Establishment and growth of seedlings of Eucalyptus obliqua: Interactive effects of litter,water, and pathogens

We studied how leaf litter, water and fungal pathogens affect the establishment, survivorship, and growth of Eucalyptus obliqua using glasshouse and field experiments. In a glasshouse experiment, the presence of leaf litter of E. obliqua increased the establishment of seedlings when the seeds were placed beneath the litter, but dense litter layers reduced seedling emergence when seeds were placed on top. Results from a complementary experiment suggest that the effect of litter may be mediated by the high humidity microenvironment it creates, which may enhance imbibition of unburied seeds. In another glasshouse experiment, litter in combination with excessive water applications reduced seedling emergence and increased seedling mortality, but the addition of fungicide reverted these effects. Litter with lower water application or fungicide added increased seedling emergence. In all cases higher biomass in seedlings growing in litter treatments was associated with earlier emergence. In two recently burnt sites, the density of seedlings of E. obliqua was positively correlated to the presence of litter. Experimental addition of litter in the field reduced soil temperature during the summer, and together with watering, increased soil water content, and the water potential of established seedlings. However, neither treatment had any positive effect on their growth or survivorship. On the contrary, litter addition reduced the height and number of leaves, probably as a result of increased herbivory. We conclude that in Mediterranean-type systems litter may produce various effects on seedling establishment depending on their developmental stage, and the season.     

Effect of grass litter on seedling recruitment of the critically endangered Cistus heterophyllus in Spain

Cistus heterophyllus Desf. is an Iberian-north African endemic plant species with only one wild population left in Spain. A sowing experiment under controlled conditions was designed to assess both the seedling emergence rate and the growth fitness of C. heterophyllus on a litter layer of Brachypodium retusum (Pers.) Beau., which forms dense carpets around the remnant population of C. heterophyllus. In order to test the physical effects of B. retusum litter on Cistus seedling recruitment, six sowing treatments in pots were compared that combined three doses of litter (0, 0.6 and 1.1 kg m⁻²) and different physical locations of the seeds (above, below the litter layer or within a litter–peat mixture). B. retusum litter interfered with the seedling recruitment of C. heterophyllus, negatively affecting both seedling emergence and their early growth. Such an effect was more severe when the seeds were located above a 0.6 kg m⁻² litter layer, although seedlings from seeds sown under the litter showed a growth reduction ranging from 58% to 100% compared to the no-litter treatment. A significant negative effect of the litter–peat mixture on the number of leaves, leaf size and number of branches of seedlings was also inferred. These data provide experimental evidence on the serious threat that B. retusum litter represents for the conservation and regeneration of the last population of C. heterophyllus in Spain.     

Seedling recruitment of Succisella inflexa in fen meadows: Importance of seed and microsite availability

Abstract. The effects of different forms of land use on germination and establishment of the rare fen species Succisella inflexa were investigated in seed introduction experiments in a mown and an abandoned fen meadow in SE Germany. Treatments included abandonment, mowing in fall and mowing with creation of gaps in the moss and litter layer. Floating capacity of seeds was tested in order to estimate potential dispersal by water. On the mown meadow, gaps had a slightly positive effect on germination rates, while greatly increasing seedling survival until the next spring. At the abandoned site, litter inhibited germination, whereas mosses had a negative effect on germination and a positive effect on survival rates during the first year after germination. Both germination and seedling establishment were negatively affected by the presence of slug herbivores. On the abandoned site, no seedlings at all survived until the next spring. Even though seeds of Succisella inflexa were capable to float for several weeks and to germinate thereafter, the situation at the field sites indicates that longdistance dispersal is highly unlikely. Our results showed that not only direct effects of abandonment, such as accumulation of litter, may have led to poor germination and poor seedling establishment of the species. Additionally, indirect consequences of changes in land use, such as higher seedling herbivory by slugs and successional vegetation changes due to abandonment, were important in determining habitat quality and availability of microsites for seedling recruitment. Furthermore, early mowing imposed seed limitation on plant populations.     

