Seasonal variation of the soil seed bank of grasses in central Argentina as related to grazing and shrub cover

The objectives of this research were to (a) study seasonal variation of the seed bank of grasses in two areas with different grazing history, (b) analyse the effect of the presence of shrubs on the seed bank and (c) study seed viability and dormancy in buried seeds of Piptochaetium napostaense and Stipa tenuis. The seed density of grasses was in general low and showed seasonal variation. Most of the grasses showed maximum seed density in December when seed dispersal occurs. Shrub cover did not show a marked influence in seed accumulation. The relatively high number of damaged seeds suggests that predation may be an important factor in seed bank dynamics in the Caldenal. Seeds of dominant perennial forage grasses showed dormancy that could explain, at least in part, persistence of these species in spite of periodical disturbance.     

Distribution of cacti in Larrea spp. shrublands in Mendoza, Argentina

In three Larrea shrublands, cacti grow under Larrea canopy at northern and southern exposures and in open spaces were counted. Cacti distribution was related to environmental variables. Seven out of the 15 cacti present were found to be associated with shrubs, whereas the other species were found in the open spaces. The lower temperature and radiation occurring under shrubs at south exposures would explain the absence of cacti in these places. The higher organic matter, nutrients, and moisture in the soils under shrubs appeared to be correlated with higher cacti density. The presence of cacti seedlings only under shrubs could be attributed to the favourable environmental conditions for cacti seed germination and seedling survival that occurred in these places.     

The role of facilitation in seedling recruitment and survival patterns, in the Strandveld Succulent Karoo, South Africa

Several studies in arid ecosystems have demonstrated that seedling establishment and survival were greater underneath the canopies of shrubs than in the open spaces between shrubs. These results led to the formulation of the hypothesis that seedling recruitment and survival were dependent on the shelter provided by shrub species (nurse-plant hypothesis). Seedling emergence and survival beneath and between the canopies of selected perennial shrub species were investigated at five localities in the Strandveld Succulent Karoo, each dominated by a different shrub species. Seedling emergence and survival were also examined at three localities dominated by annual species. In general, species richness and seedling densities were significantly higher in open areas than underneath shrubs, while seedling survival percentages did not differ significantly between microsites. Therefore, no evidence could be found to support the hypothesis that seedling recruitment and survival were facilitated by the presence of shrub species. Although most species will be able to establish in the absence of shrubs, the presence of woody species may have other advantages when restoring vegetation in mined areas.     

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.     

Seed banks in a degraded desert shrubland: Influence of soil surface condition and harvester ant activity on seed abundance

We compared seed banks between two contrasting anthropogenic surface disturbances (compacted, trenched) and adjacent undisturbed controls to determine whether site condition influences viable seed densities of perennial and annual Mojave Desert species. Viable seeds of perennials were rare in undisturbed areas (3–4 seeds/m²) and declined to <1 seed/m² within disturbed sites. Annual seed densities were an order of magnitude greater than those of perennials, were one-third the undisturbed seed densities on compacted sites, but doubled on trenched sites relative to controls. On trenched sites, greater litter cover comprising the infructescences of the dominant spring annuals, and low gravel content, enhanced seed densities of both annuals and perennials. Litter cover and surface ruggedness were the best explanations for viable perennial seed densities on compacted sites, but litter cover and the presence of a common harvester ant explained annual seed densities better than any other surface characteristics that were examined. Surface disturbances can have a varied impact on the condition of the soil surface in arid lands. Nevertheless, the consistently positive relationship between ground cover of litter and viable seed density emphasizes the importance of litter as an indicator of site degradation and recovery potential in arid lands.     

Annual growth zones in stems of Hypericum irazuense (Guttiferae) in the Costa Rican páramos

The high peaks of the Cordillera de Talamanca in Costa Rica support shrub- and grass-dominated páramo ecosystems that experience stand-replacing wildfires. The dry season that facilitates these fires results in dormancy in plant growth and provides an opportunity to use dendrochronological analyses to determine ages of plants in burn sites to support studies of fire history and postfire vegetation recovery. This study investigates the formation of annual growth zones in stems of the common shrub Hypericum irazuense. Unlike other páramo shrubs, H. irazuense rarely resprouts following fire, instead recovering through seedling recruitment following seed dispersal from the unburned periphery. Laboratory analysis of 19 prepared cross sections from 15 stems shows that samples of H. irazuense from burned areas can provide an estimate of the minimum number of years since the previous fire, supporting earlier work based on field examination of stems. Including a time lag for seedling recruitment or resprouting refines that estimate. Counts of growth zones in most sections coincided with dates of the last known fires. The presence of annual growth zones in H. irazuense places the species within a relatively small group of woody Neotropical species for which annual rings or growth zones have been demonstrated.     

