The role of fire in preventing transitions from a grass dominated state to a bush thickened state in arid savannas

We investigated the role of fire in controlling bush thickening in arid savannas. Three controlled fires were initiated to investigate the responses of planted seedlings, saplings and mature shrubs. The stem diameters of seedlings, saplings and mature shrubs prior to the fire were measured. Other dimensions of the saplings and mature shrubs such as leaf mass were also estimated. The intensity of each fire was estimated. Fire temperatures adjacent to seedlings saplings and mature shrubs were measured using fire paints on steel plates. All three fires were moderately hot. One year after each fire, the mortality of seedlings in the fire treatments was very high (97.1–99.3%) and significantly greater than in adjacent controls (16.1–51.6%). Mortality and topkill of saplings and mature shrubs were negatively related to stem diameter. Fire temperature did not significantly affect mortality and topkill in most cases. Surviving saplings showed a much greater resprout response, relative to pre fire size, than larger shrubs and trees. Our results suggest that fire is crucial in interrupting the transition from open grassy savanna to thicket in arid savannas. Managers who prevent fires at this stage are likely to experience bush thickening in the future.     

Water requirements for germination and early seedling establishment in four African savanna woody plant species

The aim of this study was to determine the water requirements for germination and early seedling establishment of four African savanna tree species, namelyAcacia karroo, A. nilotica, A. tortilisandMundulea sericea. The acacias are characteristic of nutrient-rich, andM. sericeaof nutrient-poor savannas. Imbibition times of scarified seeds ofA. karroo(4 h),M. sericea(6 h) andA. tortilis(8 h) were rapid relative toA. nilotica(28 h) and were inversely correlated with seed size. Imbibed water is lost in about 2 h after drying at 25°C for all species exceptA. nilotica(only 70% moisture loss). Seeds ofA. karrooandM. sericeakept at 40°C dried to below their original moisture contents within 2 h. Seeds ofA. niloticaandA. tortilisdried at 40°C lost viability relative to (undried) control seeds, while there was no significant loss of viability forA. karrooandM. sericea. For fully imbibed seeds to germinate in sandy savanna soils, all four species required at least the equivalent of 3 mm rainfall every 2 days under mild greenhouse conditions. However, most ungerminated seeds were still viable despite drying-out from a fully imbibed state. Maintenance of soil at 50% field capacity (FC), or watering to FC every 9th day is the maintenance requirements for 2-week-old seedlings ofA. niloticaandM. sericeato continue growth for a further 5 weeks under greenhouse conditions, althoughA. niloticamostly survived 25% FC with one seedling surviving 12·5% FC. Root penetration was rapid in sandy soils, withA. niloticaandM. sericeaattaining a depth of 40 cm within 15 days. Frequent, but not necessarily high, rainfall appears to be essential for germination and seedling survival over the first 7 weeks.     

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.     

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.     

Seed germination of five Helianthemum species: Effect of temperature and presowing treatments

The germination behaviour of five Helianthemum species (H. almeriense, H. appeninum, H. cinereum, H. hirtum, H. squamatum) has been studied under controlled conditions. Constant 15, 20 or 25 °C and alternating 25/15 °C temperature regimes and 16/8 h light/dark photoperiod conditions were used. Presowing treatments applied were manual scarification, boiling water, hot water, dry heat and sulphuric acid. Germination values recorded were final germination percentage and germination rate expressed as days to reach 50% of the final germination percentage (T₅₀). Incubation temperature had no significant effect on final germination percentage for untreated seeds of the five Helianthemum species. However, variation due to temperature was significant for scarified seeds, with the lowest germination percentage attained at 25 °C. In all Helianthemum species studied, the highest germination percentages were obtained by manual scarification of seeds. Germination rate of scarified seeds decreased as germination temperature increased. The different presowing treatments investigated allowed some germination in some species, but none were any better than manual scarification. The high germination among most species studied, following mechanical rupture of the seed coat, shows that the mechanism of dormancy lies in the seed coat. The physical dormancy caused by impermeable seed-coat appears to be the main reason of poor germination of untreated seeds of Helianthemum species studied.     