Germination requirements and responses to leaf litter of four species of eucalypt

We studied how the responses of four species of eucalypt to leaf litter related to their germination responses to light and water availability. Two of the species (Eucalyptus obliqua and E. baxteri) have a mesic distribution, while the other two (E. oleosa, and E. incrassata) are more xerophytic. We studied the effect of litter on emergence of the four species in a glasshouse experiment. Litter did not affect the emergence of E. incrassata and E. oleosa, but enhanced the emergence of E. obliqua and E. baxteri. Litter increased the seedling mortality of all four species. Germination responses to light and water availability were studied in growth cabinets under controlled conditions. The germination of E. obliqua and E. baxteri was substantially lower under fluorescent light than in darkness, but that of E. oleosa and E. incrassata was not affected by the light environment. The germination of E. obliqua and E. baxteri was significantly reduced by reduced water potential (a). Substantial germination of E. oleosa and E. incrassata occurred even at very low a (less than –1.05 MPa). We conclude that both the shade and the humid micro-environment provided by litter may have contributed to the emergence responses of the four species to litter, and these responses may correspond to ecological adaptations to the different environments in which they live.     

Importance of seed size for the establishment of seedlings of five deciduous broad-leaved tree species

Effects of seed size and phenology on the establishment of five deciduous broad-leaved tree species were examined in deciduous woodland. Treatments included absence and presence of litter in the forest understory, a small gap, and a large gap. Seedling emergence of large-seeded speciesQuercus mongolica var.grosseserrata andAcer mono was not reduced by accumulation of litter in the forest understory, but was promoted in the large gap where litter was less. Seedling emergence of small-seeded species,Alnus hirsuta, Cercidiphyllum japonicum andBetula platyphylla var.japonica, was reduced by the litter in almost all of the sites. Seedlings of large-seeded species avoid shade stress phenologically by unfolding all of their large leaves in a short period before canopy closure in the forest understory. These species had little mortality after seedling emergence. In contrast, small-seeded species have a longer duration of leaf emergence, shorter leaf longevity, and rapid leaf turnover in all the sites. These seedlings attained similar height to those of the large-seeded species at the end of the second year in the large gap, but survival and height growth rate decreased after canopy closure in the forest understory. We suggest that the importance of seed size in determining seedling establishment largely depends on the relationships between seasonal changes of environmental conditions and phenological traits of seedlings, which are related to seed size.Abbreviations Ah Alnus hirsuta - Am Acer mono - Cj Cercidiphyllum japonicum - Bp Betula platyphylla var.japonica - Qm Quercus mongolica var.grosseserrata     

Rodent seed predation and seedling recruitment in mesic grassland

Seedling recruitment of two grasses (Arrhenatherum elatius and Festuca rubra) and two herbs (Centaurea nigra and Rumex acetosa) was measured in areas with and without rodents to which seeds of each species were sown at three seed densities (1000, 10,000 and 50,000 seeds m⁻²) in two seasons (spring and autumn 1995). Seed removal was measured for 10-day periods and the fate of seedlings was followed for 15 months after sowing. The proportion of seed removed ranged from 6 to 85% and increased with increasing seed density for each species. Rodents had no effect on seedling emergence or survival in the spring sowing. In the autumn sowing, rodents reduced seedling emergence of all four species sown at 1000 and 10,000 seeds m⁻² but had no impact at 50,000 seeds m⁻², presumably because of microsite limitation. We suggest the difference between spring and autumn arose because emergence was seed limited in autumn but microsite limited in spring; microsite availability was higher in autumn because a summer drought killed plants, reduced plant biomass and opened up the sward. Fifteen months after the autumn sowing, fewer A. elatius and C. nigra seedlings survived on plots exposed to rodents. This result reflected not only the reduced seedling emergence but also increased seedling mortality (seedling herbivory) in sites exposed to rodents. In contrast, F. rubra and R.acteosa showed density-dependent seedling survival which compensated for initial differences in seedling emergence, so that no effect of rodents remained after 15 months. The results suggest that rodent seed predation and seedling herbivory exert strong effects on seedling recruitment of A.elatius and C. nigra when recruitment conditions are favourable (conditions that lead to high microsite availability) and may contribute to both species being maintained at low densities in the grassland. The results also demonstrate that highly significant impacts of rodent seed predation at the seedling emergence stage can disappear by the time of plant maturation. Received: 2 March 1998 / Accepted: 28 September 1998     

