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.     

Seed and microsite limitations of recruitment and the impacts of post-dispersal seed predation at the within population level

Seed predation may cause important seed losses in plant populations, but its impact on the dynamics of populations will depend on the degree of seed or microsite limitations for recruitment. Seed losses will only affect recruitment if it is seed limited. The nature of recruitment limitation (seeds or microsites) is usually ascribed to whole plant populations but it may vary within populations among microhabitats and habitats. Thus, the potential impact of seed predation will also vary within the population, being highest where recruitment is seed limited. The impact to the whole population will depend on the spatial concordance between the intensity of seed predation and that of seed limitation. Recruitment limitations (with seed addition experiments), seed predation (with seed removal experiments), and the dynamics of seed availability in the soil (with soil samples taken both after seed dispersal and before the following dispersal event) of the shrub Corema album (Empetraceae) were investigated in dunes in NW Spain, at microhabitats ‘open ground’, ‘underneath C. album ♀’, and ‘underneath C. album ♂’ at two habitats, sparse and dense scrub. The nature of recruitment limitation (seeds vs. microsites) varied within the population. It was seed limited in the microhabitat ‘open ground’ and microsite limited under shrub cover. The spatial patterns of seedling recruitment were unrelated to seed availability but strongly affected by germination requirements. The spatial discordance between seed availability and recruitment implies a crucial constraint for processes affecting seed availability (seed predation but also e.g., dispersal) to impact recruitment. They will not affect its spatial pattern but only its quantity as long as they act in those sites selected by seeds to germinate. Seed predation was highest underneath mother plants and lowest in open ground. Thus, its potential impact is low, as it is centred where recruitment is not seed limited. This study shows that the analysis of seed predation in relation to recruitment limitations at smaller spatial scales within the population provides more insight to understand its impact.     

Post-dispersal seed predation: consequences for plant demography and evolution

Post-dispersal seed predation is only one of many factors underlying plant demography and evolution. Nevertheless, the generalist feeding habits of many post-dispersal seed predators and the limited ability of plants either to compensate for or to respond to post-dispersal seed losses directly suggest that post-dispersal seed predation may have a considerable impact on plant populations. Seed predators probably have little direct influence on the demography of plants that regenerate exclusively by vegetative means or are buffered by a large active seed bank, but such species are only a minority in most plant communities.In general, ants are significant post-dispersal seed predators in arid and semi-arid ecosystems while they act mainly as seed dispersers rather than as predators in temperate ecosystems. Although studies have probably underestimated the importance of invertebrates and birds as seed predators, rodents appear to have greater potential to influence seed dynamics, and are particularly important in temperate ecosystems. For example, production of mast seed crops is more effective at satiating specialist invertebrate seed predators than generalist vertebrates, and recruitment may be limited by post-dispersal seed predation even during mast years.Both spatial variation in post-dispersal seed predation and differences in predation between species are important elements which facilitate the coexistence of different plant species. Where microsites are limiting, selective post-dispersal seed predators can influence pre-emptive competition for these microsites. Seed size determines the extent of density-dependent predation and the exploitation of buried seed. This suggests that post-dispersal seed predators may also play a role in the evolution of seed characteristics. However, conclusions regarding the ecological and evolutionary impact of post-dispersal seed predators will remain speculative without a more substantial empirical base.     

Microsite and litter cover effects on seed banks vary with seed size and dispersal mechanisms: implications for revegetation of degraded saline land

