Seed removal patterns in burned and unburned desert habitats: Implications for ecological restoration

In desert ecosystems, selective foraging by seed consumers affects distribution and composition of soil seed banks, influencing plant population dynamics. However, the roles of consumers in burned habitats where direct seeding is used during ecological restoration to replenish depleted seed banks have not been well established. We evaluated seed removal for nine seed species over 12 months in burned and unburned shrublands in the Mojave Desert, USA. Percentage of total seed removed was highest during spring (16% of offered seed) and summer (21%). Rodents removed much of the large-seeded Coleogyne ramosissima in both burned and unburned habitats, while seed removal of this species by ants was low in burned and moderate in unburned habitats. Ants removed the greatest amount of small-seeded species (Penstemon bicolor, Encelia farinosa, and Sphaeralcea ambigua) in unburned habitat, indicating that ants can exploit different seed masses. Seed removal imposes limitations on seed availability, particularly for large-seeded species, as both rodents and ants selected seeds of C. ramosissima. Successful restoration seeding projects in arid lands may require protecting seed from granivore pressure, and seed species selection and season of seeding warrant consideration to reduce seed loss.     

Seed dispersal and seed fate in Joshua tree (Yucca brevifolia)

Joshua tree (Yucca brevifolia) is a charismatic symbol of the Mojave Desert. Despite its familiarity, we know little about the reproduction of this species, including mechanisms of seed dispersal. Here we examine mechanisms of seed dispersal and resulting seed fate. We experimentally tracked fruit and seed removal and followed the fates of Joshua tree seeds using radioactive tracers. The majority of Joshua tree fruits monitored were taken directly from the tree canopy by white-tailed antelope squirrels, and seeds and fruits on the soil surface were quickly removed by animals. Rodents given seeds labeled with scandium-46 cached them between 0.1 cm and 4.1 cm deep. Seedling emergence was most common for seeds planted 1 cm deep, whereas seeds placed on the soil surface seldom germinated. Wind dispersal is unlikely because fruits and seeds lack adaptations for wind dispersal; wind speeds required to move Joshua tree seeds and fruits across the soil surface were higher than those typically found in the Mojave Desert. Further, rodents removed most seeds before abiotic burial was possible. We conclude that most Joshua tree seeds are dispersed by scatter hoarding by rodents, and that caches made by rodents are suitable sites for seedling emergence.     

阿拉善高原荒漠植被组成分布特征及其环境解释 Ⅳ.土壤种子库特征研究

阿拉善荒漠土壤种子库种子密度主要受到水、热变化梯度的影响而呈现水平地带性分布规律。土壤种子库的分布主要集中在0～5cm的土层中,深层土壤很少有种子库的存在。阿拉善土壤种子库共统计到26种植物种子,分属9科22属,以禾本科最多;生活型以一年生草本植物种子的比例最高,达到51.36%。极端干旱环境条件下不存在有效的土壤种子库。土壤种子库多样性(Simpson指数)在东阿拉善草原化荒漠区为最高,在0.546～0.917之间;西阿拉善荒漠区次之为0.417～0.915;额济纳绿洲区为0.855,说明阿拉善荒漠区从东到西,土壤种子库物种多样性指数呈现下降的趋势。土壤种子库相似性研究表明,阿拉善荒漠不同区域的土壤种子库相似性差异较大,东阿拉善荒漠化草原植被区,相似性系数范围为0.17～0.67,西阿拉善典型荒漠区为0～0.75,额济纳戈壁荒漠区和东、西阿拉善土壤种子库的相似性系数范围是0～0.4。说明阿拉善荒漠区,由于生境和植被的异质性程度高,导致土壤种子库之间的差异性显著。阿拉善荒漠的土壤种子库和地上植被组成上差异显著,地上植被的优势种以旱生、中生灌木、半灌木为主,土壤种子库中以一年生、多年生草本为主,没有统计到灌木种子的存在,表明土壤种子库对于灌木层片更新和演替的贡献非常小。这些发现对了解干旱荒漠植被自然修复特点具有重要的潜在贡献。     

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.     