Long-term vegetation response to mesquite removal in Desert Grassland

Forty-six years of vegetation response to mesquite removal at dry, low elevation sites on the Santa Rita Experimental Range in southern Arizona was only slightly different than the vegetation dynamics where mesquite trees were left intact. Only the density of threeawn grass species (Aristida spp.) was greater in the mesquite removal areas: and that difference persisted even after the cover of mesquite was no longer different between treatment and control areas. Cover of shrubs and perennial grasses, and density of all other perennial grasses did not differ between mesquite treatments throughout the study period. Mesquite cover on treated areas was not different than untreated areas 40 years after tree removal. The long-term results support the interpretation that vegetation dynamics at these dry locations, are not limited by the abundance of neighbouring mesquite. Alternatively, mesquite abundance is self-limiting at levels less than would influence grass abundance and precipitation anomalies may override any effects of neighbouring mesquite. Practically, these results suggest that areas with 350 mm year⁻¹ of annual precipitation and <20% mesquite cover may have very little potential for increasing grass abundance through the removal of mesquite trees.     

Abiotic conditions, neighbour interactions, and the distribution of Stipa tenacissima in a semiarid mountain range

Abiotic conditions and interactions among plants are important factors determining plant geographical distributions. With the aim of analyzing the distribution of a perennial grass species in a semiarid mountain range, we experimentally tested above- and belowground effects of shrubs on the grass along a gradient of aridity. We measured growth, physiology and reproductive output of Stipa tenacissima tussocks surrounded by intact or manipulated neighbour shrubs. Performance of Stipa improved under mesic conditions, but the net effect of shrubs on Stipa did not change along the gradient. Belowground, we did not find significant effects, suggesting belowground resource partitioning between tussocks and shrubs. Aboveground net effects of shrubs on Stipa were generally positive and independent from abiotic conditions. The improved performance of Stipa at the mesic end of gradient suggests that its absolute physiological optimum lies out of its realized range. Overall, the arid end of the geographical distribution of Stipa in this mountain range may be determined by environmental severity. The coincidence of the upper limit of Stipa with the lower tree limit suggests that the more mesic end of Stipa distribution may depend on other factors; most likely competition with trees, which, unlike shrubs, could be excluding Stipa tenacissima from forests.     

Reproductive and recruitment traits as indicators of the invasive potential of Kalanchoe daigremontiana (Crassulaceae) and Stapelia gigantea (Apocynaceae) in a Neotropical arid zone

Populations of two ornamental exotic species, Kalanchoe daigremontiana and Stapelia gigantea, have been recently detected inside a protected area containing arid ecosystems in Venezuela. We indirectly tested their invasive potential by characterizing their reproductive biology and recruitment patterns and comparing our estimates against Baker's Law and reproductive profiles reported for invasive plants. K. daigremontiana is autogamous, produces >16,000 seeds per plant and also reproduces clonally. Despite low seed viability (17.9%) and germination rates (11.9%), seeds were present in the seed bank. Plantlets of asexual origin showed high survival (75–100%) compared to seedlings of sexual origin (10%). S. gigantea is self-incompatible, xenogamous and produces close to 1500 seeds per m² of plant tissue. Seed viability (77%) and germination rates (62%) were high, but this species is not represented in the seed bank. It has a vegetative growth. A combination of reproductive and recruitment attributes, which match those considered in Baker's Law and others reported for invasive plants, confer K. daigremontiana and S. gigantea the potential to invade Neotropical arid zones; the former mainly through selfing and production of numerous asexual plantlets, and the latter through an association with a locally abundant pollinator and production of wind dispersed seeds.     

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.     

Fire in Chihuahuan Desert grassland: Short-term effects on vegetation, small mammal populations, and faunal pedoturbation

A prescribed burn resulted in significant decreases in canopy cover of the grasses: Bouteloua eriopoda, Sporobolus flexuosus, and Aristida purpurea. One year post-burn, basal cover of B. eriopoda remained significantly lower in burned patches than in unburned areas but there were no differences in basal cover of the other perennial grasses. Only one species of the 14 summer annual species occurred in both burned and unburned plots. There were six species of spring annuals in burned patches but no spring annuals in the unburned grassland ten months post-burn. Fire killed 100% of the snakeweed shrubs (Gutierrezia sarothrae), 77% of the Ephedra torreyana shrubs, and 36% of the Yucca elata. All mesquite shrubs that were top-killed by fire, resprouted one month post-burn. Fire had no effect on abundance and species richness of rodents. There were fewer wolf spider, Geolycosa spp. burrows in burned areas than in unburned grassland. The area and volume of soil in termite galleries and sheeting were significantly larger in the unburned grassland than in the burned areas.     

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.     

