Differential seedling performance and environmental correlates in shrub canopy vs. interspace microsites

Shrubs in semi-arid ecosystems promote micro-environmental variation in a variety of soil properties and site characteristics. However, little is known regarding post-fire seedling performance and its association with environmental variation in former shrub canopy and interspace microsites. We compared post-fire seeding success and various soil properties important for seedling establishment between shrub canopy and interspace microsites in Wyoming big sagebrush (Artemisia tridentata) plant communities in southeast Oregon, U.S.A. We burned 5, 20 × 20 m sites and established paired canopy and interspace micro-transects seeded with bluebunch wheatgrass (Pseudoroegneria spicata, 193 seeds/m) or crested wheatgrass (Agropyron cristatum, 177 seeds/m). At one year post-fire, seedling density was 69% higher (p = 0.012) for crested wheatgrass (compared to bluebunch wheatgrass) and 75% higher (p = 0.019) for interspace microsites (compared to canopy). However, tiller and leaf area production were over twice as high (p < 0.05) in canopy microsites. Soil color and soil temperature, explained 19–32% of variation in seedling performance metrics. Shrub effects on seeding success are complex and interact with abiotic disturbances, but patterns of increased seedling performance in canopy microsites and their relationships to soil variables may suggest tactics for increasing success of restoration practices.     

Light responses of Carex planostachys from various microsites in a Juniperus community

Juniperus communities are found on over 50 × 10⁶ ha in arid and semiarid habitats in southwestern North America. The drought tolerant sedge Carex planostachys occurs below the canopy in some of these communities. Cover and biomass of C. planostachys are high below the canopy and low in associated gaps. The purposes of this study were to investigate the temporal and spatial physiologic response of C. planostachys to abiotic changes, and determine it's light response characteristics from four contiguous microsites. Net photosynthesis was highest in spring when temperature was cooler and soil water higher, but low carbon uptake continued during summer drought. In addition, C. planostachys demonstrates a capacity to recover from extreme drought, despite water potential measured below ?9.0 MPa. Based on physiological light response curves and gas-exchange measurements, C. planostachys appears tolerant of shaded and full sun habitats. Light levels below the canopy were reduced compared to the gaps, but light saturation of C. planostachys did not change and net CO₂ uptake was only reduced slightly. Carbon uptake was coupled to light levels and not soil moisture. Observed differences in physiological attributes and variation in C. planostachys cover and biomass correspond to the presence or absence of the canopy. Low light compensation points, coupled with reduced respiratory demand, maximize photosynthetic gain in low light microsites. C. planostachys appears to acclimate across a range of light regimes, suggesting photosynthetic plasticity, allowing growth and survival in diverse light microhabitats. C. planostachys, tolerant of drought, appears anisohydric and demonstrates a capacity to acclimate to sun and shaded habitats, which could allow it to occur across a wider range of arid areas.     

Winter establishment of the alien annual Schismus barbatus is not affected by insect herbivory in Northern-Central Monte Desert

We studied the effect of ant herbivory on the establishment and survival of the annual plant Schismus barbatus. We hypothesized that ants may control this species when biomass of native plants is generally low during their vegetative period. We predicted that ant herbivory will decrease plant survival and reproduction. We tested our prediction with an insect exclosure experiment in a sandy desert of Northern-Central Monte.We found more than 12 000 established seedlings per square meter on early May, after two consecutive rain pulses of ca. 20 mm each. Overall, we found that almost 75% of recruited plants survived by the end of the cool season (September), and that 22–24% of the initially established plants survived as mature reproductive plants by the end of the growing season (December). Contrary to our expectation, insect herbivory did not affect plant establishment, plant survival or the proportion of flowering and fruiting individuals of S. barbatus.The large number of seedlings reported, the ability to exploit a temporal window free of plant competitors and enemies, and the availability of microsites where this species can succeed, all suggest that S. barbatus may have the potential to become an important plant invader in the Monte Desert.     