Fire and seedling population dynamics in western Oregon prairies

Abstract. Through an experiment in three prairie vegetation types in western Oregon, USA the effect of prescribed fire on the timing and rates of seedling emergence and mortality was examined. Seeds of common exotic and native prairie species were sown into burned and unburned plots in late September, 1995. Emerged seedlings were censussed the following winter, early spring and late spring. Results indicated that spring population levels could not be forecast by fall seedling flushes, as winter survival was important in seedling establishment. The bulk of emergence for all grass and annual forb species occurred in the fall, followed by low to severe winter mortality. Perennial forbs were more variable in emergence times but, once emerged, perennial forb seedlings were likely to become established. Burning caused a statistically significant increase in seedling accumulation through emergence and survival in 11 of 23 cases. Burning improved seedling winter survival for most grass and short‐lived forb species and increased emergence of perennial forb species. These patterns were most conspicuous on the two sites dominated by exotic species, where burning significantly improved the accumulation of seedlings from most native species tested. Thus, prescribed burning might be a useful restoration tool in these communities. In contrast, two of the three species increased by burning in the native bunchgrass site were exotic pest plants, suggesting that fire should be prescribed with caution.     

The coupling of recruitment and disturbance by fire in two resprouting Proteaceae species

Recruitment in plant populations is often tightly coupled to major disturbances such as fires. For species with persistent seed banks, fire-related cues may allow or enhance germination. The litter layer influences germination and may modify the impact of seed predators on seeds and seedlings. The litter layer is obviously affected by fire, providing one mechanism by which disturbance can determine recruitment. We tested the role of litter in the disturbance–recruitment coupling of two species with contrasting seed release timing after fire—Banksia serrata (canopy seed bank) and Telopea speciosissima (transient seed bank) by planting their seeds both early and late in the post-fire recruitment period (PRP) and manipulating litter density in orthogonal treatments. Vertebrate seed predators were excluded. Both species established more seedlings late in the PRP, although results were strongly influenced by very poor establishment at one site. Invertebrate seed predators consumed more T. speciosissima seeds in sites early (69.5%) than late in the PRP (51.2%), while consumption of B. serrata seeds was lower overall and comparable across sites (average 47.3%). Surprisingly, litter had very little effect on establishment and none on invertebrate seed predation, suggesting that other factors are more important. Recruitment was only loosely coupled to disturbance for the canopy seed bank species; for the transient seed bank species, the coupling was tighter but separated in time from the disturbance. Understanding both the strength and temporal aspects of the disturbance–recruitment coupling is necessary for appropriate management of plant functional diversity in fire-prone habitats.     

The role of seed depth,litter, and fire in the seedling establishment of amphicarpic peanutgrass (Amphicarpum purshii)

Summary Amphicarpum purshii is an annual grass which mostly grows in disturbed areas of the New Jersey Pine Barrens, USA. It is amphicarpic, producing spikelets (and seeds) both above and below the soil surface. Previous research has shown that subterranean seed production ensures reproduction in the event of a major disturbance such as fire and results in rapid post-burn colonization of these sandy habitats. The effects of fire, litter, and seed depth were further examined by planting subterranean seeds at four depths in 16 litter-covered flats buried at ground level and comparing plants arising from burned flats with those in undisturbed litter-covered flats. At 0 and 1 cm depth, rates of seedling emergence were lowest in burned flats. Surface-sown seeds produced seedlings more likely to desiccate. Sowing depth had a greater influence on most measured characters than burning treatments. The mean depth of subterranean seed placement by Amphicarpum is 3.5 cm and this coincides with the seed depth from which plants showed the greatest height growth, shoot biomass, and reproductive output. In a second experiment, subterranean seeds on the bare soil surface in clay pots were more likely to lose viability and less likely to germinate than seeds protected by litter or burial in soil. In addition to providing protection from fire, placement of seeds below ground in the sandy habitat of peanutgrass provides conditions more suitable for seed survival and subsequent seedling establishment.     

