Seed dynamics of resprouting shrubs in grassy woodlands: Seed rain,predators and seed loss constrain recruitment potential

Abstract Measuring the fate of seeds between seed production and seedling establishment is critical in understanding mechanisms of recruitment limitation of plants. We examined seed fates to better understand the recruitment dynamics of four resprouting shrubs from two families (Fabaceae and Epacridaceae) in temperate grassy woodlands. We tested whether: (i) pre‐dispersal seed predation affected seed rain; (ii) post‐dispersal seed predation limited seed bank accumulation; (iii) the size of the seed bank was related to seed size; and (iv) viable seeds accumulated in the soil after seed rain. There was a distinct difference in seed production per plant between plant families with the legumes producing significantly more seeds per individual than the epacrids. Seed viability ranged from 43% to 81% and all viable had seed or fruit coat dormancy broken by heat or scarification. Pre‐dispersal predation by Lepidopteran larvae removed a large proportion of seed from the legume seed rain but not the epacrids. Four species of ants (Notoncus ectatomoides, Pheidole sp., Rhytidoponera tasmaniensis and Iridomyrmex purpureus) were major post‐dispersal seed removers. Overall, a greater percentage of Hardenbergia (38%) and Pultenaea (59%) seeds were removed than the fleshy fruits of Lissanthe (14%) or Melichrus (0%). Seed bank sizes were small (<15 seeds m^?2) relative to the seed rain and no significant accumulation of seed in the soil was detected. Lack of accumulation was attributed to seed predation as seed decay was considered unlikely and no seed germination was observed in our study sites. Our study suggests that seed predation is a key factor contributing to seed‐limited recruitment in grassy woodland shrubs by reducing the number of seeds stored in the soil.     

Episodic recruitment of the seedling banks in balsam fir and white spruce

• Premise of the study: In ecosystems where seed production is low and masting years are sporadic, or with species that have short-lived seeds, regeneration is assured by seedling banks rather than seed banks. Seedling establishment and survival play a critical role in determining the composition of these plant communities by supplying new individuals for their maintenance. Seedling emergence and mortality were investigated to test the hypothesis that recruitment into the seedling bank is periodic. • Materials and methods: Seed production and seedling emergence and survival was monitored during 1994-2007 in balsam fir (Abies balsamea) and white spruce (Picea glauca) in four pristine stands of the boreal forest of Quebec, Canada. Measurements were collected twice per month by sampling one permanent plot of 20 × 20 m per stand. • Key results: Seed-rain abundance reached 9 × 10³ seeds m⁻² year⁻¹, and was characterized by synchronous sequences of low and high seed production. New seedlings appeared only during the year following a seed production of at least 1 × 10³ and 1.5 × 10² seeds m⁻² year⁻¹ for balsam fir and white spruce, respectively. Seedlings emerged in July and survived 34-52 d on average, with balsam fir showing a longer lifespan and lower mortality, although 85–99% of seedlings died before completing one year of life. • Conclusions: The emergence of young seedlings was coupled with massive seed rains, which allowed synchronous replenishment of the seedling banks among stands and species, and generated different cohorts, yielding a discontinuous age structure.     

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     

The invasive potential of Australian banksias in South African fynbos: A comparison of the reproductive potential of Banksia ericifolia and Leucadendron laureolum

