Seed predation limits post-fire recruitment in the waratah (<Emphasis Type="Italic">Telopea speciosissima</Emphasis>)

Seed predation may reduce recruitment in populations that are limited by the availability of seeds rather than microsites. Fires increase the availability of both seeds and microsites, but in plants that lack a soil- or canopy-stored seed bank, post-fire recruitment is often delayed compared to the majority of species. Pyrogenic flowering species, such as Telopea speciosissima, release their non-dormant seeds more than 1 year after fire, by which time seed predation and the availability of microsites may differ from that experienced by plants recruiting soon after fire. I assessed the role of post-dispersal seed predation in limiting seedling establishment after fire in T. speciosissima, in southeastern Australia. Using a seed-planting experiment, I manipulated vertebrate access to seeds and the combined cover of litter and vegetation within experimental microsites in the 2 years of natural seed fall after a fire. Losses to vertebrate and invertebrate seed predators were rapid and substantial, with 50% of seeds consumed after 2 months in exposed locations and after 5 months when vertebrates were excluded. After 7 months, only 6% of seeds or seedlings survived, even where vertebrates were excluded. Removing litter and vegetation increased the likelihood of seed predation by vertebrates, but had little influence on losses due to invertebrates. Microsites with high-density vegetation and litter cover were more likely to have seed survival or germination than microsites with low-density cover. Recruitment in pyrogenic flowering species may depend upon the release of seeds into locations where dense cover may allow them to escape from vertebrate predators. Even here, conditions suitable for germination must occur soon after seed release for seeds to escape from invertebrate predators. Seed production will also affect recruitment after any one fire, while the ability of some juvenile and most adult plants to resprout after fire buffers populations against rapid declines when there is little successful recruitment.     

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.     

Spatial dynamics of <Emphasis Type="Italic">Ilex aquifolium</Emphasis> populations seed dispersal and seed bank: understanding the first steps of regeneration

The objective of this study was to analyse quantitatively the spatial distribution of holly (Ilex aquifolium L.) seed rain and seed bank, and to detect the relationships between these consecutive processes. We measured seed dispersal by birds and fallen fruits, and also density and viability of seed bank in two Ilex populations in central Spain. Analysis was made distinguishing the following microhabitats: holly woodland, edge of holly woodland, open grassland 10 m and 100 m from the woodland, fleshy fruit shrubs, dry fruit shrubs, and adjacent non-holly woodland. Spatial distribution of dispersed and in-soil seeds was measured by the clumping index. Seed rain and seed bank under holly woodlands were significantly higher than in the other microhabitats. Forest edges and fleshy fruit shrubs were the next microhabitats with the highest seed rain and seed bank density. Interannual and interlocality variations were not significant. The relative importance of the different dispersal methods varied between microhabitats, with a similar support of bird dispersed seeds and fallen fruits within the woodland and a greater influence of cattle dispersal in open areas. Seed spatial aggregation was significant in both dispersed seeds and soil seeds from holly woodlands and the edge of the forest. Aggregation under shrubs, grasslands and the adjacent forests evidenced a general random distribution of holly seeds (only in some cases clumping index was significant). Quantitative differences between seed rain and seed bank are important. Post-dispersal seed predation did not modify seed rain distribution, which was mirrored in the seed bank pattern. These two phases of holly regeneration had a heavy spatial influence, determined by the landscape structure and activity of the dispersal agents, that reflects a differential recruitment potential. Comparisons between both populations suggest that in the southern locality (Robregordo) holly has a weaker capacity to colonize open areas, and a stronger recruitment limitation due to propagule availability.     

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     

Seed dispersal by lizards on a continental‐shelf island: predicting interspecific variation in seed rain based on plant distribution and lizard movement patterns

