The impact of shade on establishment of shrubs adapted to the high light irradiation of semi-arid environments

Light may be an important limiting resource that influences community structure of chenopod shrublands. As part of a larger study that aimed to determine the factors that influence chenopod community structure, the focus of this study was the influence of plant canopy on the growth and establishment of smaller plants. We therefore measured the height and cover of three chenopods (Enchylaena tomentosa, Maireana brevifolia and Maireana georgei) when growing within and outside of the canopy of Atriplex bunburyana under field conditions. All three chenopods had lower cover and E. tomentosa was taller when growing within the canopy of A. bunburyana in comparison to those growing outside of the canopy. The chenopods were then grown under three artificial shade regimes. Plant height, cover, biomass, relative leaf area and photosynthetic surface area measurements showed that each species responded differently to shade. E. tomentosa biomass was facilitated by shade. This was inferred by an increase in total plant biomass. M. brevifolia, in contrast, tolerated shade by increasing above-ground biomass allocation. M. georgei was adversely affected by the shade regimes: root biomass decreased in response to shade. Competition for light is, therefore, likely to influence chenopod community structure of semi-arid and arid environments.     

Phenology ofMedicago minimaandErodium cicutariumin semi-arid Argentina

The phenology ofMedicago minimavar.minimaandErodium cicutariumwas studied at two different field sites in the Calden District (approx. 10 million ha), a temperate semi-arid phytogeographical region in central Argentina. One site had been protected from wild and domestic herbivores for the 6 previous years. The other adjacent site had been exposed to continuous grazing by cattle for several decades. A phenological key was made for each species in 1988, and this key was improved during the 1989 growing season when phenological observations were made every 1–3 weeks.The growth cycle ofM. minimaandE. cicutariumwas similar at the two studied sites. Initiation of the cycle occurred in autumn and appeared to be associated with soil water availability. Phenological patterns were very dynamic at the reproductive stage in both species. This stage started earlier and was faster inE. cicutariumthan inM. minima. The end of the growing season occurred during late spring, concomitant with high maximum air temperatures. Our results suggest that these species hasten their development when air temperature increases and soil water availability decreases. This could be an important strategy in these species which allows them to persist as seeds, and produce a new generation under favourable environmental conditions.     

Catastrophic drought-induced die-off of perennial chenopod shrubs in arid Australia following intensive cattle browsing

Cover and survivorship of perennial chenopods, Atriplex vesicaria and Maireana astrotricha, declined markedly during a prolonged dry period at Roxby Downs in arid South Australia. Despite their resilience to browsing in favourable seasons, only 2% of the A. vesicaria shrubs browsed heavily by cattle survived the drought. M. astrotricha exhibited greater drought survivorship, although the post-drought cover retention in browsed shrubs was only half that of unbrowsed controls. Survivorship was highest for unbrowsed chenopods growing in moisture-enhanced run-on or dune-base environments. Maintenance of these patchy areas with reduced water-stress is hence important for the persistence of browsed chenopod shrublands. Light browsing by either European Rabbits (Oryctolagus cuniculus) or Greater Stick-nest Rats (Leporillus conditor) had no measurable impact upon chenopod survivorship.     

Microfungal community associated with rhizosphere soil of Zygophyllum qatarense in arid habitats of Bahrain

Population density and species diversity of microfungal communities were investigated in the rhizosphere soil of the halophytic plant Zygophyllum qatarense inhabiting saline and nonsaline habitats of the arid desert environment of Bahrain. Unlike the nonsaline habitat which is situated in the physiographic zone of multiple escarpment and backslopes, the saline site is located in the coastal lowlands and is featured by high chloride content, electrical conductivity, total soluble salts and low organic matter. Soils of both habitats are sandy, slightly alkaline, poor in nutrient sources, low in water-holding capacity and mainly dominated by a salt-tolerant flora. Quantification of data for the recovery of fungi were based on colony identification and counts by a series of ten-fold dilutions plate method, using various natural, synthetic and selective media. A total of 2780 isolates, fluctuating between 25 and 1109 per sample, were recovered during the present study among all habitats, seasons and plant sizes. Grouping of these isolates has resulted in a maximum of 28 fungal taxa varied between 5 and 15 species, of which 24 were hyphomycetes, 3 ascomycetes and one was an unknown species. All the recorded species in this study, excluding the genus Fusarium, are newly reported from the arid terrestrial habitats of Bahrain. Of the encountered fungi,Cladosporium sphaerospermum was the most dominant and frequent genus, among all plant sizes, followed by Penicillium citrinam and Aspergillus fumigatus, a finding with consistent documented data from similar arid Sahara ecosystems. Examination of data, supported by analysis of relative density values, percentage recovery rates, polar ordination and diversity indices revealed that the nonsaline habitat during the dry period yielded the highest isolate frequency, species abundance, and diversity when compared with the saline habitat. Moreover, a progressive increase in colony occurrence and species diversity was equivalently associated with increment in plant size in the nonsaline habitat. Apparently, the vast majority (moderate to low occurrence class) of the reported species are rhizosphere indigenous saprophytic cellulose-decomposers, whilst the sparse taxa (high occurrence class), e.g. C. sphaerospermum, are regarded as thermo-osmotolerant. Comparison of species richness among samples suggests that small plants inhabiting nonsaline habitats during the wet and dry season were richest in species composition. It is proposed that inter- and intra-specific variation in fungal community between the above habitats reflect not only the influence of plant age and season but also extends to critical multi-soil edaphic and biotic factors involving essentially soil moisture, salinity and root growth dynamic and exudates.     