Clubmoss, precipitation, and microsite effects on emergence of graminoid and forb seedlings in the semiarid Northern Mixed Prairie of North America

Clubmoss (Selaginella densa Rydb.) provides extensive ground cover throughout the Northern Mixed Prairie of North America, but little is known about how this cryptogam affects seedling emergence relative to precipitation. Seedbed treatments including a control, Glyphosate applied to kill clubmoss, and clubmoss removal were factorially combined with ambient precipitation, precipitation reduced, and irrigation. Graminoid and forb emergence in the seedbeds and the microsites of live clubmoss, dead clubmoss, bare soil, litter, and caespitose grasses was determined. Forb emergence was greatest with clubmoss removal and irrigation. About 2.6-3.6-fold more forbs emerged from bare soil microsites, but clubmoss reduced forb emergence by 32-76%. Litter reduced forb emergence in 2 of 3 years. Graminoid emergence did not vary among seedbeds (P ≥ 0.780), but irrigating increased seedling emergence. Graminoid emergence was unaffected or improved in live clubmoss, dead clubmoss, bare soil, and litter microsites. In the semiarid Northern Mixed Prairie, above-average precipitation combined with bare soil will favor emergence of forbs. By contrast, clubmoss does not limit graminoids because seedlings emerge from a range of seedbeds and microsites provided diaspores are present and precipitation is above average. Emergence of graminoids and forbs is constrained primarily by precipitation in the semiarid Northern Mixed Prairie.     

Recruiting mechanisms of Cylindropuntia leptocaulis (Cactaceae) in the Southern Chihuahuan Desert

Sexual reproduction and vegetative propagation require the establishment of morphologically diverse reproductive structures (seedlings and stem segments). This study assesses both types of Cylindropuntia leptocaulis recruitment in two microenvironments protected from herbivory (in open space and under the canopy of Larrea tridentata) and with two soil types (collected below L. tridentata canopy and from open space).An observational study revealed that sexual establishment was very limited (only 8 juveniles/ha were found), while clonal establishment was more frequent (59 juveniles/ha); both types of propagule were found only under shrub canopy.Field experiments showed that, as in other Cylindropuntia species but contrasting with findings for Opuntia species, seeds germinated and seedlings survived under nurse plants as well as in open space, but they grew faster in the former environment. Faster growth under canopy shade provides better chances of successful recruitment to later life-cycle stages. Stem segments never established in open space, requiring the shade of nurse plants. This may be due to physiological constraints imposed by their small volume. In synthesis, the results indicate that clonal propagation is the most important recruitment mechanism in C. leptocaulis populations, and that it takes places under shrub canopy.     

Optimising seed broadcasting and greenstock planting for restoration in the Australian arid zone

Vegetation within some parts of Western Australia has been degraded by resource extraction, and ecological restoration is necessary to prevent erosion and reinstate plant biodiversity. Two restoration approaches, seed broadcasting and planting of seedlings, were tested with plant species (Acacia tetragonophylla F. Muell., Atriplex bunburyana F. Muell. and Solanum orbiculatum Poir.) known to have been dominant prior to mining activities in the World Heritage Area at Shark Bay. For broadcast seeding, soil raking and/or ripping increased seedling emergence, but only after sufficient rainfall. Survival of A. bunburyana seedlings (≤92%) was higher than A. tetragonophylla (≤13%) almost two years after planting and soil ripping partly alleviated soil impedance and resulted in increased seedling survival. Shoot pruning, fertiliser and moisture retaining gel had a reduced or detrimental effect on survival. Seedling survival differed between the three experimental sites, with electrical conductivity being the most noted soil difference between the sites. Restoration in the arid environs of the World Heritage Area at Shark Bay in Western Australia is challenging, but this study shows that seedling establishment is technically feasible and provides methodology useful to other arid restoration projects.     