Abstract Several taxa of the Australian Proteaceae have invaded South African fynbos and require costly management programmes to bring under control. Banksia spp. have been introduced only recently to fynbos regions. The invasive potential of Banksia ericifolia (Proteaceae) was investigated by comparing its recruitment potential with that of an indigenous proteaceous shrub, Leucadendron laureolum. Both species are overstorey shrubs that are killed by fire and rely on canopy-stored seeds (serotiny) for recruitment. Eight year old B. ericifolia shrubs produced an average of 16 500 seeds per plant, which is thirty times more than the average of 570 seeds produced by 10 year old L. laureolum shrubs. The seed bank of B. ericifolia was not only larger than that of L. laureolum (1098 vs 525 viable seeds m^?2 projected canopy cover), but also considerably larger than that described for the species in its native environment (200–330 seeds m^?2 in a 9 year old stand north of Sydney). Leucadendron laureoleum released most of its seed a few days after the cones were burnt, whereas seed release in B. ericifolia was spread over 12 weeks. The seeds of B. ericifolia had lower wingloading and fall rates than L. laureolum and were dispersed over greater distances. The relative seedling growth rates of the two species were very similar (0.03 g per day), but below-ground biomass was greater and proteoid roots were more developed in B. ericifolia seedlings than in L. laureolum after 100 days. Four year old B. ericifolia plants growing in the field had attained over twice the height of indigenous pro-teoids and accumulated up to 10 times the fresh biomass of L. xanthoconus, a species which is ecologically similar to L. laureolum. The Bioclimatic Prediction System (BIOCLIM) was used to create a bioclimatic profile of B. ericifolia and identify climatically suitable areas in the Cape Province. Results show that its potential distribution covers most fynbos areas in the southwestern Cape. It is concluded that B. ericifolia has the potential to be highly invasive in fynbos.     

Seed and microsite limitation in Rheum nobile,a rare and endemic plant from the subnival zone of Sino-Himalaya

Background: Species persistence, particularly in monocarpic species, depends on the successful recruitment of individuals. An understanding of the factors that limit the recruitment of rare monocarpic plant species is therefore vital for their conservation.

Aims: To identify the factors limiting the recruitment of Rheum nobile, a rare and highly specialised monocarpic giant herb endemic to the high eastern Himalayas.

Methods: Seed sowing (seeds added or not added) and seedling transplanting experiments were conducted in disturbed (vegetation removed) and undisturbed plots in the vicinity of established populations of R. nobile to explore the mechanisms of recruitment limitation. Four levels of photosynthetically active radiation (0, 15, 30 and 50 μmol m^?2 s^?1) and two sowing positions (beneath and above grass litter or moss layer) were manipulated in the laboratory to determine how ground cover limited seedling emergence.

Results: Seed addition increased seedling recruitment. Disturbance significantly increased seedling emergence and establishment. Seed germination significantly decreased with the reduction of light availability, but 31.7% of all seeds germinated in complete darkness. Seedling emergence was close to zero when seeds were sown on top of a layer of grass litter or moss, but rose to 34.5% when the seeds were sown beneath such layers.

Conclusions: Our results indicate that the recruitment of R. nobile is limited by a combination of seed and microsite availability. Therefore, in order to conserve this species, we suggest adding seeds to suitable sites and implementing soil disturbances in existing populations to create suitable microsites.     

Early postfire seed dispersal, seedling establishment and seedling mortality of Pinus coulteri (D. Don) in central coastal California, USA

Seedling recruitment in many highly serotinous populations of Pinus coulteri on California's central coast depends almost entirely on periodic, stand-replacing fire. Compared to serotinous pines of the Mediterranean Basin, little detailed information is available on the postfire demography of California closed-cone pines, including P. coulteri. In September 1996 a wildfire burned the 760-ha American Canyon Research Natural Area (RNA). Using aerial photography, we mapped burn severity of P. coulteri-chaparral woodlands and forests within the RNA. From May to September of 1997, we also quantified seedling establishment and mortality in relation to biophysical site characteristics including fire severity. Seventy-six percent of P. coulteri forests and woodlands experienced high-severity burns, 9% moderate-severity burns, and 15% low-severity or unburned. Of the 53 plots used for seedling counts, 70% were high-severity, 26% moderate-severity, and 4% low-severity. Seedling densities 13 months postfire were low (0.21 m^–2), but seedling mortality also was low (8.4%). Aerial seed bank size increased from north-facing to south-facing slopes and from high-severity to low-severity burns. Seedling recruitment was unrelated to burn severity and increased with the size of the canopy seed bank (cone density). Many seedlings established from rodent seed caches; 23% of the seedlings established in clumps from seeds cached by Dipodomys agilis, Chaetodipus californicus and Peromyscus maniculatus. Pinus coulteri seeds have low potential for dispersal by wind, but secondary dispersal by rodents moves seeds away from source trees and into neighboring chaparral. We discuss the potential importance of rodent seed caching to postfire demography of California and Mediterranean serotinous pines.     