Seed movements and fates are important for restoration as these determine spatial patterns of recruitment and ultimately shape plant communities. This article examines litter cover and microsite effects on seed availability at a saline site revegetated with Eucalyptus sargentii tree rows interplanted with 5?C6 rows of saltbush (Atriplex spp.). As litter accumulation decreases with increasing distance from tree rows, soil seed banks were compared between paired bare and litter-covered zones within three microsites; tree row, saltbush row closest to tree row and saltbush mid-row (middle row of saltbush between tree rows). Germinable seed banks of the four most abundant species with contrasting seed sizes and dispersal mechanisms were assessed to test the hypotheses that: (i) microsites with litter cover contain higher seed densities than bare areas, but that (ii) microsite and litter effects will vary depending on seed size and dispersal mechanisms. Overall, litter cover increased seed densities, however, litter effects varied with seed size, with no effect on small-seeded species and litter increasing densities of large-seeded species. Seed bank composition also differed between tree and shrub microsites due to differences in seed morphology and dispersal mechanisms. Water-dispersed species were unaffected by microsite but densities of wind-dispersed species, including Atriplex spp., were higher in saltbush microsites. Densities of wind-dispersed species also differed between the two saltbush microsites despite similar litter cover. Future plantings should consider row spacing and orientation, as well as the dimensions of seeding mounds and associated neighbouring depressions, to maximize litter and seed-trapping by microsites.     

Habitat effects on second-order predation of the seed predator Harpalus rufipes and implications for weed seedbank management

Seed predators provide a valuable ecosystem service to farmers by reducing densities of weed seeds, and, in turn, densities of weed seedlings they must manage. The predominant invertebrate weed seed predator in Maine, USA, agroecosystems is the carabid beetle Harpalus rufipes DeGeer. Pitfall trapping has shown that H. rufipes prefers sites with vegetative cover to fallow sites, preference speculated to be driven by predator avoidance behavior. To test this hypothesis, ‘second-order predation assays’ were developed, in which live H. rufipes prey were presented to second-order predators. Field experiments were conducted to determine foremost if H. rufipes was subject to second-order predation, and secondly, whether (a) vegetative cover affords H. rufipes protection from second-order predators, and (b) high rates of second-order predation correspond with decreased invertebrate seed predation rates. Two 72-h experiments were conducted (mid August and September 2012) at crop and non-crop sites across a 28 ha diversified farm in Stillwater, ME, USA.Second-order predation was 2.8% per day. Based on images from motion-sensing cameras, H. rufipes’ predators included birds and small mammals. Neither a relationship between second-order predation and vegetative treatment, nor an empirical relationship between second-order predation and invertebrate seed predation were detected. However, a simulation model predicted that 2.8% per day second-order predation could increase the number of seeds entering the seedbank by more than 17% annually. Additionally, complex habitats supported higher rates of second-order predation than did simple habitats.     

Post‐dispersal granivory in a tall‐tussock grassland: A positive feedback mechanism of dominance?

Questions: Is post‐dispersal seed predation a factor contributing to a positive feedback mechanism for dominance of tall‐tussock grasslands? Do seeds dispersed into neighbouring microhabitats of contrasting dominance differ in their probabilities of being predated? Does predation rate vary among predator groups? Do seed eaters selectively forage among the available seeds? Location: The southern and flooding sub‐regions of the Argentinian Pampas. Methods: We examined seed predation by vertebrate and invertebrate predators within two microhabitats of grassland mosaics (highly dominated tall‐grass patches vs. scarcely dominated short‐grass matrix) for different seed species in semi‐natural grasslands. Proportion of seeds eaten by different predator groups was estimated through exclusion experiments and analysed using ANOVA for split‐split‐plot designs. Experiments were performed during the autumn of two consecutive years in both Pampa sub‐regions. Results: Removal of seeds after a five day trial was two to four times higher in the tall‐grass patches than in the short‐grass matrix. During the same period, vertebrate predation was six times higher than invertebrate predation in the tall‐grass phase of the Southern Pampa, but it did not differ in the short‐grass matrix. Relative predation among phases showed the same pattern in the Hooding Pampa, where preferences by seed species also varied according to phase. Conclusions: The highest predation intensity shown by vertebrates in the tall‐grass patches indicates that they are the main seed predators in these systems, possibly because this microhabitat grants them refuge against carnivorous predators. This could significantly reduce the available seeds for recruitment of subordinate species after different disturbance events (fires, trampling by large herbivores, burrowing), representing an active filter to the floristic composition of the patch and contributing with other mechanisms to the dominance of tall‐grass species.     

