从大到小依次是:驼绒藜、琵琶柴、芨芨草、芦苇、合头草、蒿叶猪毛菜、白刺、盐爪爪、小嵩草、麻黄、沙拐枣、柽柳和梭梭,其中,驼绒藜在土壤含水量、容重和有机质上占据最高值,琵琶柴在土壤pH、全盐、全氮维上占据最高值;②生态位宽度较大的物种在生态位重叠上并不占据最高重叠值,而是处于一个中游的位置;③驼绒藜和琵琶柴在生态位宽度和生态位重叠上均占相对高的位置,对群落的建群起重要作用;④78个种对在每一个土壤因子上的生态位重叠值0.3的占总对数的比例为76%,柴达木盆地东部植物总体生态位重叠偏低,竞争较小,群落处于一个相对稳定的状态。