Arbuscular mycorrhizal fungal abundance in the Mojave Desert: Seasonal dynamics and impacts of elevated CO2

Arbuscular mycorrhizal fungi (AMF) play important roles in ecosystem processes. However, little is known about AMF abundance in arid, nutrient-poor environments like the Mojave Desert of North America. We conducted two AMF studies: one examined AMF responses to elevated atmospheric CO₂ at the Nevada Desert FACE (Free-Air-Carbon dioxide-Enrichment) Facility (NDFF), and the second examined seasonal dynamics at nearby sites. In both studies, AMF measurements (root colonization, extra-radical hyphal [ERH] length, and two measures of glomalin-related soil protein [GRSP]) and environmental factors (root length, soil water content, and precipitation) were measured across microsites (beneath shrubs, shrub interspaces) for two species (Larrea tridentata, Ambrosia dumosa). Elevated CO₂ did not significantly affect AMF measurements. Other NDFF studies show no change in fine root production under elevated CO₂ but show increased available nitrogen. We infer that additional fixed carbon under elevated CO₂ is not allocated to soil resource foraging. However, AMF varied seasonally. ERH seasonally declined across species and microsites, but GRSP declined only beneath L. tridentata. Our results combined with previous results indicate that drought negatively affects AMF root colonization. Robust inter-relationship among AMF measurements only occurred between the two measures of GRSP, indicating that resources are independently allocated to different AMF structures.     

Growth and water relation parameters in drought-stressed Coriaria nepalensis seedlings

In the present study, growth and water relation parameters were analysed in drought-stressed Coriaria nepalensis Wall. seedlings. C. nepalensis seedlings were subjected to four drought cycles of 7, 14, 21, and 28-days, and to one control level (watered on alternate days) in a glasshouse. The seedlings failed to survive a 28-days drought during summer. Osmotic adjustment (defined as the decrease in osmotic potential at zero or full turgor in response to water deficit) was measured as the difference between the osmotic potential of seedlings watered on alternate days (control) and those subjected to 21-days drought cycle. Seedlings subjected to 21-days drought had a predawn water potential of −2.60 MPa, and showed an osmotic adjustment of −1.95 MPa at full turgor and −2.17 MPa at zero turgor. The growth of seedlings was positively related to moisture and with water potential. With decline in soil moisture the root:shoot ratio increased while leaf weight ratio decreased. Leaf characteristics, such as leaf number, leaf area, leaf area ratio, specific leaf area and leaf drop, were also affected by moisture stress. This study has indicated that osmotic adjustment is a major adaptive mechanism of C. nepalensis that aids successful regeneration of seedlings in degraded sites with inhospitable soil conditions.     

Growth response of three globose cacti to radiation and soil moisture: An experimental test of the mechanism behind the nurse effect

Cactus seedlings often establish under nurse plants which modify environmental conditions by increasing moisture and decreasing solar radiation, which may cause beneficial and detrimental effects, respectively, on seedling growth. Three soil moisture treatments (5%, 25% and 60%) and two solar radiation levels (100% exposure=243 μmol m⁻² s⁻¹, and 40%=102 μmol m⁻² s⁻¹) were used in a factorial design to analyze seedling growth response of three rare cactus species (Mammillaria pectinifera, Obregonia denegrii and Coryphantha werdermannii). The variables evaluated were relative growth rate (RGR), root/shoot ratio (R/S), and K (RGR_roots/RGR_shoot), obtained from an initial seedling harvest (6-month-old seedlings) and a final harvest 6 months after treatment application. All three species had slow RGRs (0.002–0.012 g g⁻¹ day⁻¹). O. denegrii had the lowest RGR values, but was the only species for which R/S and K varied with soil moisture. While all seedlings responded markedly to soil moisture, no response was observed to radiation treatments. The latter might have been related to the relatively low solar radiation levels present in the greenhouse. Yet, our results suggest that the main benefit nurse plants offer to seedlings is the increase in soil moisture.     