Fire and flood: Soil‐stored seed bank and germination ecology in the endangered Carrington Falls Grevillea (Grevillea rivularis,Proteaceae)

Abstract Seed germination is dependent on the interaction between the dormancy state of a seed and the presence of favourable environmental conditions. Thus, the spectacular pulse of seedling recruitment in many Australian vegetation communities following disturbances such as fire can be attributed to changes in microsite conditions and/or the dormancy‐breaking effect of the disturbance on accumulated seed banks. Grevillea rivularis is a threatened species endemic to the area immediately above Carrington Falls in the NSW Southern Highlands. Most of the population is confined to the riparian vegetation zone in woodland and heath, and is therefore subject to periodic disturbance from fire and flood. For this species, a pulse of seedling recruitment has been recorded after fire, flood and mechanical soil disturbance. The aims of this study were to examine the density and vertical distribution of the soil‐stored seed bank and to investigate the role of heat and scarification as cues for germination of fresh and soil‐stored seed. There was a large seed bank under the canopies of established individuals (194 ± 73 seeds m^?2) and most seeds were found in the 0–2 cm and leaf‐litter layers of the soil profile. The germination response of soil‐stored and fresh seed was examined using a hierarchical series of laboratory experiments. Seeds of G. rivularis showed marked dormancy polymorphism. Thirty‐six percent of soil‐stored seed germinated without treatment, whereas no untreated fresh seeds germinated. Scarification or heating caused significant germination of dormant soil‐stored seed, but only scarification resulted in germination of dormant fresh seeds. These results highlight important differences in the dormancy state of soil‐stored and fresh seed. Thus, being a riparian species in a fire‐prone environment, the dormancy mechanisms in seeds of G. rivularis suit this species to disturbance by both fire and flood.     

Experiments on the mechanism of tree and shrub establishment in temperate grassy woodlands: Seedling emergence

Field experiments were designed to examine tree and shrub seedling emergence in temperate grassy woodlands on the New England Tablelands. The effects of study sites, intensity of previous grazing, removal of ground cover by fire or clearing, burial of seeds and ant seed theft on seedling emergence were tested in two field experiments. Six tree and seven shrub species were used in the experiments and their cumulative emergence was compared with laboratory germination studies. All species used in field experiments had lower cumulative emergence than those in laboratory germination studies despite prolonged periods of above average rainfall before and after seeds were sown. Eucalypt species emerged faster in the field than the shrub species and generally attained higher cumulative emergence than the shrubs. Spatial effects of sites and patches within sites, and of previous grazing history did not strongly influence patterns of seedling emergence in most species. Ground and litter cover generally did not enhance or suppress the emergence of seedlings, although the removal of cover in recently grazed areas enhanced the emergence of some species. Burning enhanced the emergence of some tree and shrub species where plots had more fuel and intense fires, but this effect was not strong. Compared with other treatments, seedbed manipulations produced the strongest effects. In the absence of both invertebrate and vertebrate predators, seedling emergence was lower for surface‐sown seed, compared with seed sown on scarified soil surfaces. Higher seedling emergence of buried seeds in the presence of invertebrate predators probably resulted from the combined effects of predator escape and enhanced moisture status of the germination environment. Some promotion of emergence was achieved for all species in most sown treatments probably as a result of a prolonged above average rainfall. In contrast, the natural recruitment of trees and shrubs was negligible in experimental plots, highlighting the importance of seed supply and dispersal as ultimate determinants of recruitment.     