Patch colonisation in a savanna grassland

Patches (1 m diameter) were cleared in a heavily grazed and a lightly grazed savanna in South Africa; half of them were covered once with grass litter. The greatest colonisation over three years occurred on heavily grazed patches with litter, the least on open, lightly grazed patches. Annual rainfall affected colonisation rate. Basal cover was lower on patches than in the surrounding vegetation. Patch composition was weakly affected by treatment, and was partly related to the composition of adjacent vegetation, but the proportional representation of species on patches differed from the surrounding vegetation. The stoloniferous Digitaria eriantha and the obligate seed reproducers Aristida bipartita and Heteropogon contortus were major components of patch communities but were uncommon in the surrounding vegetation and in the seed bank. Setaria incrassata and Themeda triandra were the predominant components of the surrounding vegetation and of the seed bank, but Setaria established very poorly and Themeda established well only on heavily grazed patches with litter. The pattern of seedling establishment was the same as that of colonisation. Most seedlings emerged atthe beginning of the wet season, with ca. 50% mortality soon thereafter. The colonising species can be partly predicted from the availability of propagules (vegetative or seed), emergence and establishment success, and subsequent growth.     

Bottlenecks and spatiotemporal variation in the sexual reproduction pathway of perennial meadow plants

Sexual reproduction is important for the growth of populations and the maintenance of genetic diversity. Several steps are involved in the sexual reproduction pathway of plants: the production of flowers, the production of seeds and the establishment of seedlings from seeds. In this paper we quantify the relative importance and spatiotemporal variability of these different steps for four grassland perennials: Centaurea jacea, Cirsium dissectum, Hypochaeris radicata and Succisa pratensis. We compared undisturbed meadows with meadows where the top soil layer had been removed as a restoration measure. Data on the number of flower heads per flowering rosette, the numbers of flowers and seeds per flower head, and the seedling establishment probabilities per seed were collected by field observations and experiments in several sites and years. Combination of these data shows that H. radicata and S. pratensis have higher recruitment rates (1.9 and 3.3 seedlings per year per flowering rosette, respectively) than the more clonal C. dissectum and C. jacea (0.027 and 0.23, respectively). Seedling establishment is the major bottleneck for successful sexual reproduction in all species. Large losses also occurred due to failing seed set in C. dissectum. Comparison of the coefficients of variation per step in space and time revealed that spatiotemporal variability was largest in seedling establishment, followed closely by flower head production and seed set.     

The importance of seed reserves for seedling performance: an integrated approach using morphological,physiological, and stable isotope techniques

To investigate how seed reserves affect early seedling performance, we conducted a factorial greenhouse experiment using Lithocarpus densiflora (Tanoak). Seedlings were grown from large (5.8±0.7 g) and small (3.2±0.4 g) seeds and, following shoot emergence, seeds were either removed or left attached. Seedlings were harvested for quantification of biomass and ¹³C at seven time periods following seed removal (2, 4, 8, 16, 32, 64, 128 days) and seedling photosynthesis was measured three separate time periods (2–4, 49–82, 95–128 days after seed removal). Biomass increased for all seedlings, but the increase was significantly larger for seedlings with attached seeds than with removed seeds. Seed removal just after shoot emergence significantly decreased seedling biomass, but seed removal 64 days after shoot emergence had no effect on seedling biomass. Seedling photosynthesis per unit leaf area varied by time and seed presence, but not by seed size. At the first period, seedlings with attached seeds had significantly higher photosynthetic rates than seedlings with removed seeds, at the second period there was no effect of seed removal, and at the third time period seedlings with attached seeds had significantly lower photosynthetic rates than seedlings with removed seeds. Despite temporal variation in photosynthesis per unit leaf area, seedlings with attached seeds always had significantly greater leaf area than seedlings with removed seeds, resulting in significantly higher total plant photosynthesis at all three time periods. The ¹³C values of both the leaves and roots were more similar to that of the seed for seedlings with attached seeds than for seedlings with removed seeds, however, seed removal and seed size strongly affected root ¹³C. This study demonstrates that seed reserves have important effects on the early growth, physiology, and ¹³C of L. densiflora seedlings.     