Buried seed population in the herbaceous lomas on Loma Ancon in the coastal desert of central Peru

Herbaceous lomas in the Peruvian coastal desert, of South America establish in spring, and its habitat is limited to the southern or southwestern slopes along the coast that are affected by thick fog. The time of appearance, the duration and the thickness of the fog vary greatly from year to year, so the lomas can grow only in habitats with enough water to, sustain seed germination and plant growth. This paper studies the species composition and density of the buried seed population, of the herbaceous lomas of Loma Ancon in order to clarify the mechanisms of the lomas' establishment. The mean number of species with viable seeds was about, 12 spp. m⁻² and that of dead seeds was about 22 spp. m⁻². The dominant species wereSolanum tuberiferum, S. pinnatifidum andNolana humifusa, both in viable and dead seeds. Viable seed density was about 5000–8000 seeds m⁻², which is comparable with the seed densities of other herbaceous communities. Dead seed density was about 15000–27000 seeds m⁻², or nearly three times the viable seed density, because the rate of decomposition was slow in the extremely dry conditions. The net increase of viable seeds by seed production was estimated at about 5000 seeds m⁻² in 1980, and the increase in the number of dead seeds was 2200 seeds m⁻².     

Lantana camara and the fruit- and seed-damaging fly,Ophiomyia lantanae (Agromyzidae): Seed predator,recruitment promoter or dispersal disrupter?

The seed damaging fly, Ophiomyia lantanae is a widespread agent that infests the fruits and seeds of Lantana camara. Its effectiveness in reducing seed dispersal or seedling recruitment has not been comprehensively tested. We determined the effects of O. lantanae damage on fruit removal rates of L. camara in a field study. We also investigated recruitment responses (seed germination in growth cabinets and seedling emergence in pot trials) of two common L. camara biotypes, pink and pink-edged red (PER) to O. lantanae damage. We found that fewer damaged fruits were removed than undamaged fruits, suggesting that frugivores select against damaged fruit. In all recruitment experiments, the responses to damage were biotype dependent. In germination experiments, these differential responses were primarily due to significantly reduced germination of damaged PER seeds. Seedling emergence from pots of the PER biotype was also lower in damaged seeds (27.5%) than undamaged seeds (56.1%). In contrast, emergence increased with seed damage in the pink biotype, from 28.2 to 41.1%. O. lantanae damage significantly reduced seed mass only in the smaller seeded PER biotype. In the larger seeded pink biotype, it may have promoted seedling emergence by interfering with dormancy mechanisms. We also examined in situ seed banks at L. camara populations prior to peak seed production. Mean viable seed densities ranged from 78.6 to 402.8 seeds m⁻². Of these, non-dormant seeds comprised 24.6 to 98.2 seeds m⁻², suggesting that recruitment is unlikely to be seed limited. We conclude that while O. lantanae influences recruitment and dispersal processes, the magnitude of the responses measured are unlikely to greatly influence plant densities in south-east Queensland infestations.     

Effects of plant competition,seed predation,and nutrient limitation on seedling survivorship of spotted knapweed (<Emphasis Type="Italic">Centaurea stoebe</Emphasis>)

We measured seed germination and seedling survivorship of spotted knapweed, Centaurea stoebe, in a series of laboratory and field experiments to evaluate the efficacy of seed limitation as a management focus. This work was initiated 6 years after introduction of several biological control agents. The soil seed bank of the site used in this study contained a mean density of 5,848 seeds/m² (ranging from 0 to 16,364 seeds/m²), and 92% of the seeds isolated from soils were shriveled, discolored, and/or partially decayed. Additionally, none of the intact seeds germinated, suggesting that the viable seed bank at our field study site has been exhausted. Centaurea stoebe seeds were planted into pots under a range of soil nitrogen (N) availability, with half of the pots containing a single density of previously established seedlings of a native cool-season grass, slender wheatgrass (Elymus trachycaulus). A watering regime mimicking local precipitation was applied. Spotted knapweed exhibited large biomass responses to N addition, but the presence of grasses suppressed the ability to exploit this N. Surprisingly, low soil N conditions improved knapweed survivorship in the presence of grasses. Nevertheless, recruitment and biomass were still far below the levels reached in the absence of competition. To evaluate the effect of density on successful recruitment, Centaurea stoebe seed was introduced into a meadow at three densities matching reduced levels of seed production under the constraints of seed predators. These densities were sown with or without a seed mixture of native species, into an existing plant community lacking C. stoebe, and seedling recruitment was recorded over 2.5 years. Across all plots and densities sown (568–2,272 seeds m⁻² year⁻¹), seedling recruitment was less than 1%. The invasion potential of spotted knapweed was greatly diminished when realistic levels of plant competition and biological control limit seed production. We therefore conclude that a combination of seed limitation and shortage of ‘safe sites’ within undisturbed vegetation can limit densities of C. stoebe.     