Role of nurse shrubs in restoration of an arid rangeland: Effects of microclimate on grass establishment

A perennial forage grass, Agropyron desertorum was sown under the canopies of four shrub species or in open areas to test for facilitation during seedling establishment in an arid rangeland in Karnakh, Northeast Iran. Height, area and volume of shrubs were measured. Microclimate conditions and seedling establishment were assessed three times within two consecutive growing seasons. Near surface light intensity and air temperature were lower under shrubs, which led to initially higher soil moisture and grass establishment under the canopy of some shrub species. The leguminous shrub (Astragalus gossypinus) showed facilitation during moderate stress (summer 2009), but shifted to a negative effect during the severe drought (summer 2010). Competition, possibly for light, reduced the establishment of Agropyron seedlings under the cushion-like shrub (Acantholimon prostegium). Salsola arbusculiformis and Artemisia kopetdaghensis respectively showed a neutral and a facilitation effect in the first season, but a combined effect of allelopathy and drought led to the high mortality of Agropyron seedlings under their canopy. In conclusion in this arid rangeland, shrubs may facilitate establishment of understory plants under moderate drought stress, and for non-resource factors (light and temperature), but these positive effects are suppressed due to competition under severe drought conditions.     

The association of native and non-native annual plants with Larrea tridentata (creosote bush) in the Mojave and Sonoran Deserts

Creosote bushes (Larrea tridentata) form islands of fertility that influence annual plant abundances in desert ecosystems, and the distribution of native and non-native annuals with respect to creosote may differ. We established plots on the north and south facing sides of L. tridentata in the Mojave and Sonoran Deserts within four different microhabitats spanning from under the shrub canopy into the open. We counted native and non-native annuals twice during the winter growing season to determine the effects of L. tridentata on the spatial distribution of native and non-native annuals. In both deserts, annual plant abundances were higher on the north side of L. tridentata and in open areas. Native annuals in the Mojave were most abundant near the edge of the shrub canopy, whereas native annuals in the Sonoran were most abundant in open areas. The effects of L. tridentata fertility islands on annual abundances were not consistent between the two deserts we studied. Our study emphasizes the importance of research on native and non-native annuals in multiple regions before generalizations can be made about the effects of L. tridentata on annual plant abundances in desert scrub ecosystems.     

Microclimate, freezing tolerance, and cold acclimation along an elevation gradient for seedlings of the Great Basin Desert shrub, Artemisia tridentata

Vegetation, microclimate, seedling frequency, freezing tolerance, and cold acclimation were compared for seedlings of Artemisia tridentata collected from 1775, 2175, and 2575 m elevation in the eastern Sierra Nevada, California. Data were used to test the hypothesis that ecotypic differences in stress physiology are important for seedling survival along gradients from desert to montane ecosystems. The vegetation canopy cover and A. tridentata seedling frequency were greatest at 2575 m, compared to 1775 and 2175 m. Snow cover ameliorated temperatures near the soil surface for part of the winter and depth varied across elevations. Freezing tolerance was compared for seedlings maintained in growth chambers at day/night air temperatures of 25°C/15°C. The temperature at which electrolyte leakage and Photosystem II function (F_V/F_M) from leaves were half-maximum was approximately −13·5°C for leaves of seedlings from all three elevations. Shifting day/night air temperatures from 25°C/15°C to 15°C/5°C initiated about 1·5° of acclimation by plants from all three altitudes, with seedlings from the highest elevation exhibiting the greatest acclimation change. Measurements of ambient air and canopy temperatures at the three elevations indicated that wintertime average low temperatures were consistent with the measured degree of freezing tolerance. At small spatial scales used in this study, pollen and seed dispersal between study sites may have precluded resolution of ecotypic differences. Patterns of freezing tolerance and cold acclimation may depend on a combination of mesoclimate and microclimate temperatures, canopy cover, snow depth, and snow melt patterns.     

Pine plantation bands limit seedling recruitment of a perennial grass under semiarid conditions

Pine plantations coexist with Stipa tenacissima grasslands in many semiarid western Mediterranean areas. We compared three microsites created by a 30-year-old Pinus halepensis plantation: below pine plantation line (BP), below canopy of pines (BC) and interline bare band (BA). They were evaluated in terms of soil properties, pine litter and suitability as recruitment niches for S. tenacissima. Next, in a manipulative experiment in growth chambers we tested the hypothesis that pine litter interferes with the seedling emergence of S. tenacissima. Three treatments in pots were compared: (a) soil from BA; (b) intact soil + litter from BP; and (c) soil + litter from BP, which was mixed in the laboratory (BPMX). In the field the main microsite differences were pine cover and litter cover and thickness. Seedling emergence was significantly greater in BA than in BP. Emergence and litter depth fits a linear regression model. In the growth chamber litter did not interfere with the emergence of S. tenacissima. However, seedlings grown without litter were 28% longer and their mass was 27% greater than in the litter treatments. The detected pine litter interference may be relevant for plant dynamics and might be considered in forestry management programs.     