Soil seed flora, germination and regeneration pattern of woody species in an Acacia woodland of the Rift Valley in Ethiopia

The objectives of this study were to determine the composition, density and spatial distribution of the soil seed bank of woody species, as well as their regeneration pattern in two different land use systems, controlled (ranch) and open grazing, in an Acacia woodland of the Rift Valley in Ethiopia. We also compared the species composition of the soil seed bank and the above-ground vegetation to find out if differences exist in the soil seed bank and advance regeneration between the two land use systems. The germination requirements of seeds of the woody species were also investigated under laboratory conditions. Acacia senegal, Acacia seyal, Acacia tortilis, Dichrostacys cinerea and Balanites aegyptiaca were encountered in the above-ground vegetation in both systems. Seeds of only A. tortilis (90±32 seeds m⁻²) from the ranch, and A. senegal (5±3 seeds m⁻²) and A. tortilis (72±34 seeds m⁻²) from the open system were found in the soil seed bank along transects with patchy horizontal pattern. The two systems were not significantly different in density of soil seed banks of A. senegal andA. tortilis . Jaccard's similarity index showed that only a few woody species were common in the soil seed flora and above ground vegetation. However, all of the species accumulated seeds (58±43–331±130 seeds m⁻²) in/on the soil under the canopy. Very large numbers of seeds of A. tortilis (19382±9722 seeds m⁻²) and D. cinerea (1278±494 seeds m⁻²) were also found in barns. Most of the seeds recovered from the soil samples (60–80%) were found in the litter layer. Acid and mechanical scarification improved legume germination (36–99% and 60–99%, respectively) over boiling water (0–48%). Treatment means differed significantly for all the legumes (p<0·001) but not for Balanites. Height and diameter class distribution of regeneration of A. tortilis and D. cinerea in both systems and A. senegal in the open system had a negative exponential correlation (r_s=−0·5, −0·25 and −0·86, respectively). A. seyal and B. aegyptiaca showed poor regeneration. Horizontal distribution of advance regeneration of all the species was patchy. Advance regeneration of A. seyal, A. tortilis and B. aegyptiaca were not significantly different, while that of A. senegal and D. cinerea were significantly different between the two systems. Poor representation of species in the soil seed bank along transects and in the different height and diameter classes may be attributed to the low density of mature trees as well as the mode and strategy of seed dispersal. Ungulate and wind dispersed species (e.g. A. senegal, A. tortilis and D. cinerea) were highly favoured. Patchiness in the distribution of seeds and advance regeneration was also a result of endozoochory. Dispersal of non-ungulate dispersed seeds (A. senegal, A. seyal and B. aegyptiaca) was restricted to the canopy zone. Piles of seeds ofA. tortilis and D. cinerea that were found in barns were a result of consumption of their pods by cattle. High concentration of seeds in the litter layer may be due to low soil disturbance and larger size of seeds. The height and diameter class distribution of A. senegal (in the open system), A. tortilis and D. cinerea also indicated that the species have good regeneration. Results from the germination tests indicated that seeds of the legumes require pre-sowing treatments to give a rapid, uniform and improved germination. Intervention through artificial regeneration should be employed to improve the density and regeneration capacity of those species with hampered regeneration at both systems.     

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.     

Germination ofAcacia origena,A. pilispinaandPterolobium stellatumin response to different pre-sowing seed treatments, temperature and light

Seeds ofAcacia origena,A. pilispinaandPterolobium stellatumhave a hard seed coat that hampers imbibition of water and prevents maximum, uniform and rapid germination. Therefore, the seeds were subjected to different treatments involving mechanical scarification, concentrated sulphuric acid and boiling water to break the dormancy imposed by the hard seed coat. To determine the effects of different constant temperature regimes on germination, scarified seeds were incubated on a thermogradient at 10, 15, 20, 25 and 30°C. Germination of scarified seeds was also tested in light and dark. Mechanical scarification resulted in 100% germination for theAcaciaspecies and 94% forP. stellatum. ForA. origena, sulphuric acid treatments for 60, 90 and 120 min resulted in significantly higher per cent germination (>95%) than all the other treatments. ForA. pilispina, sulphuric acid treatments for 45 and 60 min, as well as immersing seeds for 1 min in hot water, gave significantly higher per cent germination (97%) than all the other treatments. ForP. stellatum, mechanical scarification and all the acid treatments applied resulted in significantly higher per cent germination (>85%) than the control and all hot water treatments. All seeds ofP. stellatumimmersed in hot water, even for 1 min, rotted soon after sowing, indicating that they are sensitive to high temperatures. Scarified seeds of the twoAcaciaspecies germinated to a higher degree (≥90%) between 10 and 30°C, while germination of scarified seeds ofP. stellatumranged from 48 to 82% between 10 and 30°C. Scarified seeds of the three species germinated to a large extent (>70%) both in light and dark, indicating that seeds do not require light for germination.     