Dispersal and recruitment dynamics in the fleshy‐fruited Persoonia lanceolata (Proteaceae)

Question: What is the role of dispersal, persistent soil seed banks and seedling recruitment in population persistence of fleshy‐fruited obligate seeding plant species in fire‐prone habitats? Location: Southeastern Australia. Methods: We used a long‐term study of a shrubby, fleshy‐fruited Persoonia species (Proteaceae) to examine (1) seed removal from beneath the canopy of adult plants; (2) seedling recruitment after fire; (3) the magnitude and location of the residual soil seed bank; and (4) the implications for fire management of obligate seeding species. We used demographic sampling techniques combined with Generalised Linear Modelling and regression to quantify population changes over time. Results: Most of the mature fruits (90%) on the ground below the canopy of plants were removed by Wallabia bicolor (Swamp wallaby) with 88% of seeds extracted from W. bicolor scats viable and dormant. Wallabies play an important role in moving seeds away from parent plants. Their role in occasional long distance dispersal events remains unknown. We detected almost no seed predation in situ under canopies (< 1%). Seedling recruitment was cued to fire, with post‐fire seedling densities 6‐7 times pre‐fire adult densities. After fire, a residual soil seed bank was present, as many seeds (77‐100%) remained dormant and viable at a soil depth where successful future seedling emergence is possible (0‐5 cm). Seedling survival was high (> 80%) with most mortality within 2 years of emergence. Plant growth averaged 17 cm per year. The primary juvenile period of plants was 7–8 years, within the period of likely return fire intervals in the study area. We predicted that the study population increased some five‐fold after the wildfire at the site. Conclusions: Residual soil seed banks are important, especially in species with long primary juvenile periods, to buffer the populations against the impact of a second fire occurring before the seed bank is replenished.     

The dispersal of a dry-fruited shrub by red deer in a Mediterranean ecosystem

Seed dispersal of dry-fruited shrubs has received little attention in Mediterranean areas despite their frequency in the vegetation and the consideration given to the dispersal of fleshy-fruited shrubs in the area. Red deer faeces has recently been found to contain large numbers of seeds from one of the most common shrubs of this group, gum cistus Cistus ladanifer, although its importance in the reproduction of the species is unknown. This study examines the role of the red deer as an effective disperser of C. ladanifer. For this purpose, we quantify i) the C. ladanifer seed content in red deer dung over a year, ii) the seed shadow generated by the red deer wish their faeces during the same period, and iii) the ability of the dung-borne seeds to germinate and establish as seedlings under field conditions within five years following excretion. The results reveal an extremely high seed Content of the species in red deer dung (up to 80.5 ± 41.9 germinable seeds g^-1), which is virtually confined to the summer (July–August), when we estimate that a red deer defecates up to 24 000 seeds of the species per day. Furthermore, red deer mainly deposit gum cistus seeds amongst plant formations lacking the species: over the year, red deer excrete <2600 seeds m^-2 in C. ladanifer-dominated scrub and 7400–8800 seeds m^-2 in other plant formations. Under natural conditions, the dung-borne seeds have a more sniggered among-years germination pattern than free seeds in the soil. Though no seedling survived its first summer drought, the survival of seedlings sprouted from dung was significantly longer than that of control seedlings in the first and third years after deposition, and indistinguishable from it the second, fourth and fifth years. This is the first quantification of the importance of red deer to C. ladanifer dispersal and establishment, and suggests that endozoochory by mammalian herbivores can be very valuable for dry-fruited shrubs in the Mediterranean region.     