Effect of environmental heterogeneity on field germination of Opuntia tomentosa (Cactaceae,Opuntioideae) seeds

We studied in Opuntia tomentosa the effect of plant shade (orientation and distance), and the place where the seeds remained before germination. Seeds were collected in 1998 and a) sown immediately on soil (non-buried seeds), b) stored in the laboratory and sown on soil during the 1999 rainy season (control) or c) buried after collection (under the shade of a tree, a rock and in an open space), exhumed 7 months later and sown in 1999 (exhumed seeds). Seeds were sown under or outside the shade of several trees; or on the northern, eastern and western sides of a 14-m-tall tree. In 1999, germination was high in all samples, but the control seeds had delayed germination. After two rainy seasons, non-buried seeds had the lowest germination rate. Germination was not affected by the distance from the tree. Outside of the plant's shade, germination was low. Germination was affected more by the environmental heterogeneity than by the individual plant shade. Germination probabilities were the highest at 16–19 °C; field germination was zero at 20.6 °C. Seed polymorphism and microenvironmental heterogeneity spread germination across two rainy seasons creating a permanent seed bank.     

Environmental control of seed dormancy and germination in the short-lived Olimarabidopsis pumila (Brassicaceae)

The purpose of our study was to better understand seed germination ecology of the spring annual short-lived Olimarabidopsis pumila, which grows in the Gurbantonggut Desert, China. Seeds underwent after-ripening at 4, 20 ± 2 (room temperature) and 30 °C. After dormancy was broken, germination capacity was a function of temperature and presence of light. For the temperature range studied (4–30 °C), germination capacity was significantly higher between 15 and 25 °C than at temperatures above or below them. Alternating temperatures of 20/10, 25/10 and 25/15 °C were favorable for germination. Although GA₃ did promote dark germination of seeds, GA₃ did not replace light for germination. Seeds germinated to >75% if light exposure time was over 8 h and temperature and moisture conditions were favorable. Seeds were able to germinate at relatively low water potentials (83% at ?0.41 MPa), but decreasing water potentials produced detrimental effects on germination percentage and rate. Thus, dormancy characteristics and germination behavior of O. pumila seeds ensure that germination occurs in the desert only when soil moisture conditions are favorable for seedling establishment and survival.     

Effects of the abandonment of the burrowing mounds of fat sand rat (Psammomys obesus cretzschamar 1828) on vegetation and soil surface attributes along the coastal dunes of North Sinai,Egypt

Fat sand rats (Psammomys obesus) are strictly herbivorous and live in densely complex burrows in the desert of North Africa and the Middle East. Little is known however about the effect of their foraging and burrowing activities on surface morphology and plant community attributes. This study evaluated such effects by comparing burrow and mound surface morphology, canopy of the main host chenopod shrub Anabasis articulata, vegetation cover, and plant abundance and species richness on and off both active and abandoned colonies in the semi-stabilized sand dunes of the Northern Sinai. In general, active burrow systems were characterized by reduced A. articulata canopy area, and more soil disturbance, with higher and larger burrow mounds dominated by bare ground, dung, and dead and fresh litter. The abandonment of mounds for five years has resulted in significant increases of plant cover, canopy height, abundance and species richness. Vegetation structure and plant species assemblages differed between mound and non-mound patches of both active and abandoned sites. The results suggest that fat sand rats can have significant direct and indirect, short-term and long-term effects on vegetation dynamics and structure through their mound building and foraging.     

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.     