Growth response of the saltbush Atriplex nummularia L. to inoculation with the arbuscular mycorrhizal fungus Glomus intraradices

Plantlets of Atriplex nummularia were inoculated with the arbuscular mycorrhizal fungi Glomus intraradices in a pot experiment. Plants were grown in a low P soil. Highly significant growth response of a Chenopodiaceae was recorded for the first time. Mycorrhizal colonization of roots was well developed, internal hyphae and vesicles were observed, but not arbuscules. These observations suggest that arbuscules are not necessary to obtain significant growth stimulation from mycorrhizal inoculation. Atriplex nummularia is already used as forage crops, its high mycorrhizal dependency offers possibilities to develop this production and revegetation strategies.     

Short-term effects of shrub control on two different plant communities in Argentina

The aim of this work was to study the effect of controlling jarilla (Larreaspp.), nonforage shrubs for livestock, on two plant communities with different forage production in the arid mid-west of Argentina. Total vegetation cover, forage species cover and carrying capacity were determined during three growing seasons, for cleared and uncleared plots. At the end of the study, significant differences were found in theLarrea cuneifoliacommunity, where average forage species cover increased by 156% over the control, and greater carrying capacity (129%) was also obtained from the cleared treatment. No significant differences were found in theLarrea divaricatacommunity. Data lead us to recommend shrub control as an improvement practice only in areas with low forage species cover and poor carrying capacity.     

Ecophysiological studies inHesperaloe funifera(Agavaceae): a potential new CAM crop. The effect of light availability on photosynthesis and growth

Hesperaloe funifera(Agavaceae) is being investigated as a new specialty-fiber crop for arid lands. The objective of this study was to examine the effects of reduced solar radiation on photosynthesis and growth in this CAM species.Peak CO₂uptake integrated through 24 h was found to be highest during the fall. Although shading greatly reduced CO₂uptake in fall and winter, the decrease was not statistically significant for the spring and summer. Total biomass was significantly different for the three light treatments, with average fresh weights of 5243 g, 4488 g, and 3671 g for the full-sun, 53%-sun, and 20%-sun treatments, respectively. The greatest differences were found in the crown growth, which would affect future production of both leaves and flower stalks.Photosynthesis and growth results suggest that the cultivation ofHesperaloe funiferawill be favored in well-lit environments characteristic of arid or semi-arid climates.     

Annual growth rings of the shrub Anarthrophyllum rigidum across Patagonia: Interannual variations and relationships with climate

Annual growth rings are well-defined in some shrubby species distributed along the Patagonian steppe and provide useful information to identify the environmental factors that influence the radial growth of shrubs in the region. However, little is known about variations in ring width from these species and their relationships with local environment. In this paper we present 18 growth ring width chronologies covering the last 2–3 decades from the shrub Anarthrophyllum rigidum (Fabaceae) encompassing most of its natural range of distribution in Argentina. Interannual variations in growth were first analyzed collectively to identify common regional patterns. In addition, shrub growth at each site was compared with local climate records and site-specific characteristics. Based on a principal component analysis (PCA), sites were joined in four major groups related to the site proximity to the Andes (foothills versus Patagonian plateau) and latitude (north versus south Patagonia). The relatively similar percentage of PCA variance associated with each group reflects environmental differences among sites. Consistent with this observation, we noted large variability in the responses of the A. rigidum woody growth to regional climatic fluctuations. At most sites, this species is favored by abundant winter precipitation, warmer spring and autumn temperatures (controlling the duration of the growing season), cool summers, or a combination of these factors. However, deviations from these patterns are also recorded. Our study represents the first large-scale dendroclimatological analysis of a shrubby species in southern South America. The results presented here provide information on the main patterns and factors affecting the growth of A. rigidum in Patagonia, and may be useful for determining well-suited practices for management and conservation of this ubiquitous species.     