Factors affecting seedling recruitment in an invasive grass (Pennisetum setaceum) and a native grass (Heteropogon contortus) in the Hawaiian Islands

In the Hawaiian Islands, native Heteropogon contortus (pili grass) is being replaced by alien grasses, one of which is Pennisetum setaceum (fountain grass). Both grasses depend on seeds for population growth. To help understand factors promoting the spread of the alien and decline of the native, we investigated the effects of physical disturbance, nutrient addition, and seed supplementation on seedling recruitment in experimental field plots. In the first year, our field site experienced an unusual drought, and seedling recruitment was greater for H. contortus than for P. setaceum under all treatments. Disturbance increased recruitment of H. contortus seedlings during some sampling periods. Recruitment was not significantly increased by seed additions for either species despite our finding of only 49 and 4 seeds m^–2 in the seed bank for H. contortus and P. setaceum, respectively. In the first year, most P. setaceum seedlings died between monthly surveys. We resurveyed our field plots in a second, wetter year and found the pattern was reversed: recruitment of P. setaceum seedlings was greater than H. contortus seedlings in most treatments. Greenhouse comparisons of seedling survival under three drought regimes (water every 5,7 and 10 days) revealed that H. contortus seedlings tolerate drought better than P. setaceum seedlings. Seedling recruitment for these species in the leeward Hawaiian Islands appears to be primarily dependent on water availability, with the alien having the advantage in wetter years. Once seedlings of the long-lived alien become established, the alien seems capable of maintaining its dominance over H. contortus, even during periods of drought.     

SEED BANK,SURVIVORSHIP, AND SIZE DISTRIBUTION OF A NEBRASKA POPULATION OF IMPATIENS CAPENSIS (BALSAMINACEAE)

The growth and survivorship of a population of Impatiens capensis were followed in eastern Nebraska. Soil cores were taken within the population to estimate the size of the seed bank. The number of seeds in the soil remained relatively constant from April through October, with a mean of 200 seeds/m². The seed bank size corresponded to observed seedling densities in the following years. Twenty permanent plots were established within the population and censused for three years, 1983–1985. Seedling recruitment to the population was similar in 1983 and 1984 but 30% higher in 1985. The seedling densities of plots were uncorrelated from year to year. Survivorship in 1983 followed a Deevey Type I pattern. Almost one-third of the population survived until chasmogamous reproduction. Plant size (stem height) was measured monthly on all individuals during 1983. The frequency distribution of stem height was positively skewed as long as growth was exponential. When the growth rate slowed, the distribution became more normal.     

Effects of seed dispersal on Juniperus communis recruitment on a Mediterranean mountain

Abstract. The recruitment of the relict shrub Juniperus communis on a mountain in SE Spain was studied during the period 1994–1998. The main objective was to determine both the quantitative and qualitative effects of bird dispersal on seedling establishment. Seed removal by birds, seed rain, post‐dispersal seed predation, germination, and seedling emergence and survival were analysed in different microhabitats. Birds removed 53 ‐ 89% of the seeds produced by plants. Seed rain was spatially irregular as most seeds accumulated near stones used by birds as perches and below mother plants while a few seeds were dropped in wet meadows and open ground areas. Post‐dispersal seed predation by rodents affected < 10% of dispersed seeds but varied significantly among microhabitats. Only 3.6 ‐ 5.5% of dispersed seeds appeared viable, as many seeds had aborted or showed wasp damage. Seeds germinated in the second and third springs after sowing, reaching a germination percentage of 36%. Seedling emergence was concentrated in wet meadows. Seedling mortality was high (75–80%), but significantly lower in wet meadows, the only microhabitat where seedlings could escape from summer drought, the main mortality cause. Seed abortion, germination and seedling mortality proved to be the main regeneration constraints of J. communis on Mediterranean mountains. Birds exerted a strong demographic effect, although their qualitative effect was limited by abiotic factors which caused the pattern of seed rain to differ from the final pattern of recruitment between microhabitats.     