Fire survival of Cape Proteaceae-influence of fire season and seed predators

Many Cape Proteaceae store seed reserves in closed cones on the plant and rely entirely on these reserves for episodic recruitment after fires. Population size is sensitive to intervals between fires but also to fire season. Populations can be nearly eliminated by successive winter or spring fires. Three hypotheses explaining seasonal variation in recruitment were tested: (a) seeds germinate immediately after fire but seedlings die from summer drought; (b) seeds remain dormant over summer but the longer the delay between seed release after fire and germination 1) the greater the competition between seedlings and resprouts, or 2) the greater the seed losses to predators and/or decay before germination.Drought-avoiding dormancy occurred in 9 of 11 Cape Proteaceae studied, all of which delayed germination to autumn or winter. Seedling emergence and survival was not significantly increased after removal of competitors by methyl bromide poisoning. Seed predation, measured by exclosures, however, significantly reduced seed reserves before germination and also number of seedlings emerging. Post emergence seedling predation was negligible in the burn in contrast to adjacent mature vegetation where seedling predation was very heavy.The role of germination cues and rodent behaviour in controlling population recruitment is discussed and it is concluded that a knowledge of both is essential for predicting vegetation dynamics in this system.Acknowledgements: I am grateful to J. Vlok and R. America for field assistance and J. Breytenbach, F. Kruger, P. Slingsby and J. Yensch for observation and comment. This work was supported by the Directorate of Forestry, South Africa as part of their conservation research programme.     

The fear diet: Risk,refuge, and biological control by omnivorous weed seed predators

Weed seed biocontrol by omnivorous mice and insects can limit weed seedbanks, but this ecosystem service can be difficult to predict given the broad diet breadth of seed predators and their potential for intraguild predation. Seed foraging behavior is further modified by fluctuating cues of predation risk from higher trophic levels and the availability of refuge habitat. Uncertainty about whether co-occurring insects and mice additively contribute to weed biocontrol or interfere with each other via intraguild predation limits our ability to recommend habitat management strategies that reliably promote seed destruction. Using seed removal assays, fluorescent powder tracking, and stable isotope analyses, we assessed effects of a predation risk cue (moonlight) on mouse foraging patterns in a patchwork of vegetated and exposed plots in a cultivated field. Mouse foraging activity decreased on exposed ground during the full moon, compared to dark nights, yet foraging movements were unaffected by moon cycle within refuge patches. Weed seed consumption was more than three times higher in cover than exposed soil, and 78% of that difference was attributable to invertebrate granivores. Mice and invertebrate granivores both exhibited higher foraging activity in cover, indicating co-occurrence of intraguild predators and prey. However, stable isotope analyses of fecal samples revealed that mice captured in refuge habitats fed at slightly lower trophic levels than those in exposed habitats (suggesting minimal intraguild predation in refuge habitat), and mouse diet was unaffected by moonlight. Despite increased availability of invertebrate prey in cover patches, mice do not appear to preferentially exploit prey when avoiding their own predators or interfere with weed seed predation. Therefore, functional redundancy of mice and invertebrate seed predators in cover crops and other refuge habitats may strengthen and stabilize weed seed biocontrol.     

Litter effects on seedling establishment interact with seed position and earthworm activity

Seedling establishment is influenced by litter cover and by seed predators, but little is known about interactions between these two factors. We tested their effects on emergence of five typical grassland species in a microcosm experiment. We manipulated the amounts of grass litter, seed sowing position and earthworm activity to determine whether: (i) the protective effect of litter against seed predation depends on cover amount and seed sowing position, i.e., on top or beneath litter; (ii) seed transport by earthworms changes the effect of seed sowing position on seedling emergence; and (iii) seeds transported into deeper soil layers by earthworms are still germinable. Litter cover and presence of earthworms lowered seedling emergence. The impact of seed position increased with seed size. Emergence of large-seeded species was reduced when sown on the surface. Additionally, we found an important seed position × earthworm interaction related to seed size. Emergence of large-seeded species sown on top of the litter was up to three times higher when earthworms were present than without earthworms. Earthworms also significantly altered the depth distribution of seeds in the soil and across treatments: on average 6% of seeds germinated after burial. In contrast to the seed position effect, we found no size effect on mobility and germinability of seeds after burial in the soil. Nevertheless, the fate of different-sized seeds may differ. While burial will remove large seeds from the regeneration pool, it may enhance seed bank build up in small-seeded species. Consequently, changes in the amount of litter cover and the invertebrate community play a significant role in plant community composition.     