Use of large Acacia trees by the cavity dwelling Black-tailed Tree Rat in the Southern Kalahari

Recent extensive harvesting of large, often dead Acacia trees in arid savanna of southern Africa is cause for concern about the conservation status of the arid savanna and its animal community. We mapped vegetation and nests of the Black-tailed Tree Rat Thallomys nigricauda to assess the extent to which the rats depend on particular tree species and on the existence of dead, standing trees. The study was conducted in continuous Acacia woodland on the southern and eastern edge of the Kalahari, South Africa. Trees in which there were tree rat nests were compared with trees of similar size and vigour to identify the characteristics of nest sites. Spatial analysis of tree rat distribution was conducted using Ripley's-L function. We found that T. nigricauda was able to utilize all available tree species, as long as trees were large and old enough so that cavities were existing inside the stem. The spatial distribution of nest trees did not show clumping at the investigated scale, and we therefore reject the notion of the rats forming colonies when inhabiting continuous woodlands. The selection of a particular tree as a nest site was furthermore depending on the close proximity of the major food plant, Acacia mellifera. This may limit the choice of suitable nest sites, since A. mellifera was less likely to grow within a vegetation patch containing a large trees than in patches without large trees.     

Refuge selection by two sympatric species of arid-dwelling land snails: Different adaptive strategies to achieve the same objective

Terrestrial gastropods, especially those dwelling in dry habitats such as the Mediterranean, are highly susceptible to dehydration. In this study, the use of refuges by Iberus gualtieranus gualtieranus and Sphincterochila candidissima is compared. These two species belong to families with different distribution ranges, but they live in sympatry on a mountain in southern Spain with a dry Mediterranean climate. I. g. gualtieranus used mainly rock crevices as refuges, consistently throughout the year. S. candidissima hibernated beneath humus during autumn and winter (I. g. gualtieranus did not hibernate). In spring, S. candidissima seemed active, while I. g. gualtieranus was already aestivating. In summer, S. candidissima aestivated on vegetation, while I. gualtieranus aestivated deep within crevices. These differences in the use of refuges may be explained on the basis of the need for more protection by I. g. gualtieranus because of its morphology, which is less suited to the dry habitat, but has evolved to enable this species to exploit karstic crevices as refuges more efficiently. These results illustrate that behaviour and morphology interact allowing both species to coexist in sympatry in an arid environment using different adaptive strategies.     

The effects of seasonal changes to in-stream vegetation cover on patterns of flow and accumulation of sediment

In-stream macrophytes are typically abundant in nutrient-rich chalk streams during the spring and summer months and modify the in-stream environment by altering river flows and trapping sediments. We present results from an inter-disciplinary study of two river reaches in the River Frome catchment, Dorset (UK). The investigation focused on how Ranunculus (water crowfoot), the dominant submerged macrophyte in the study reaches, modified patterns of flow and sediment deposition. Measurements were taken on a monthly basis throughout 2003 to determine seasonal patterns in macrophyte cover, associated changes in the distributions of flow velocities and the character and amount of accumulated fine sediment within stands of Ranunculus.Maximum in-stream cover of macrophytes exceeded 70% at both sites. Flow velocities were less than 0.1 m s^− 1 within the stands of Ranunculus and accelerated to 0.8 m s^− 1 outside the stands. During the early stages of the growth of Ranunculus, fine sediment mostly accumulated within the upstream section of the plant but the area of fine sediment accumulation extended into the downstream trailing section of the plant later in the growing season. The fine sediment accumulations were dominated by sand (63–1000 μm) with silts and clays (0.37–63 μm) comprising < 10% by volume. The content of organic matter in the accumulated sediments varied within stands, between reaches and through the growing season with values ranging between 9 and 105 mg g^− 1 dry weight. At the reach scale the two sites exhibited different growth forms of Ranunculus which created distinctive patterns of flow and fine sediment deposition.     