Effects of fire and environmental variables on plant structure and composition in grazed salt desert shrublands of the Great Basin (USA)

Fire in shrub-dominated portions of the Great Basin, largely fueled by non-native annuals such as Bromus tectorum, has become an important structuring force altering vegetation composition and soil characteristics. The extent to which fire affects native species in drier portions of the Great Basin, termed salt desert, is poorly documented. We conducted a survey of grazed salt desert habitat in northwestern Nevada 5 years after wildfires burned 650,000 ha, with the goal of investigating community response to fire and factors correlating with post-fire recovery. We found that recruitment of a dominant shrub, Artemisia spinescens, is severely restricted following fire: it occurred in only 2 of the 24 burned sites. The co-dominant shrub, Atriplex confertifolia, occurred in most burned sites although on average its percent cover was one-third lower than adjacent unburned sites. Biotic soil crust cover was four times lower, and non-native species cover 5 times higher, in burned sites compared to unburned. Ordination analyses confirmed differences among plant communities in burned versus unburned sites, with environmental variables soil conductivity, plant litter, soil potassium (K⁺) and pH explaining 38% of the variance in community composition. However, we found no environmental predictors of recovery for native species in burned sites. Future recruitment is likely to be further limited, as fire frequency in the salt desert is expected to increase with invasion by non-native annual grasses and with global climate change.     

Soil seed bank longevity of the exotic annual grass Bromus rubens in the Mojave Desert,USA

Although arid environments are often considered among the least invaded terrestrial biomes, the impacts of exotic plant species can be severe and long lasting. Bromus rubens (red brome) is an exotic annual grass species in the Mojave Desert known to outcompete native plant species, alter habitat, and promote accumulation of fuel that contributes to increasing fire frequency and severity. We assessed longevity of the exotic B. rubens seeds in the soil by burying seeds at four depths (0, 2, 5, and 10 cm) and recovering seeds 6, 12, 18, and 24 months after burial. Seed viability was reduced with greater burial depth and greater time since burial. A relatively small proportion of seeds retained viability for two years, suggesting that while the B. rubens seed bank can be large, it is relatively short-lived. Although B. rubens apparently relies more on the annual production, dispersal, and germination of seeds than on a long-lived seed bank for its annual recruitment, the numerous seeds produced by individual plants indicate that even a small proportion of seeds remaining viable for more than a year can aid recruitment from the seed bank and is an important factor in understanding population dynamics.     

Pre- and post-dispersal seed loss and soil seed dynamics of the dominant Bulnesia retama (Zygophyllaceae) shrub in a sandy Monte desert of western Argentina

In deserts and semi-deserts dominant plants seldom rely on a persistent seed bank to ensure their recruitment from sexual reproduction, which is generally based on the seeds produced in the last reproductive event. We studied pre- and post-dispersal seed loss of Bulnesia retama, a dominant shrub of the sandy Monte deserts (western Argentina). We hypothesised that pre- and post-dispersal seed depletion in B. retama regulate soil seed density and in turn decrease soil seed reserves. We also hypothesised that spatial patches induced by standing vegetation and other microsites additionally affect post-dispersal seed loss and soil seed density.We observed that as much as half the total seed production was lost due to pre-dispersal seed predation. Seed dispersal plus predation showed a negative exponential function with 61% of seeds falling during the first 28 days, and 99% after 90 days, when 34% of the seeds remained inside excluded branches. A significant patch effect was observed on soil seed density and seed dynamics, with a greater number of seeds and increased seed depletion rates below B. retama shrubs. However, spatial patches did not affect seed germination although they increased with burial time, from 22% to 70% after 75 and 276 days, respectively. Likewise, a similar trend was found in potential seed germination; seeds collected directly from mother plants had a low germination rate (less than 25%), which increased to 75% after 276 days of soil burial. About 15% of the dispersed seeds should persist to the next rainy season, thus contributing to a low-density but persistent seed bank.The delay in seed germination and temporal decrease in soil seed density that we observed for B. retama suggest that a set of mechanisms have been developed promoting germination events whenever soil water is available, at any moment of the year.     