Factors affecting the distribution, abundance and seedling survival of Mammillaria gaumeri , an endemic cactus of coastal Yucatán, México

The population status of an endemic cactus (Mammillaria gaumeri) was assessed in coastal Yucatán, México. Similar numbers of mature plants were found in open and full sun microhabitats. Plants in full sun had more ramets and were of larger diameter. There was a correlation between M. gaumeri abundance and the biomass of a disturbance indicator, Myrmecophila tibicinis (Orchidacea). Higher seedling survival in shade microhabitats and aggregated distribution of adults suggests that shrubs serve as nurse plants. Initial mortality rates of seedlings were many times higher in full sun microhabitats than in shade conditions. Herbivory had no effect on seedling survival.     

The influence of elevation, shrub species, and biological soil crust on fertile islands in the Mojave Desert, USA

We quantified soil nutrients and biological crust cover (bryophytes and lichens) under the canopies of three species of Mojave Desert shrubs and in interspaces between shrubs at three elevations to determine the effects of shrub species, soil crust, and elevation on islands of soil fertility. Means of pH, organic matter, total Kjeldahl nitrogen, nitrogen mineralization, and gravimetric soil moisture are significantly greater in soils under Ambrosia dumosa (Gray) Payne, Larrea tridentata Cov., and Coleogyne ramosissima Torr. than soils from adjacent interspace microhabitats. Although soil moisture and soil organic matter increase by a factor of 1.5 from the low elevation to the high elevation site, the ratio of shrub to interspace concentrations, or the difference in mean soil variables between shrubs and interspaces, is effectively constant and independent of elevation. Total bryophyte and lichen cover is relatively low (24.5%), however, there are 11 species of bryophytes and two species of lichens distributed across three elevations with the highest species richness and cover at the low-elevation site. Bryophyte and lichen cover is correlated with silt but is not related, consistently, to soil nutrients. Overall, the balance of processes controlling spatial aggregation of soil nutrients under shrubs is remarkably insensitive to potential differences in organic inputs among elevations, shrub species, and soil crust surfaces.     

Plant–water relationships and growth strategies of Jatropha curcas L. seedlings under different levels of drought stress

Although Jatropha curcas, an important tropical biofuel crop, is reputed for its drought resistance, its ability to perform under dry conditions has hardly been investigated. In a greenhouse experiment we investigated the plant–water relationships of Jatropha seedlings from different accessions under different levels of drought stress. There was little difference in plant–water relations between accessions. Drought significantly reduced leaf area, biomass and relative growth rate, but had no effect on specific leaf area, daily range in leaf water potential, leaf water content, transpiration efficiency or aboveground biomass water productivity, corrected for atmospheric conditions. Stem wood density was equally low (0.26 g cm^?3) for all treatments. Stem water content was lowest for dry treatment seedlings. Based on these results, Jatropha could be characterized as a stem-succulent tree. In contrast to other stem-succulent deciduous trees, leaves were not shed immediately after the seedlings were confronted with drought. Instead, at the onset of drought, leaves with a higher adaxial stomatal density were formed, after which leaves were only gradually shed. The role of the succulent stem in the water economy of Jatropha was confined to balancing the small water losses of the leaves during drought.     

Moderate water supply and partial sand burial increase relative growth rate of two Artemisia species in an inland sandy land

Artemisia ordosica and Artemisia sphaerocephala seedlings were buried in sand to depths of 0, 0.25, 0.5, 0.75, 1 and 1.25 times the seedling height (H) and supplied with 2.5, 5.0, 7.5 and 10.0 mm of water every three days. For A. ordosica, the highest relative growth rate (RGR) and seedling height were achieved with 75 mm/month water supply and 0.5 H burial at 50 days, and the RGR decreased with 100 mm/month water supply. For A. sphaerocephala, the highest RGR was reached with 50 mm/month water supply and 0.5 H burial, but plants from this species had the greatest seedling height with 25 mm/month water supply and 0.75 H burial. A. ordosica and A. sphaerocephala have different proportions of their biomass in roots and shoots and thus are adapted to sandy environments in different ways. Partial sand burial significantly increased the RGR and growth height of the two Artemisia species, but complete burial lead to seedling death. Differences in the growth response to burial and moisture levels help explain why A. sphaerocephala is primarily found on drifting dunes and A. ordosica on semi-fixed and fixed dunes.     