Seed depredation negates the benefits of midstory hardwood removal on longleaf pine seedling establishment

Midstory hardwoods are traditionally removed to restore longleaf pine on fire‐excluded savannas. However, recent evidence demonstrating midstory hardwood facilitation on longleaf pine seedling survival has brought this practice into question on xeric sites. Also, midstory hardwoods could facilitate longleaf pine seedling establishment, as hardwood litter may conceal seeds from seed predators or improve micro‐environmental conditions for seedling establishment. However, little is known about these potential mechanisms. In this study, we tracked longleaf pine seed depredation and germination in artificially seeded plots (11 seeds/m²) in a factorial design fully crossing hardwood retention or removal with vertebrate seed predator access or exclusion in the Sandhills Ecoregion of North Carolina, U.S.A. Seed depredation averaged 78% across treatments and was greatest in unexcluded plots. Hardwood retention did not affect seed depredation. Longleaf pine averaged 3.6 germinants/4 m² across treatments, and was six times more abundant where vertebrates had been excluded. Hardwood removal had a strong positive effect on seedling germination, likely due to the removal of litter, but only when vertebrates were excluded. Our results indicated midstory hardwoods are not facilitating longleaf pine seedling establishment. Nevertheless, our results indicated that hardwood removal may not increase longleaf pine seedling establishment, as seed depredation diminished the effectiveness of hardwood removal under mast seed availability. Collectively, these results demonstrate the underlying complexity of the longleaf pine ecosystem, and suggest that planting may need to be part of the restoration strategy on sites where seed depredation limits longleaf pine natural regeneration.     

Empty Seeds Are Not Always Bad: Simultaneous Effect of Seed Emptiness and Masting on Animal Seed Predation

Seed masting and production of empty seeds have often been considered independently as different strategies to reduce seed predation by animals. Here, we integrate both phenomena within the whole assemblage of seed predators (both pre and post-dispersal) and in two contrasting microsites (open vs. sheltered) to improve our understanding of the factors controlling seed predation in a wind-dispersed tree (Ulmus laevis). In years with larger crop sizes more avian seed predators were attracted with an increase in the proportion of full seeds predated on the ground. However, for abundant crops, the presence of empty seeds decreased the proportion of full seeds predated. Empty seeds remained for a very long period in the tree, making location of full seeds more difficult for pre-dispersal predators and expanding the overall seed drop period at a very low cost (in dry biomass and allocation of C, N and P). Parthenocarpy (non-fertilized seeds) was the main cause of seed emptiness whereas seed abortion was produced in low quantity. These aborted seeds fell prematurely and, thus, could not work as deceptive seeds. A proportion of 50% empty seeds significantly reduced ground seed predation by 26%. However, a high rate of parthenocarpy (beyond 50% empty seeds) did not significantly reduce seed predation in comparison to 50% empty seeds. We also found a high variability and unpredictability in the production of empty seeds, both at tree and population level, making predator deception more effective. Open areas were especially important to facilitate seed survival since rodents (the main post-dispersal predators) consumed seeds mostly under shrub cover. In elm trees parthenocarpy is a common event that might work as an adaptive strategy to reduce seed predation. Masting per se did not apparently reduce the overall proportion of seeds predated in this wind-dispersed tree, but kept great numbers of seeds unconsumed.     

Lack of fire and insect herbivory constrain recruitment in a rare legume from Western Australia

We investigated factors influencing recruitment in the rare endemic shrub Acacia insolita subsp. recurva (Fabaceae) from the heavily cleared south‐west of Western Australia. We examined annual seed production over 4 years, including the impact of herbivory on reproductive output. Conversion of bud to fruit was low (overall 0.02%). The lack of significant difference in canopy dimensions between caged and uncaged plants suggested that vertebrate herbivore grazing was negligible, but invertebrate predation had a negative impact on seed production (loss of >80% of fruiting potential). Soil cores determined the presence of soil‐stored seeds and an experimental burn confirmed that plants are fire‐killed and can regenerate from the seed bank. This seed reserve was found to contain <5 seeds m^?2, and both freshly collected seeds and seeds retrieved from the soil had high viability (99% vs. 91%) when subjected to a germination test. Seedling recruitment 29 months post‐fire resulted in a ratio of three seedlings for every adult killed by fire. We also compared reproductive success in this rare Acacia with its common conspecific and although the rare species produced more flowers, the success of flowering did not translate into better fruit set. We conclude that insect damage to reproductive branchlets and lack of appropriate disturbance are major factors constraining recruitment. Active site management may be required for the continued persistence of this fire‐dependent legume.     