Leaf litter of <Emphasis Type="Italic">Kageneckia angustifolia</Emphasis> D. Don (Rosaceae) inhibits seed germination in sclerophyllous montane woodlands of central Chile

Leaf litter accumulation can have either positive, negative or neutral effects on seed germination and seedling recruitment. In montane woodlands of the Mediterranean zone of central Chile, large amounts of leaf litter accumulate beneath the crowns of the summer semi-deciduous tree Kageneckia angustifolia and no regeneration of this or other plant species has been observed beneath this tree throughout the year. In a sample plot of 5000 m² we selected ten K. angustifolia trees and measured (1) leaf litter accumulation beneath and outside canopy; (2) the effects of time elapsed since burial on viability of K. angustifolia seeds with and without a leaf litter cover; (3) field seed germination with presence or absence of leaf litter and (4) the possible chemical effects of K. angustifolia leaf litter leachates on seed germination of its own seeds and of other two co-occurring native shrubs species (Guindilia trinervis and Solanum ligustrinum). Our results show that a considerable accumulation of leaf litter occurred beneath K. angustifolia, and litter negatively affected seed viability and germination of this species in the field. Under laboratory conditions, K. angustifolia leaf litter leachates inhibited seed germination of its own seeds and of the two native shrub species. Chemical effects are likely involved in the negative effects of leaf litter on the recruitment of K. angustifolia in the montane sclerophyllous woodland of central Chile.     

Seedling bank dynamics of Fagus grandifolia var. mexicana before and after a mast year in a Mexican cloud forest

Abstract. Fagus grandifolia var. mexicana seedling dynamics were studied in a relict forest inside the crater of a volcano where it is the only canopy tree species. The objectives of the study were (1) to determine changes in seedling density before and after a mast year, as well as to monitor seedling recruitment, mortality and growth rate in relation to the micro‐environment, and (2) to discuss the impact of masting on seedling dynamics of the population. Before masting, seedling density was 8.8 seedlings m^–2, afterwards seedling density peaked at 51.5 seedlings m^–2, then decreased by 50% in 3 months and by 85% after 8 months, down to levels of before the mast year (9.9 seedlings m^–2) after 16 months. Seedlings in the forest floor before the mast‐seeding event showed a low relative growth rate – 0.14 mm mm^–1 mo^–1 against 1.27 mm mm^–1 mo^–1 for seedlings that emerged immediately after masting. Seed germination was high (83%) and rapid (L₅₀= 7 days) following masting. Seedling growth was positively correlated with soil water content but not with temperatures and relative humidity. Data suggested that seed production in mast years is important in maintaining the seedling bank, and thus the viability of relict Fagus populations.     

How important is seed predation to recruitment in stable populations of long-lived perennials?

Summary The many evolutionary modifications to seed biology in response to seed predation do not necessarily imply that seed predators have an important impact on population recruitment. This is because competition between individual plants for rare safe sites can cause an oversupply of seeds so far as a population is concerned. The importance of seed losses to population recruitment at any point in time is related to the abundance of safe sites (Fig. 1): it is zero when safe sites are absent, negligible when safe sites are rare, and greatest when safe sites are numerous enough for recruitment to be limited by seed supply. Here I interpret the impact of severe seed losses on population recruitment in four species of long-lived perennials (Eucalyptus baxteri, Leptospermum juniperinum, Casuarina pusilla and L. myrsinoides) by considering these losses in terms of the overall seed dynamics of the populations. I focus on seed supply and seedling survival, as a measure of the current abundance of safe sites, and the maintenance of seed banks, as a measure of the ability of populations to exploit any future changes in safe site abundance. Insect seed predators destroyed about 95% of total seeds in each case. However, these losses do not necessarily have an important impact on population recruitment, because: (i) in most years recruitment appears to be limited by a rarity of safe sites, and not by seed supply (which was as high as 43 germinable seeds/m²/yr); and (ii) the losses did not prevent the establishment of large seed banks (ranging from >30 to >1000 viable seeds/m²) potentially capable of exploiting temporary conditions favourable for recruitment. In contrast to the situation with many annual plants, patterns of recruitment in stable populations of long-lived perennials are often extremely complex, and the significance of seed losses therefore difficult to determine.     