Germination and early growth of <Emphasis Type="Italic">Ailanthus</Emphasis> and tulip poplar in three levels of forest disturbance

Increasing rates of forest disturbance may provide greater opportunity for invasion of nonnative species, thereby altering the successional trajectory of native plant communities. In the eastern U.S., invasive Ailanthus altissima and native Liriodendron tulipifera have similar life histories and niches and often co-occur. To examine how disturbance affects the establishment of these species, we performed field experiments to evaluate the response of sown seeds and transplanted seedlings to three levels of disturbance on north- and south-facing aspects. L.␣tulipifera germination was severely limited by low seed viability and had significantly lower germination than A. altissima in all sites. The effect of disturbance regime on A. altissima germination depended on aspect in the second growing season. In contrast, mean seedling survival, biomass, leaf area and leaf area ratio were greater for L. tulipifera in all field sites. Overall, the north-facing selective cut forest provided a disproportionately large number of suitable microsites for L. tulipifera establishment. Collectively, this study demonstrated that different timber harvest practices produce heterogeneous mosaics of suitable microsites for germination and establishment. Limited L. tulipifera germination may be a serious constraint to population establishment if seeds are deposited for the first time immediately after a disturbance event. However, if sufficient viable seeds of both species exist, L. tulipifera out-performs the invasive in the first two years following disturbance. This may explain why A. altissima has shown explosive population growth in a limited number of sites in the past century.     

An invasive frog, Eleutherodactylus coqui, increases new leaf production and leaf litter decomposition rates through nutrient cycling in Hawaii

A frog endemic to Puerto Rico, Eleutherodactylus coqui, invaded Hawaii in the late 1980s, where it can reach densities of 50,000 individuals ha⁻¹. Effects of this introduced insectivore on invertebrate communities and ecosystem processes, such as nutrient cycling, are largely unknown. In two study sites on the Island of Hawaii, we studied the top-down effects of E. coqui on aerial, herbivorous, and leaf litter invertebrates; herbivory, plant growth, and leaf litter decomposition rates; and leaf litter and throughfall chemistry over 6 months. We found that E. coqui reduced all invertebrate communities at one of the two study sites. Across sites, E. coqui lowered herbivory rates, increased NH₄⁺ and P concentrations in throughfall, increased Mg, N, P, and K in decomposing leaf litter, increased new leaf production of Psidium cattleianum, and increased leaf litter decomposition rates of Metrosideros polymorpha. In summary, E. coqui effects on invertebrates differed by site, but E. coqui effects on ecosystem processes were similar across sites. Path analyses suggest that E. coqui increased the number of new P. cattleianum leaves and leaf litter decomposition rates of M. polymorpha by making nutrients more available to plants and microbes rather than through changes in the invertebrate community. Results suggest that E. coqui in Hawaii has the potential to reduce endemic invertebrates and increase nutrient cycling rates, which may confer a competitive advantage to invasive plants in an ecosystem where native species have evolved in nutrient-poor conditions.     

CONTRASTING GERMINATION AND SEEDLING GROWTH OF BETULA ALLEGHANIENSIS AND RHUS TYPHINA SUBJECTED TO VARIOUS AMOUNTS AND TYPES OF PLANT LITTER

Several characteristics of the environment, such as plant litter, may interact with species characteristics, such as seed size, to determine patterns of plant establishment and thereby influence abundance and spatial location of plants long after the factor has itself disappeared. To determine if litter might differentially affect seedling establishment of Betula alleghaniensis Britton and Rhus typhina L., two woody species with different sized seeds, we conducted two greenhouse experiments. We documented emergence, morphology, and growth patterns of seedlings covered by three amounts each of Fagus grandifolia Ehrh. leaves or Tsuga canadensis L. needles. All amounts of litter significantly reduced and delayed final Betula alleghaniensis seedling emergence, and the resultant seedlings had reduced root: shoot ratios, longer hypocotyls, and were less robust. Litter did not affect Rhus typhina emergence but did alter biomass allocation: stem, leaf, and root biomass of Rhus seedlings was reduced by greater amounts of litter, and by leaf litter relative to needle litter. We attribute the different responses of Betula and Rhus to differences in seed size (Betula mean seed size = 1.0 mg; Rhus mean seed size = 8.5 mg) and germination cues (Betula requires light, Rhus requires chemical or heat scarification), and suggest that differential response to litter may contribute to coexistence among these sympatric species.     