Does predation contribute to tree diversity?

Seed and seedling predation may differentially affect competitively superior tree species to increase the relative recruitment success of poor competitors and contribute to the coexistence of tree species. We examined the effect of seed and seedling predation on the seedling recruitment of three tree species, Acer rubrum (red maple), Liriodendron tulipifera (yellow poplar), and Quercus rubra (northern red oak), over three years by manipulating seed and seedling exposure to predators under contrasting microsite conditions of shrub cover, leaf litter, and overstory canopy. Species rankings of seedling emergence were constant across microsites, regardless of exposure to seed predators, but varied across years. A. rubrum had the highest emergence probabilities across microsites in 1997, but Q. rubra had the highest emergence probabilities in 1999. Predators decreased seedling survival uniformly across species, but did not affect relative growth rates (RGRs). Q. rubra had the highest seedling survivorship across microsites, while L. tulipifera had the highest RGRs. Our results suggest that annual variability in recruitment success contributes more to seedling diversity than differential predation across microsites. We synthesized our results from separate seedling emergence and survival experiments to project seedling bank composition. With equal fecundity assumed across species, Q. rubra dominated the seedling bank, capturing 90% of the regeneration sites on average, followed by A. rubrum (8% of sites) and L. tulipifera (2% of sites). When seed abundance was weighted by species-specific fecundity, seedling bank composition was more diverse; L. tulipifera captured 62% of the regeneration sites, followed by A. rubrum (21% of sites) and Q. rubra (17% of sites). Tradeoffs between seedling performance and fecundity may promote the diversity of seedling regeneration by increasing the probability of inferior competitors capturing regeneration sites.     

动物与植物种子更新的关系Ⅱ．动物对种子的捕食、扩散、贮藏及与幼苗建成的关系

植物的繁殖体总是面临来自各类生物（如昆虫、脊椎动物、真菌）的捕食风险。因动物捕食引起的种子死亡率影响植物的适合度、种群动态、群落结构和物种多样性的保持。种子被捕食的时间和强度成为植物生活史中发芽速度、地下种子库等特征的主要选择压力,而种子大小、生境类型等因素也影响动物对植物种子的捕食。捕食者饱和现象被认为是植物和种子捕食者之间的高度协同进化作用的结果,是限制动物破坏种子、提高被扩散种子存活率的一种选择压力。大部分群落中的大多数植物种子被动物扩散。种子扩散影响种子密度、种子被捕食率、病原体攻击率、种子与母树的距离、种子到达的生境类型以及建成的植株将与何种植物竞争,从而影响种子和幼苗的存活,最终影响母树及后代植物的适合度。种子被动物扩散后的分布一般遵循负指数分布曲线,大多数种子并没有扩散到离母树很远的地方。捕食风险、生境类型、植被盖度均影响动物对种子的扩散。植物结实的季节和果实损耗的过程也体现了其对扩散机会的适应。许多动物有贮藏植物种子的行为。动物贮藏植物繁殖体的行为,一方面调节食物的时空分布,提高了贮食动物在食物缺乏期的生存概率;另一方面也为种子萌发提供了适宜条件,促进了植物的扩散。于是,植物与贮食动物形成了一种协同进化关系,这种关系可能是自然界互惠关系（mutualism)的一种。影响幼苗存活和建成的因子包括种子贮蒇点的微生境、湿度、坡向、坡度、林冠盖度等。许多果食性动物吃掉果肉后,再将完好的种子反刍或排泄出来。种子经动物消化道处理后,发芽率常有所提高。     

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.     