Seed reserves in the central Monte Desert, Argentina: implications for granivory

It has been suggested that lower rates of granivory in South American warm deserts compared to their North American counterparts arose from a reduction of seed reserves in the former owing to the extinction of argyrolagid marsupials. We measured seed reserves in two habitats of the central Monte Desert in an attempt to detect such seed decline, but to no avail. After moderate rainfall, maximum seed standing crops reached 16,000 and 23,000 seeds m⁻²in shrublands and open forests, respectively. Under the canopy of trees and shrubs there were 19,000 and 37,000 seeds m⁻²; whereas in exposed areas there werec. 10,000 seeds m⁻²in both habitats. Seed banks in other South American semi-arid areas showed similar values. Total grass seeds as well as those presumably preferred by ants also seem to be similar in both continents. Hence, granivory in South America, as has been already reported for Australia, is lower than in North America in spite of the great similarity of seed bank sizes. Moreover, argyrolagids were unlikely seed-hoarding granivores, therefore some other reason than argyrolagid extinction should be sought to explain the lack of specialized seed-eating mammals, and the smaller overall seed consumption rates in South American deserts.     

Gas exchange responses of a desert herbaceous perennial to variable sunlight in contrasting microhabitats

Over the course of a day, aridland plants experience a range of incident photosynthetic photon flux (PPF) spanning three orders of magnitude. Rapid photosynthetic responses to changes in PPF have large effects on individual plant carbon gain and water use patterns, hence are important to plant distribution and success. We investigated the response over time of photosynthesis (A), stomatal conductance (g), and inter-cellular CO₂ concentration (C_i) to step changes in PPF in a long-lived aridland perennial that typically occurs in two contrasting microhabitats, shade under shrubs of other species and unshaded inter-shrub spaces. An initial rapid response in A and C_i for plants in both microhabitats occurred after abrupt changes in PPF. This was followed by slower changes in these parameters during the rest of the light or dark period. Stomatal conductance responded more gradually to step changes in PPF. The initial increase in A after a dark cycle was significantly greater for plants under shrubs than for plants in inter-shrub spaces, but other changes in A, g and C_i did not differ. We attribute the similar responses in plants from different microhabitats to natural variations in solar radiation and limited selection for differentiation due to population gene pools dominated by plants in the open. Our results support the hypothesis that variable light regimes select for photosynthetic gas exchange processes that closely track changes in incident PPF. Our data also support the hypothesis that gas exchange responses to variable light regimes in aridland plants minimize trade-offs between carbon gain and water loss.     

Is the analysis of plant community structure better than common species-diversity indices for assessing the effects of livestock grazing on a Mediterranean arid ecosystem?

Grazing in arid Mediterranean ecosystems brings about changes in species diversity, but the best way to measure such changes is unclear. In this study we compared various methods in order to identify indicators that might be useful for the management and conservation of grazed arid Mediterranean ecosystems.Changes in community structure and composition were compared along a previously studied grazing gradient in Cabo de Gata-Nijar Natural Park. Plant diversity was measured by calculating diversity (Shannon information index), evenness (Pielou index) and richness (species number). Rarefaction curves were used to measure plant species richness and the results were compared with traditional richness values. Community structure was measured as the percentage of bare ground and the coverage of overall perennial grasses, Stipa tenacissima L., perennial shrubs and annual plants. Our results showed that the proportion of bare ground, the cover of perennial shrubs and the relative abundance of S. tenacissima are good indicators of grazing effects on vegetation. Overall plant community structure was more sensitive to the grazing gradient than were the diversity indices. Finally, the adequacy of standardized methods for comparing species richness (i.e. rarefaction curves) along a grazing gradient are discussed.     

The influence of plant removal on succession in Wyoming big sagebrush

Restoration treatments are based on the largely untested notion that desired recovery of plant communities following disturbance wouldn’t occur in the absence of active intervention. We identified rate of short-term (10 year) floristic changes following removal of plant functional groups in Wyoming big sagebrush plant communities in 1999-2005 and 2008. Treatments imposed on 6 × 6 m plots were: 1) removal of all plant functional groups, 2) perennial grass removal, 3) shrub removal and 4) control. Our data suggest recovery of the shrub component on shrub removal plots could take decades. Similarly, perennial grass cover and density on perennial grass removal plots was less than half that of unaltered plots 10 years after treatment. When all functional groups were removed, cover of annual forbs, annual grasses, and shrubs returned to unaltered levels within ten years or less. Perennial forbs were unaffected (p > 0.05) by treatment. The fact that natural recovery of some components occurred within a relatively short post-disturbance time interval (i.e. <10 years) suggests that intervention may not be necessary for some functional groups. Restoring shrubs in areas dominated by perennial grasses may require targeted reductions of competing perennial grasses. Conversely, shrub dominance may limit perennial grass re-establishment.     