Seed germination responses of Periploca sepium Bunge, a dominant shrub in the Loess hilly regions of China

Experiments were conducted to determine the effects of temperature, light, osmotic stress, hydration-dehydration regime and stress relief on the seed germination of Periploca sepium Bunge (Chinese silk vine Asclepiadaceae), which is a native shrub in the Loess Plateau of Northwest China. Freshly harvested seeds germinated equally well in both light and darkness. Seeds germinated under all of tested temperature regimes, and much faster when temperature was maintained at 30, 35, and 30/20, 30/25, 35/25, 35/30 °C range, with more than 90% of seeds germinating within 2 days. Moderate osmotic stress did not inhibit germination at any tested temperature. At 30 °C, seed germination was not affected significantly above −0.8 MPa. The hydration-dehydration pretreatment showed none side effects on P. sepium seeds and the seeds germinated more rapidly as soon as optimal water condition was restored. These results show the advantages of P. sepium, which is a competitive and widespread shrub in dry areas, and provide useful information for vegetation restoration in these drought-prone regions.     

Factors controlling the regenerative cycle of Thymus funkii subsp. funkii in a semi-arid gypsum steppe: A seed bank dynamics perspective

We researched the soil seed bank dynamics of Thymus funkii subsp. funkii, a gypsovag shrub endemic to the southeastern Iberian Peninsula, to determine possible reasons for low recruitment in isolated populations growing on crusted gypsum soils. Recruitment-restriction hypotheses involving main factors in the seed bank dynamics were analysed, and a diagrammatic dynamics model drafted from a population perspective. Viable seed production averaged 362 per plant. Seed shadows resulting from primary dispersal were concentrated under the mother plant. There were two thyme-seed predators: coleopteran larvae (pre-dispersal phase) and Messor bouvieri ants (pre- and post-dispersal phases), which predated 16.4% and 3.8% of population yield, respectively. However, ants may simultaneously contribute to the spatial secondary dispersal, since they lost a small fraction of seeds they harvested. Seeds showed innate physiological dormancy to high summer temperatures, although this disappeared rapidly. The response of germination to other temperature × light conditions was wide and rapid. No permanent seed bank was detected in the soil. Seeds buried in the upper soil layer (2 cm) germinated highly and fast. In contrast, a significant fraction of seeds buried deeply (8 cm) remained viable for over one year. Germination in the field accounted for only 1.0% of the seed output, and seed death due to pathogen attack and aging was also low (3.0%). Seedling emergence was confined to autumn and showed a high correlation with total plant cover, irrespective of plant species. Seedling survival was very low. The small spatial correspondence between primary dispersal and seedling emergence patterns may reflect the importance of secondary dispersal in T. funkii seed bank dynamics. Hypotheses and consequences of such horizontal seed movements for conservation management are discussed.     

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.     

Mucilaginous seed coat structure ofCarrichtera annuaandAnastatica hierochunticafrom the Negev Desert highlands of Israel, and its adhesion to the soil crust

Carrichtera annuaandAnastatica hierochunticaare two desert annuals, the seeds of which are dispersed by rain. The seed coat surface and cross-section structure of the dry seeds of these plants were examined by SEM. The seeds were wetted by placing them on natural saturated desert soil crust in Petri dishes. The Petri dishes were placed in a freeze-dryer when the upper part of the seed surface was still dry. This enabled study by SEM of the developmental stages of the mucilage, from the first stage until the full formation of the mucilage. We argue that the ecological significance of the mucilage differs in winter and summer. In winter the main effect is in water retention and soil particle contact, while in summer dew may allow damage repair and priming for the germination process.     

Desert pavement development and landscape stability on the Eastern Libyan Plateau, Egypt

Desert pavement surfaces of the eastern Libyan Plateau in central Egypt represent a stable landscape preserving Middle and Upper Paleolithic artifacts. Detailed measurements of pavement clasts indicate significant variability in clast size, density, lithology and orientation between pavements, but no spatial relationship among any of these pavement variables over the study area. Pavement characteristics are unrelated to local geomorphic features including slope gradient and aspect, suggesting a desert pavement surface that has developed without significant influence from transporting mechanisms such as overland flow and slope failure. Meridional vertical cracks in surface clasts implicate thermal stresses due to diurnal solar variation as a mechanical weathering process, whereas the presence of a clast-free silty layer within all soil profiles indicates that these are accretionary pavement surfaces that have grown upward over time. The desert pavement in this region has likely developed in situ through mechanical breakdown of surface clasts and desert pedogenesis, indicating long-term stability for this region and minimal taphonomic effects on artifacts > 2 cm in diameter deposited on this surface over the last ca. 100 ka.     