Effects of salt and alkali stress on Reaumuria soongorica germination

Seed germination and early seedling growth are crucial stages for plant establishment. Two neutral (NaCl and Na₂SO₄) and two alkali (NaHCO₃ and Na₂CO₃) salts were selected to investigate their effects on germination and recovery responses in Reaumuria soongorica. Results show that both salt types significantly reduced germination and radicle elongation. The rate of germination and emergence of R. soongorica seeds continuously decreased as salinity increased, and the time to achieve maximum germination rate was delayed. The speed of seed germination dropped rapidly as salt concentration increased. Alkaline salts restricted the germination rate of R. soongorica seeds, and stresses resulting from alkaline salts and high concentrations of neutral salts resulted in many deformed seedlings. The length of the radicle and germ decreased with increasing salt concentration, but certain concentrations of salt and increased pH promoted germ growth. The results of regression analysis show that salt concentration was the dominant factor inhibiting R. soongorica seed germination rate. Salinity, buffering capacity, and pH all affected embryo growth, but salinity had the most pronounced effect. Seed viability under highly saline conditions appears to be a better indicator of adaptation to saline environments than seed germination under saline conditions.     

Effect of sand burial depth on seed germination and seedling emergence of Calligonum L. species

A greenhouse study was conducted to determine the effects of sand burial on seed germination and seedling emergence of ten Calligonum L. species. The results suggested that the deeper the seeds in sand, the lower and slower their germination and seedling emergence. Seedling emergence occurred from a maximum depth of 12 cm with most seedlings emerging from 2 and 4 cm depths. The percent seedling emergence, number of days of first emergence, percent seed germination and percent of dormant seeds were significantly affected by different species and burial depth. There was a significant positive correlation between the number of days to first emergence and burial depth. Both mean percent germination and percent seedling emergence were negatively correlated with depth significantly for each species.     

Temporal variability in the maternal environment and its effect on seed size and seed quality in Flourensia cernua DC. (Asteraceae).

The biological phenomenon commonly referred to as maternal effect occurs whenever the environment exerts an influence upon the mother that is later expressed in characteristics of her offsprings. For example, the environmental conditions experienced by a mother plant during flowering and fruiting can modify the quality of her descendants (seeds), and even interrupt their development. Between-year variation in minimum temperature and the classic, between-year variation in precipitation represent an unpredictable environment for winter blooming plants in arid zones. In this study, we investigated the effect of maternal environment (temperature and precipitation) on seed size and seed quality in F. cernua, over a 5-year period. Results indicated that the proportion of filled seeds, as well as their size, increased with higher absolute minimum temperatures during seed formation, and that, to a lesser extent, both the precipitation occurred during the same period and the annual precipitation also have a positive influence on seed size. In this way, environmental conditions prevailing during seed formation exerted a strong maternal effect on the size and quality of seeds produced and, probably also, in the developmental possibilities of F. cernua seedlings. This effect was subject to important, between-year variation.     

大赖草(Leymus racemosus)结实格局

连续2年对新疆阿勒泰地区荒漠植物大赖草(Leymus racemosus)居群(公路附近沙丘平坦处,P1;农田周围沙丘平坦处,P2;周围积水沙丘中下部,P3;多个沙丘中间平坦处,P4;多个沙丘中间隆起处,P5)、小穗间和小穗内的结实格局进行了比较研究。结果表明:(1)2011年大赖草的穗长、穗宽、总花数、结实率和大种子比率都高于2012年,小种子数低于2012年;2年的结实率在5个居群间均表现为P2P3P1P5P4。(2)每花序内不同小穗的穗宽、总花数和结实率表现为中部下部上部。(3)每小穗均有3.7±0.56朵花、0.6±0.09粒种子,小穗基部的结实率较高。大赖草结实率在年际和不同环境下波动较大,水分和温度是影响其结实格局的主要因素;小穗间及小穗内的资源分配符合资源竞争假说。大赖草花多果少的结实格局是适应荒漠资源贫瘠、干旱少雨气候条件的一种生殖保障。     

Germination of Callitris seeds in relation to temperature, water stress, priming, and hydration–dehydration cycles

The effects of temperature, water stress, hydration–dehydration cycles and seed priming on the germination of Callitris verrucosa and Callitris preissii, two Australian semi-arid coniferous tree species, were investigated. Optimum germination occurred at 18°C, with a minimum germination time of 8–9 days for both species. At this temperature, germination was inhibited at osmotic potentials lower than −1·0 MPa, but the capacity to germinate at low osmotic stress increased as the temperature decreased. Seed priming and hydration–dehydration cycles did not reduce seed viability, and Callitris seeds appear to retain the physiological changes induced by short-term hydration, as the time to the onset of germination was decreased to about 3 days. The capacity of Callitris seeds for incremental germination is likely to increase overall germination success in a low rainfall environment.     

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.     