Germination and population structure of spear thistle Cirsium vulgare in relation to experimentally controlled sheep grazing

Summary The size and number of rosettes of Cirsium vulgare were censused in a 4 ha sheep grazing trial on lowland calcareous grassland in paddocks receiving controlled winter, spring and summer grazing treatments. Spring grazing significantly increased thistle rosette numbers, and there were always fewer rosettes in size classes 250mm–300 mm under the heavier grazing treatments. The emergence of seeds of Cirsium vulgare sown by hand into each of the grazing treatments was monitored and showed a positive effect of spring grazing. When the percent emergence of sown seeds was used as a covariate in the analysis of rosette numbers in the experimental paddocks, it was found to account for 77% of the variance in thistle numbers. It was concluded that the main effect of grazing on thistle rosette numbers is an indirect one exercised through the effects of grazing on germination conditions. Seeds of Cirsium vulgare were sown into artificial gaps at another grassland site, to determine the effect of gap-size upon seedling emergence. Germination was poor, but significantly more seedlings emerged in gaps 10–20 cm diameter, than in 5 cm gaps or in control plots without a gap. A computer simulation model was used to explore the relationship between gap density and thistle population dynamics. A threshold density of gaps was found to exist, below which thistle populations went extinct, and above which the thistle population grew geometrically. The degree of aggregation of dispersed seeds did not alter the threshold gap-density for plant extinction, but did affect the rate of increase of the thistle population when the threshold gap-density was exceeded. It is concluded that strategies for controlling Cirsium vulgare populations by grazing manangement will be most effective if aimed at reducing suitable sites for establishment in spring. The success of such attempts will depend upon the soil fertility of the site, and sudden outbreaks of Cirsium vulgare infestation can be expected if a sward gradually deteriorates through over-grazing.     

Factors affecting seed germination and seedling establishment of a long-lived desert shrub (<Emphasis Type="Italic">Coleogyne ramosissima</Emphasis>: Rosaceae)

Long-lived desert shrubs exhibit infrequent, episodic recruitment from seed. In spite of this long time scale, selection on life history attributes that affect seedling recruitment should be strong. We studied factors affecting germination phenology and seedling establishment for Coleogyne ramosissima, a dominant shrub species in the ecotone between warm and cold deserts in western North America. We also examined ecotypic differentiation in establishment strategy in response to selection regimes in two contrasting habitats. We followed patterns of dormancy loss, germination, emergence, and survival in reciprocal field experiments at warm winter Mojave Desert and cold winter Colorado Plateau study sites. Seed germination took place in late winter, under winter rain conditions at the warm desert site and under snow at the cold desert site. Distinctive germination phenologies for the two seed populations at contrasting field sites followed predictions based on laboratory germination experiments. There was no seed bank carryover across years. Seedling survival at the end of three growing seasons was remarkably high (mean survival 54%). Most seedling mortality was due to sprout predation by rodents early the first spring in unprotected caches. Emergence and establishment at each site were significantly higher for seeds from the local population, supporting the idea of ecotypic differentiation in establishment strategy. Establishment success was an order of magnitude greater overall at the Colorado Plateau site, which represents the leading edge of an upward elevational shift in distribution for this species under the current climatic regime. The Mojave Desert site is on the trailing edge of this shift, and recruitment there is apparently a much less frequent occurrence.     