An assessment of revegetation treatments following removal of invasive Pennisetum ciliare (buffelgrass)

In semi-arid regions of North America and Australia, Pennisetum ciliare (L.) Link (syn. Cenchrus ciliaris; buffelgrass) is highly invasive and has the potential to introduce fire to fire-intolerant ecosystems. Major efforts to remove P. ciliare continue and it is essential that P. ciliare be prevented from recolonizing. This study investigated potential methods to revegetate with native herbaceous plants: sowing seeds; sowing seeds and mulching; sowing seeds and transplanting seedlings; and relying on natural revegetation from the seedbank. The treatments were applied in 2009 and 2010 at sites in the Sonoran Desert which had undergone P. ciliare removal. Monsoon precipitation was below average each year and seedling emergence and establishment rates were low. There are indications that soil disturbance associated with planting seedlings promoted P. ciliare emergence and increased mortality of brittlebush (Encelia farinosa A. Gray ex Torr.), a common native perennial forb. Addition of mulch may have promoted P. ciliare over native grasses, and seeding had no effect. We did not find competition between herbaceous seedlings. Rather, native and exotic grass seedling densities were positively correlated across sites. Under prevailing conditions, low precipitation appeared to limit herbaceous plant establishment and none of the treatments reduced P. ciliare abundance.     

Nest site characteristics and nesting success of the Western Burrowing Owl in the eastern Mojave Desert

We evaluated nest site selection at two spatial scales (microsite, territory) and reproductive success of Western Burrowing Owls (Athene cunicularia hypugaea) at three spatial scales (microsite, territory, landscape) in the eastern Mojave Desert. We used binary logistic regression within an information-theoretic approach to assess factors influencing nest site choice and nesting success. Microsite-scale variables favored by owls included burrows excavated by desert tortoise (Gopherus agassizii), burrows with a large mound of excavated soil at the entrance, and a greater number of satellite burrows within 5 m of the nest burrow. At the territory scale, owls preferred patches with greater cover of creosote bush (Larrea tridentata) within 50 m of the nest burrow. An interaction between the presence or absence of a calcic soil horizon layer over the top of the burrow (microsite) and the number of burrows within 50 m (territory) influenced nest site choice. Nesting success was influenced by a greater number of burrows within 5 m of the nest burrow. Total cool season precipitation was a predictor of nesting success at the landscape scale. Conservation strategies can rely on management of habitat for favored and productive nesting sites for this declining species.     

Response of newly established Juniperus ashei and Carex planostachys plants to barrier-induced water restriction in surface soil

Plant species in the Edwards Plateau region of central Texas often face drought conditions. Some of these species reduce water stress by rooting into moist bedrock crevices. A dominant woody plant (Juniperus ashei) and co-occurring xeric sedge (Carex planostachys) were greenhouse-grown in growth tubes with or without water-blocking partial barriers in tubes with water added below the barrier (～70 cm). Soil moisture at various depths was related to growth response (above- and below-ground biomass, root length, and number of root tips) per species. Barrier treatment yielded significantly drier soil than non-barrier. Juniperus plants were significantly smaller with barrier treatment, while Carex showed no significant differences between treatments. The results suggest the sedge is more able than the shrub to increase root production in the upper layers of the soil when faced with access restriction to lower layers. The sedge may resist drought by exploring sub-superficial soil layers. When access to sub-superficial water sources is restricted, the shrub is more affected than the sedge. The plants’ growth in water-restricted surface soil for several months suggests these species may not need deep water to survive. In fact seedling growth may differ in mature plants.