Arbuscular mycorrhizal fungi affect seedling recruitment: a potential mechanism by which N deposition favors the dominance of grasses over forbs

Background and aims

Nitrogen (N) deposition usually alters plant community structure and reduces plant biodiversity in grasslands. Seedling recruitment is essential for maintaining species richness and determines plant community composition. Arbuscular mycorrhizal fungi (AMF) are widespread symbiotic fungi and could facilitate seedling establishment. Here we conducted an experiment to address whether the influence of AMF on seedling recruitment depends on N addition and plant species.

Methods

Leymus chinensis were cultivated for 5 months in the microcosms that were inoculated with or without AMF at five N addition rates. Seeds of three main species (two C₃ grasses and one non-N₂-fixing forb) of the Eurasian steppe were sown to the 5-month-old microcosms. Seedling establishment was estimated by shoot biomass, N and P contents 7 weeks after seedling germination.

Results

AMF promoted seedlings recruitment of two C₃ grasses at addition rates above 0.5 g N m^?2. In contrast, seedling recruitment of the non-N₂-fixing forb was increased by AMF at addition rates below 0.5 g N m^?2 but was decreased above 2.5 g N m^?2.

Conclusions

These results partly explain why N addition favored the dominance of grasses over forbs in perennial grassland communities. Our study indicates that AMF have the potential to influence plant community composition by mediating revegetation in the face of N deposition.     

Recovery of aSasa tsuboiana population after mass flowering and death

The recovery process of aSasa tsuboiana population after a mass flowering and death in 1977 was investigated by 15 years of observation in the Hira Mountains, Kinki district, western Japan. Seed production was high (6600–13 800 seeds m⁻² inSasa plots and 3900 seeds m⁻² in a forest plot) but emergent seedling density was low (14–21 seedlings m⁻²), probably because of seed predation byMicrotus montebelli occurring between seed shedding and the next spring. The seedling density had decreased further by the next year and theS. tsuboiana population recovered from only a limited number of seedlings. In spite of such a low initial density, theS. tsuboiana population was able to regenerate successfully and attained the previous full stand height in 7–16 years.Miscantbus sinensis invaded and delayed the recovery ofS. tsuboiana in one plot, butS. tsuboiana became dominant as it caught up with the height ofM. sinensis. Seedling growth patterns, such as frequent tillering, the onset of rhizome extension in the early stage of seedling growth and frequent culm production from rhizomes, played important roles in the successful regeneration ofS. tsuboiana.     

Effect of seed predation on seed bank size and seedling recruitment of bush lupine (Lupinus arboreus)

Whether seed consumers affect plant establishment is an important unresolved question in plant population biology. Seed consumption is ubiquitous; at issue is whether seedling recruitment is limited by safe-sites or seeds. If most seeds inhabit sites unsuitable for germination, post-dispersal seed consumption primarily removes seeds that would otherwise never contribute to the population and granivory has minimal impacts on plant abundance. Alternatively, if most seeds ultimately germinate before they lose viability, there is greater potential for seed consumption to affect plant recruitment. Of the many studies on seed consumption, few ask how seed loss affects seedling recruitment for species with long-lived seed banks. We examined post-dispersal seed predation and seedling emergence in bush lupine (Lupinus arboreus), a woody leguminous shrub of coastal grasslands and dunes in California. We followed the fate of seeds in paired experimental seed plots that were either protected or exposed to rodent granivores in grassland and dune habitats. Significantly more seeds were removed by rodents in dunes than grasslands. In dunes, where rodent granivory was greatest (65% and 86% of seeds removed from plots by rodents in two successive years), there is a sparse seed bank (6.6 seeds m⁻²), and granivory significantly reduced seedling emergence (in the same two years, 18% and 19.4% fewer seedlings emerged from exposed versus protected plots), suggesting seed rather than safe-site limited seedling recruitment. In contrast, rodents removed an average of 6% and 56% of seeds from grassland plots during the same two years, and the grassland seed bank is 43-fold that of the dunes (288 seeds m⁻²). Even high seed consumption in the second year of the study only marginally influenced recruitment because seeds that escaped predation remained dormant. Burial of seeds in both habitats significantly reduced the percentage of seeds removed by rodents. Results suggest that granivores exert strong but habitat-dependent effects on lupine seed survival and seedling emergence. Received: 24 October 1996 / Accepted: 4 February 1997