Soil disturbance by native animals along grazing gradients in an arid grassland

Domestic grazing animals that congregate around watering points in arid rangelands create clearly-defined trampling-induced grazing gradients. Grazing and trampling alter soil and vegetation condition, often leading to substantial reductions in ecological function. We measured foraging pits and mounds created by native soil foraging animals over 12 months at three watering points in a Chihuahuan Desert grassland, and hypothesized that the density and cover of their disturbances would increase with increasing distance from water. We recorded an average of 3756 disturbances ha⁻¹ and cover of 34.18 m² ha⁻¹ across the grazing gradients, which comprised mainly pits (43%) and mounds (25%) of heteromyid rodents, ants and spiders. Soil turnover was estimated at 1.43 m³ ha⁻¹. We detected no differences in density, cover, soil volume or composition of disturbances in relation to distance from water, but there were significant, though ill-defined, differences across the five sampling periods, with generally more activity in the warm–wet months. Small animal-created mounds and pits are important sources of soil and sinks for litter within grazing gradients, and may represent the only sites where plants can establish given a relaxation in grazing pressure.     

Factors influencing seed germination of Cyperus capitatus, inhabiting the moving sand dunes in southern Europe

Cyperus capitatus Vandelli (Cyperaceae) is distributed in coastal sandy habitats and mobile dunes of south Europe. Its seed germination ecology is not known, despite its potential to be used in re-vegetation projects. Laboratory experiments were conduced to assess the effects of salinity, light regime, cold stratification and burial on seed germination of this species. Overall, increasing salinity delayed germination, increased seed dormancy and mean time to germination (MTG), and reduced final germination percentage, inhibiting it completely above 1% of salinity; although it did not affect seed viability. C. capitatus seeds exhibited their greatest germination at levels between 0 and 1% in non-stratified seeds, and between 0 and 0.5% for stratified seeds. Thus, the effect of salt was greater for stratified seeds at 5 °C. Germination in light/darkness conditions was similar to that in full darkness. Finally, burial in sand of C. capitatus seeds appeared to have a significant effect on cumulative percentage of germination. Seeds buried at depths greater than 2 and 3 cm showed a lower germination success than those on sand surface or buried at shallower depths. Burial also affected the beginning and speed of seed germination.     

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.     

Invasive potential of the winter grass Schismus barbatus during the winter season of a predominantly summer-rainfall desert in Central-Northern Monte

Biological invasions are a major threat for natural ecosystems. Deserts are characterized by their scarce, highly variable rainfall and thus considerably susceptible to invasions, especially by annual plants that can quickly capitalize sporadic rainfall events. In this study we evaluated the potential of an invasive winter-annual species native to the Mediterranean region (Schismus barbatus) to invade a summer-rainfall desert in the Central-Northern Monte Phytogeographical Province, Argentina, where the native annual flora is mainly composed of summer species. We recorded the number of germinated seeds from soil samples collected in four seasons (during one year) coinciding with four important life-stages of Schismus (pre-germination, post-germination I and II and post-dispersal), and contrasted our results with data obtained from the literature. Almost 33,000 Schismus seeds were estimated to germinate per m², and about 90% percent of them germinated from samples collected in the pre-germination stage. Compared to deserts in USA and Australia, where Schismus has been a successful invader, the density of germinated seeds we observed was especially high even though the amount of rainfall for the vegetative period of Schismus was considerable lower. Our results suggest that S. barbatus has a great invasive potential in the Central-Northern Monte. The low number of annual plant populations that are able to compete for highly variable cool-season rains may be a key feature determining the invasibility of these deserts during the winter.