Distribution and Characteristics of Nitraria sphaerocarpa nebkhas in a Gobi habitat outside an oasis in Hexi Corridor region, China

Nitraria sphaerocarpa Maxim., a typical desert plant which often appears in nebkha form, is widely distributed along the periphery of oasis in arid regions of Northwest China, and plays an important role in desertification control around oases. In this paper, the characteristics and distribution patterns of N. sphaerocarpa nebkhas were investigated in a Gobi environment outside an oasis. Results shows that about 200 N. sphaerocarpa nebkhas per hectare was recorded and total sand sediments was 25 m³/ha. Nebkhas vary in size, and the mean height (H) and horizontal composition (L) were 15 cm and 80 cm, respectively. There exists a significant linear relationship between H and L, suggesting that the investigated nebkhas are mostly in the growth stage. The numbers, height and sand deposit of nebkhas decreased with increasing distance away from the oasis. Grain-size analysis indicates that the mean grain size of sediments at the crests of nebkhas increased with increasing nebkha height. Organic C and total N concentrations were related to the mean grain size. It was concluded that N. sphaerocarpa shrubs are efficient horizontal sand traps for wind-driven sediments. A better understanding of the development of nebkha can provide information for the design of biological sand control measures around oases in arid regions.     

Nutrient status of Populus euphratica growing in desert riparian forests of northwestern China

Seasonal and microhabitat variations of chemical constituents of foliar organic carbon (C), total nitrogen (N), total phosphorus (P), and total potassium (K), in Populus euphratica growing in desert riparian forests in northwestern China and their correlations were studied. Results show that ranges of C, N, P and K contents in the leaves of P. euphratica were 39.08%?46.16%, 0.28%?2.81%, 0.05%?0.18% and 0.35%?2.03%, with means of 43.51%, 1.49%, 0.102% and 1.17%, respectively. The ratio of C/N, C/P and N/P changed from 16.26 to 146.61, from 258.08 to 908.67 and from 2.89 to 26.67; the mean was 37.24, 466.27 and 15.14, respectively. The mean N content was significantly lower than of deciduous trees in China, but the mean P content was nearly equivalent. The ratio of C/N was remarkably higher than of global land plants. The ratio of N/P indicated that growth of P. euphratica was jointly limited by N and P nutrient deficiency. During the growth season, total trends of leaf C, N, P and K contents decreased. The maximum appeared in May, and the minimum in September. Among microhabitats, C, N and K contents gradually increased from riparian lowland, flatland, sandpile, Gobi and dune, but C/N ratio was opposite, and P content was not apparent. Foliar C content was extremely, significantly and positively correlated with N and K contents, respectively. The relationships of N-K and P-K were both significantly positive.     

The distribution of rare phenolic type compounds in wild and cultivated wheats

In 14 species of Aegilops, four species of Triticum and two species of Hordeum, a complex of rare phenolic compounds was found and showed quantitative differences between the wild and domestic species. There was a dramatic reduction in the cultivars to about 5% of the amount of these phenolic constituents. In the wild species these phenolic compounds may act as important germination regulators, such as water-dissolving germination inhibitors which may act as 'rain gauges'. This is especially important for the germination at the appropriate time in species inhabiting arid regions or deserts.     

Leaf nitrogen productivity as a mechanism driving the success of invasive annual grasses under low and high nitrogen supply

Invasion of the historically perennial-dominated landscapes in the Great Basin by exotic winter annual grasses is one of the most serious plant invasions in North America. Evidence suggests invasive annuals outperform native perennials under N-poor and N-rich conditions. The objective of this study was to identify key traits contributing to the success of invasive annual grasses in these environments. Three invasive annual grasses, two native perennial grasses and one introduced perennial grass were exposed to three levels of N supply. Root biomass, root length, root N uptake rate, root and leaf morphology, leaf nitrogen productivity (leaf NP) as well as biomass and N allocation were quantified over four harvests. Path analysis indicated that leaf NP was the key trait contributing to variation in N capture among the species. Species with a higher leaf NP produced more root length and consequently captured more N under a range of soil N availability. This suggests variation in leaf NP may be one critical trait determining the ability of the resident plant community to resist establishment of these invaders. Restoration programs may be able to increase weed resistance by specifically selecting for this trait in revegetation efforts.     

Water shortage and associated changes in organic nitrogen betweenPappophorum caespitosum(Gramineae) provenances

Three provenances ofPappophorum caespitosum, a forage native grass from the temperate arid region of Argentina, were studied to gain a physiological knowledge of the seasonal organic N status in response to adequate water supply and water stress conditions; and to assess the extent to which variations in the N status between these provenances arise under different water regimes. Plant part samples were analysed for free and bound organic N fractions. Water shortage resulted in a significant depression of plant growth in all provenances. Total N concentration declined along the annual cycle in both water treatments, but it was higher under water stress. The size of the free N pool was very large, suggesting that it may act like an osmoticum. Nitrogen use efficiency increases along the growing period, while the rate of N intake and the specific leaf N content decrease. Variability was found between provenances in most of the physiological and biochemical traits examined under both water regimes.     