Effects of experimental and real land use on seedling recruitment of six fen species

Seedling emergence and early establishment of six fen species differing in seed mass and growth form were investigated under experimental land use with changed vegetation structure and under real land use in a calcareous fen. Seeds of all six species were sown in plots with different experimental land-use treatments: summer and autumn mowing with or without litter removal, trampling and abandonment. Additionally, emergence and survival of experimentally sown seeds was investigated under real land use on adjacent sites managed by mowing, grazing, intense trampling or abandonment.On abandoned plots and on plots without litter removal of the land-use experiment, emergence rates of all species were negatively affected either by high litter and moss cover or by tall canopy. No differences were found between autumn and summer mowing. Gap creation by experimental trampling did not increase germination rate. Under real land use, establishment of seedlings of most species was positively affected by litter cover and tall canopy. Trampling, in contrast, had a severe negative effect on seedling survival.The investigated species differed in their germination ability which was tested in the germination chamber and in their response to land use. Succisa pratensis with the highest seed mass germinated well in the chamber and in the field more or less regardless of land use. The low germination rate of Parnassia palustris in the germination chamber indicated a limitation of viable seeds. In the field, however, seedling emergence was additionally limited by microsite availability. Seeds of Serratula tinctoria and Primula farinosa germinated well in the germination chamber, but seedling recruitment in the field was hampered in the presence of a high litter or moss cover. Seeds of Tofieldia calyculata and Pinguicula vulgaris were strongly dependent on the availability of suitable microsites in the field. They hardly germinated under natural conditions, in spite of a high number of germinable seeds in the germination chamber.     

东祁连山灌丛凋落叶水浸提液对垂穗披碱草种子萌发和幼苗生长的影响

为了解灌丛凋落叶在灌草群落结构维持中可能存在的潜在化感作用，以东祁连山3种优势灌木和灌下优势草种垂穗披碱草(Elymus nutans)为供试材料，分析了不同浓度(0.01 g/L、0.025 g/L、0.05 g/L、0.075 g/L、0.1 g/L)的金露梅(Potentilla fruticosa)、川滇柳(Salix rehderiana)、头花杜鹃(Rhododendron capitatum)及灌间草本的凋落叶水浸提液，对垂穗披碱草种子萌发、幼苗形态和生理指标的影响。结果表明：(1)垂穗披碱草种子发芽率与发芽势在金露梅与川滇柳凋落叶浸提液处理下呈“低促高抑”的浓度效应，在灌间草本凋落叶浸提液处理时表现为先升后降，而头花杜鹃凋落叶浸提液处理时均下降，并在浓度超过0.075 g/L时种子不发芽。(2)灌丛凋落叶浸提液对垂穗披碱草幼苗根长与干重均表现出抑制作用，且随浓度增大而增强；金露梅、川滇柳与灌间草本凋落叶浸提液对垂穗披碱草幼苗苗高表现出“低促高抑”的浓度效应；而头花杜鹃凋落叶浸提液处理对垂穗披碱草幼苗鲜重与苗高均呈抑制作用。(3)垂穗披碱草幼苗CAT、SOD活性随着浸提液...     

Flooding and grazing promote germination and seedling establishment in the perennial grass Paspalum dilatatum

Seed germination and seedling emergence are key processes for population recruitment. Flooding and grazing are disturbances forming gaps that may strongly influence recruitment patterns in space and time, but their combined effects and action mechanisms have rarely been addressed. In this study we analysed the effects of microhabitat conditions associated with winter flooding and spring‐summer defoliation on seed germination and seedling establishment of Paspalum dilatatum, a dominant perennial C₄ grass in native grasslands of the Flooding Pampa, Argentina. The dynamics of seedling emergence from natural seed banks and buried seeds was studied in a factorial experiment with flooding and defoliation treatments applied to soil monoliths (mesocosms) collected from natural grassland. Additional laboratory experiments were applied to investigate seed germination under different combinations of temperature, light quality and simulated flooding. Seed germination and seedling emergence of P. dilatatum were promoted by flooding and high intensity defoliation. Gaps generated by flooding were maintained by high intensity defoliation exercising a synergistic effect on survival seedlings. Flooding resulted in the breaking of seed dormancy and higher germination rates associated with alternating temperature and the activation of the phytochrome system. Our results indicate that microhabitat conditions associated with the disturbances forming gaps, such as flooding and heavy grazing, synergistically promote the recruitment process of this dominant grass species.