Effects of seed burial and fire on seedling and sapling recruitment, survival and growth of African savanna woody plant species

Seed predation and seedling mortality can act as strong demographic “bottlenecks” to sapling recruitment in African savanna woodlands. Fire also limits tree recruitment from saplings by suppressing their growth. I conducted field experiments with 13 woody plant species to assess the effects of seed burial on seedling emergence rates and effects of fire on seedling and sapling survival and growth rates over a period of 8 years at a savanna plot in central Zambia, southern Africa. Seed removal rates by small rodents varied among years and buried seeds had significantly higher emergence rates than seeds exposed to predators in most but not all the species. Annual burning reduced sapling growth in some species but in other species saplings experienced successive shoot die back even in the absence of fire. The findings show that for some woody species, seed predation is an important constraint to seedling recruitment but not for others and annual fires are important hindrances to demography and growth for some species but not others. Thus, demographic “bottlenecks” occur at different life history stages in different savanna woody species and these have the potential to alter woody tree competitive relationships and ultimately savanna structure.     

Experimental Assessment of Factors Limiting Seedling Recruitment of an Amazonian Understory Herb

Understory herbs are an important and species-rich component of tropical forests, but little is known about factors limiting recruitment of these herbs. In a Peruvian flood plain forest, we studied seedling recruitment of the widespread clonal herb Heliconia metallica . We analyzed natural recruitment and experimentally added seeds, excluded predators and removed the understory vegetation at occupied and unoccupied sites to test whether seed limitation, predation, disturbance, and the presence of mature conspecific plants influence seedling recruitment. The number of naturally recruited seedlings was higher at flooded sites and close to flowering H. metallica ramets. Predation caused considerable seed loss shortly after sowing. Seed survival strongly increased with the openness of a site, but was not affected by the presence of mature conspecifics. Seed addition increased the number of seedlings at both occupied and unoccupied sites. Two years after sowing, the number of established seedlings and their size increased with light availability at a site. In experimentally disturbed plots, light availability, seedling establishment, and survival were higher. The effect of disturbance on seedling survival was stronger at occupied sites, probably due to strong light competition from mature H. metallica plants. Herbivore damage on leaves of seedlings was lower at occupied sites, but seedlings grew faster at unoccupied sites. We conclude that seedling recruitment of H. metallica depends on natural disturbance and is limited by both seed availability and the presence of mature conspecific plants.     

Variability in post-dispersal seed predation in deciduous woodland: relative importance of location,seed species,burial and density

The considerable variability found in post-dispersal seed predation and the absence of consistent directional trends (e.g., with reference to seed size) has made it difficult to predict accurately the impact of seed predators on plant communities. We examined the variation attributable to location, seed density and seed burial on the removal of seeds of three tree species: Fraxinus excelsior, Taxus baccata and Ulmus glabra. Experiments were undertaken in five deciduous woodlands in Durham, U.K., and the relative importance of vertebrate and invertebrate seed predators was assessed using selective exclosures. In all five woodlands, seed removal was greatest from treatments to which vertebrates had access, and losses attributable to invertebrates were negligible. Rodents, in particular Apodemus sylvaticus (Muridae) and Clethrionomys glareolus (Cricetidae), were the principal seed consumers in these woodlands. Unidentified vertebrate seed predators (probably birds, rabbits and/or squirrels) appeared to be significant seed removers in three of the five woodlands. Rates of removal differed among the three tree species, increasing in the following order Fraxinus < Taxus < Ulmus but were not related to seed mass. The major effect influencing rates of seed removal was seed burial, which halved rates of seed removal overall. The effect of seed burial was a function of seed size. The larger seeds of Taxus realising little benefit from seed burial whereas encounter of the smaller Ulmus seeds fell by almost two-thirds. Removal was density-dependent for all three species. However, the relative increase in seed encounter through an increase in seed density was a negative function of seed size. This suggests that, for large seeds, the opportunity to escape seed predation via burial or reduced seed density is limited. These results reveal a number of parallels with other studies of post-dispersal predation and identify several generalities regarding the interaction between plants and post-dispersal seed predators. Comparison of the seed predation results with actual seedling distributions suggests that seed predators may influence regeneration of Ulmus glabra but probably play a lesser role in the dynamics of Taxus baccata and Fraxinus excelsior.