Rehabilitation of Rangelands in Paulshoek, Namaqualand: Understanding vegetation change using biophysical manipulations

In Namaqualand, heavy grazing of low-lying areas has resulted in the reduction of palatable perennial shrubs and the dominance of the unpalatable shrub, Galenia africana. This study investigates the process of rehabilitation and how it can be used to gain an understanding of the ecological factors that may be sustaining altered vegetation states. We intervened to change the biophysical environment through the construction of microcatchments and brushpacks to act as traps for water, seed and organic material combined with the introduction of seeds and seedlings were implemented. Two years after implementation brushpacked microhabitats improved natural recruitment of herbaceous species compared to bare areas at two out of three sites. Microhabitats did not influence natural recruitment of perennial seedlings. Emergence of palatable perennial seed was not influenced by microhabitat manipulation. Seedlings transplanted under adult G. africana shrubs showed higher survival rates at two out of three sites compared to seedlings in unmodified areas.     

Passage time, viability, and germination of seeds ingested by foxes

Seed passage through the gut of vertebrates is critical for endozoochorus seed dispersal because it may influence seed viability and germination capacity. Foxes are important fruit-eaters and presumed seed dispersers in the Chaco ecoregion. We experimentally assessed the seed passage time (SPT) and its effect on seed viability and germination in two fox species (Pseudalopex gymnocercus and Cerdocyon thous) that occur in the Chaco. Passage time was measured using both artificial seeds of three sizes and wild seeds of four native fleshy-fruit species commonly consumed by foxes (Ziziphus mistol, Acacia aroma, Celtis tala, and Syagrus romanzoffiana). The average SPT was 6.8 h for the Pampa fox and 7.7 h for the crab-eating fox. Passage time did not differ either among seeds of different sizes or between fox species. Passage through digestive tract of both fox species did not affect seed survival, whereas germination rate was slightly improved. Our data suggest that both fox species are legitimate seed dispersers of the four species studied, having a positive effect on both germination rate and dispersal distance from the parent plant.     

Invasion of the noxious weed Nicotiana glauca R. Graham after an episodic flooding event in the arid zone of Australia

During November 1997, an extreme rainfall event caused flooding in the Olary Creek and created a terminal lake within mallee vegetation on Nagaela Station, far western New South Wales. To assess the effect of this rare event on native flora, six 25×25 m² grazing exclusion plots were established on flooded (4), and un-flooded control (2) areas. We investigated (i) the distribution of Nicotiana glauca in relation to the 1997 flooding event (ii) the variation in density of N. glauca soil-stored seed between soil in control and exclosure plots (iii) the allelopathic effects of N. glauca on germination of Lactuca sativa seeds. Analysis of N. glauca stem sections revealed a maximum of five growth rings, confirming an origin following the 1997 flood event. At a September 1998 assessment, no N. glauca were found across flooded and control plots. In October 2000 N. glauca occurred at densities of 24 and 12 N. glauca seedlings ha⁻¹ in the flooded unfenced and fenced plots respectively. No N. glauca individuals were recorded either from control fenced or unfenced plots. The soil-stored seed study revealed that larger numbers of N. glauca seeds were recovered from the flooded open plot (1936±968) than in flooded fenced plots (856±428.12) but none was found in the control plots. Leachates obtained from dry leaves and twigs had a significantly greater negative impact (p>0.001) on germination of L. sativa seeds than leachates obtained from fresh leaves and bark. This study showed that the density of N. glauca is high in areas where grazing had eliminated competition from native shrubs and grasses. N. glauca is potentially a serious weed along the Olary Creek and ephemeral lake boundaries. Management strategies such as reducing grazing pressure from native and introduced animals and/or effective eradication programs should be adopted, to minimize further infestation.     

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.     

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.     

Effect of rhizobia inoculation on the seed (herbage) production of mungbean (Phaseolus aureusRoxb.) grown at Uzbekistan

In field trials conducted at Uzbekistan, inoculation of mungbean (Phaseolus aureusRoxb.) with commercial strain CIAM1901 ofBradyrhizobiumsp. (Phaseolus) increased (on average for two cultivars) the herbage mass by 46·6±6·0%, seed mass by 39·2 ± 3·6%, mass of 1000 seeds by 16·0 ± 0·8%, nitrogen content in seeds by 58·3 ± 8·9%, starch content in seeds by 30·0 ± 5·5% and number of nodules by 254%. Inoculation with this strain produced the same herbage (seed) mass as NH₄NO₃application (120 kg ha⁻¹of N), while the combined treatment of rhizobia and 60 kg ha⁻¹of N produced significantly higher yields than rhizobia inoculation alone or application of 120 kg ha⁻¹of N. The rhizobia strain M11 was isolated from Uzbekistan soils and significantly exceeded the commercial strain CIAM1901 in its influence on herbage mass by 3·9–10·6%. Two-factor analysis of variance demonstrated that the herbage mass and number of nodules are controlled mainly by the rhizobia strain genotypes, while seed yield, number of pods, mass of 1000 seeds and N and starch content in seeds are influenced by the plant cultivar genotypes.