Disentangling Fabiana imbricata (Solanaceae) regeneration: The importance of disturbance and rainfall

In Mediterranean ecosystems, fire influences plant population dynamics and changes plant community structure by affecting germination and establishment of seeder shrubs. Fabiana imbricata is a long-lived seeder shrub with a broad distribution in South America. In Northwestern Patagonia grasslands there are many records of F. imbricata recruitment after fires. We hypothesized that recruitment is promoted by: 1) soil erosion that exposes deeply buried seeds; 2) direct fire effects such as heat that could break seed dormancy; 3) indirect fire effects (i.e. increase of light availability and elimination of allelopathic substances) and 4) water availability in spring. In field conditions, we tested the effects of shading and soil disturbance on F. imbricata recruitment and seed availability and distribution in the soil profile. Under controlled conditions, we investigated the influence of leaf leachate, light, heat and water on seed germination and seedling emergence. Seed germination was inhibited by leachate, and seedling emergence was inhibited by seed heated to 80 °C and 120° for 5 min. F. imbricata seedlings only emerged in germination treatments that simulated wet and very wet spring conditions. Fire and postfire wind that favor seed exposure, combined with high precipitation early in the growing season, would be necessary for successful shrub recruitment. When F. imbricata shrubland colonizes the grassland, it strongly modifies local vegetation structure, reduces species richness and increases fuel loads. F. imbricata is a key species in the ecosystem specially related to fire regime. Information on the postfire regeneration of this species would provide valuable knowledge of the changes in biomass accumulation in a fire-prone Mediterranean ecosystem.     

Analysis of snow-vegetation interactions in the low Arctic-Subarctic transition zone (northeastern Canada)

Abstract

Recent studies have shown that northern vegetation has been growing in relation to a warming climate over the last four decades, especially across the transition zone between tundra and taiga. Shrub growth affects snow properties and the surface energy budget, which must be better studied to quantify shrub-snow-climate feedbacks. The objective of this research is to improve the characterization of the impact of shrubs on snow evolution, from its accumulation to its melt, using in-situ and satellite measurements. The research is presented for the Umiujaq site, Nunavik, representative of the low Arctic–Subarctic transition zone. Snow depth, measured along numerous transects spanning different land cover types is found to increase by a factor 2.5–3 between tundra and forest, while snow density decreases. This illustrates the trapping effect of vegetation well. Complementary, continuous snow depth measurements using weather stations from two sites (tundra with low shrubs and a small clearing with shrubs within the forest) show different site-dependent behaviors. Spatial analysis from high-resolution Pleiades images combined with Landsat (Normalized Difference Snow Index) and MODIS (Fractional Snow Cover) images suggest a slight delay in melt over open and dense forest areas compared to tundra and dense high shrubs.     

Restoration of palatable grasses: A study case in degraded rangelands of central Argentina

Restoration of palatable grasses on degraded rangelands dominated by unpalatable grasses in central Argentina is limited by low availability of seeds and safe sites for seedling establishment. The objective of our study was to determine how mechanical disturbance of unpalatable grasses (Stipa spp.) in combination with seeding of a palatable grass (Poa ligularis) influenced species composition in a degraded rangeland excluded from livestock grazing. In April 2001 10 blocks were uniformly distributed on a previously burned site dominated by unpalatable grasses, and treatments applied in 8 m×8 m experimental plots. Treatments were ‘disked and seeded’ and control (no disking, no seeding). Perennial plant cover and end-of-season standing crop, at species or species group level, were assessed in December 2002/2003 and in December 2004/2006, respectively. P. ligularis out-competed both tillers of unpalatable grasses that survived mechanical disturbance and seedlings of unpalatable grasses established after mechanical disturbance. The cover and end-of-season standing crop of unpalatable grasses was higher in the control than in the ‘disked and seeded’ treatment, whereas the cover and end-of-season standing crop of P. ligularis was higher in the latter than in the former treatment. Our results suggest that a rapid transition from a state dominated by unpalatable grasses to a state dominated by palatable grasses can be achieved by mechanical disturbance of unpalatable grasses in combination with seeding of palatable grasses in semiarid rangelands of central Argentina.     

Effects of seed age and seeding depth on growth of Stipa lagascae R. & Sch. seedlings

Native grass species are being considered in the restoration of degraded rangelands in Tunisia due to the failure of introduced species to adapt to the ecological constraints of the region. The objective of this study was to determine the effects of seed age and seedling depth on emergence and seedling vigor of Stipa lagascae, a perennial bunchgrass that is native to the region and shows promise as a species for restoration purposes. Total emergence percentage of seedlings and the coefficient of rate of emergence (CRE), the seedling vigour as well as above and below-ground biomass were measured. Younger seeds had higher emergence rates than older seeds. However, seed age did not affect variables associated with seedling vigour. Seeding depth significantly affected all parameters tested. A seeding depth of 4–6 cm was found to be optimal.