Comparison of soil bacterial communities associated with actinorhizal, non-actinorhizal plants and the interspaces in the sclerophyllous matorral from Central Chile in two different seasons

The spatial heterogeneity of resources in desert and semi-arid shrubland appears to be important in determining higher soil bacteria abundance around plants than in soil without plant cover. Thus, these bacterial communities could be important contributors to nutrient cycling in arid ecosystems. Bacterial diversity from Chilean sclerophyllous matorral was determined by Terminal Restriction Fragment Length Polymorphism (T-RFLP). Soil samples associated with the actinorhizal plant Colletia hystrix, non-actinorhizal plants and interspace soil without plant cover, were collected in May and October. The non-actinorhizal and interspace soil differed significantly in their potassium content in May and pH in October. The T-RFLP analysis revealed differences in the bacterial community structure from the different habitats. The soil bacterial communities associated with plants were the most similar, whereas the interspace soil community differed in both sampling times. The factors that best explained the groupings were potassium and pH. The greatest diversity was observed in the interspace soil. The Microbial Community Analysis showed a significant proportion of T-RFs identified as Firmicutes and Proteobacteria. Likewise, spatial and temporal differences were observed in the main groups' abundance. The dominance of Firmicutes suggests that the sclerophyllous matorral could be a different ecosystem to other arid and semi-arid soils with respect to the bacterial community structure.     

Photosynthetic responses of Larrea tridentata to a step-increase in atmospheric CO2at the Nevada Desert FACE Facility

Of all terrestrial ecosystems, the productivity of deserts has been suggested to be the most responsive to increasing atmospheric CO₂. The extent to which this prediction holds will depend in part on plant responses to elevated CO₂under the highly variable conditions characteristic of arid regions. The photosynthetic responses ofLarrea tridentata , an evergreen shrub, to a step-increase in atmospheric CO₂(to 550 μmolmol⁻¹) were examined in the field using Free-Air CO₂Enrichment (FACE) under seasonally varying moisture conditions. Elevated CO₂substantially increased net assimilation rate (A_net) in Larrea during both moist and dry periods of the potential growing season, while stomatal conductance (g_s) did not differ between elevated and ambient CO₂treatments. Seasonal and diurnal gas exchange dynamics in elevated CO₂mirrored patterns in ambient CO₂, indicating that elevated CO₂did not extend photosynthetic activity longer into the dry season or during more stressful times of the day. Net assimilation vs. internal CO₂(A/C_i) responses showed no evidence of photosynthetic down-regulation during the dry season. In contrast, after significant autumn rains, A_max(the CO₂saturated rate of photosynthesis) and CE (carboxylation efficiency) were lower in Larrea under elevated CO₂. In situ chlorophyll fluorescence estimation ofLarrea Photosystem II efficiency (F_v/F_m) responded more to water limitation than to elevated CO₂. These findings suggest that predictions regarding desert plant responses to elevated CO₂should account for seasonal patterns of photosynthetic regulatory responses, which may vary across species and plant functional types.     

Compensatory responses of an arid land crucifer, Chorispora tenella (Brassicaceae), to experimental flower removal

We studied experimentally the compensatory ability of an annual arid land crucifer, Chorispora tenella, with respect to density of conspecifics. Chorispora tenella was able to compensate for flower losses by modulating the rates of abortion of flowers, fruits and ovules. In addition, plants growing at lower density compensated more intensely against losses than plants growing at high density. Nevertheless, as a consequence of compensation, plants in all treatments produced the same quantity of seeds (equal compensation). These results suggest that, from flower production to fruit and ovule maturation, C. tenella maintained a number of ovules that exceeded those that it could eventually mature. This could help to colonize arid environments successfully, in which resources are typically scarce and spatio-temporally unpredictable.     

Comparison of the growth and biomass production of six acacia species in Riyadh, Saudi Arabia after 4 years of irrigated cultivation

The growth and biomass production of six acacia species were studied in the field for 4 years. The species used were Acacia asak, A. negrii, A. seyal, A. karroo, A. ampliceps, and A. stenophylla. The first three species are indigenous while the others are exotic. The results showed both A. ampliceps and A. asak with 100% survival while all A. negrii died. Acacia ampliceps attained the greatest height, diameter, relative growth rate and above-ground biomass while A. asak had the least. Height and diameter growth of acacia species decreased between warm and cold periods of the year.