Aim We estimated the patterns of seed deposition provided by the eyed lizard, Timon lepidus, and evaluated whether these patterns can be generalized across plant species with different traits (fruit and seed size) and spatial distributions. Location Monteagudo Island, Atlantic Islands National Park (north‐western Spain). Methods We radio‐tracked seven lizards for 14 days and estimated their home ranges using fixed kernels. We also geo‐referenced all fruit‐bearing individuals of four plant species dispersed by eyed lizards in the study area (Corema album, Osyris alba, Rubus ulmifolius and Tamus communis), measured the passage time of their seeds through the lizard gut, and estimated seed predation in four habitats (bare sand, grassland, shrub and gorse). Seed dispersal kernels were estimated using a combination of these data and were combined with seed predation probability maps to incorporate post‐dispersal seed fate (‘seed survival kernels’). Results Median seed gut‐passage times were around 52–98 h, with maximum values up to 250 h. Lizards achieved maximum displacement in their home ranges within 24–48 h. Seed predation was high (80–100% of seeds in 2 months), particularly under Corema shrub and gorse. Seed dispersal kernels showed a common pattern, with two areas of preferential seed deposition, but the importance of these varied among plant species. Interspecific differences among dispersal kernels were strongly reduced by post‐dispersal seed predation; hence, seed survival kernels of the different plant species showed high auto‐ and pairwise‐correlations at small distances (< 50 m). As a result, survival to post‐dispersal seed predation increased with dispersal distance for O. alba and T. communis, but not for C. album. Main conclusions Seed dispersal by lizards was determined primarily by the interaction between the dispersers’ home ranges and the position of the fruit‐bearing plants. As a result, seed rain shared a common template, but showed considerable variation among species, determined by their specific spatial context. Seed predation increased the spatial coherence of the seed rain of the different species, but also resulted in contrasting relationships between seed survival and dispersal distance, which may be of importance for the demographic and evolutionary processes of the plants.     

Seed and microsite limitation of recruitment in plant populations

Summary Availability of seed and microsites, respectively, are two factors that potentially may limit recruitment in plant populations. Microsites are small-scale sites suitable for germination and survival of seedlings. We discuss this dichotomy of recruitment limitation both from a theoretical and empirical point of view. Investigations of recruitment in 14 woodland species showed that 3 species were seed limited, 6 species were limited by a combination of seed and microsite availability, and 5 species were found not to be seed limited, but the limiting factor was not identified. A combination of seed and microsite limitation implies that recruitment is promoted by increasing both seed and microsite availability. We suggest that the importance of seed limitation in plant populations has been underestimated, and that the operating limiting factors may be dependent on spatial and temporal scale. We expect that many species, if adequately studied, will turn out to be both seed and microsite limited. Experimental field studies that incorporate a range of seed and microsite densities in various spatial and temporal scales are needed to examine the extent to which plant populations are seed and microsite limited.     

Composition and clumping of seeds deposited by frugivorous birds varies between forest microsites

More frequent deposition of seeds by frugivores beneath plants in fruit could impose spatial limits to the distribution of plants dispersed by animals and contribute to species coexistence. Also, differences in diet and use of microhabitats by seed dispersers could promote spatial variation in the combination of seed species deposited. We investigated patterns of seed deposition of Miconia fosteri and Miconia serrulata (Melastomataceae) by birds in the Amazon. The goal was to determine how distribution and abundance of fruiting plants, both con‐ and hetero‐specifics, affect the spatial variability in clumping and composition of multi‐specific seed deposition. We established two 9‐ha plots in undisturbed terra‐firme understory in the Ecuadorian Amazon. Seed rain was sampled with seed traps located in four microsites: below plants of the focal species, below Anthurium eminens (Araceae), and in randomly selected microsites. We examined seed deposition in these microsites in relation to habitat, fruiting neighborhood (fruit abundance, and distance to and density of plants of the target species), and crop size of M. fosteri or M. serrulata to determine if microsites differed in abundance and species composition of seeds. Seed traps below plants in fruit received more seeds than did randomly located traps. Seeds of the target species were, moreover, more commonly deposited below con‐ rather than hetero‐specific plants. Seed aggregation below fruiting plants increased in forest neighborhoods where the abundance of fruits and the combination of fruiting plant species contributed to the arrival of seeds. Microsites differed notably in the combination of seeds deposited by frugivores, and differences were less pronounced among microsites that received seeds of M. fosteri and M. serrulata than among all microsites where at least some seed species were deposited by birds. We demonstrate that two closely related, ecologically similar species possess many similarities in their patterns of seed deposition and in the factors that affect those patterns. The combination of seed species deposited below foci of dispersal depended on the fruiting plant species, and the spatial patterns of seed deposition varied with the location of the microsite and the combination of co‐dispersed species in the neighborhood. Similar species that share the same dispersers were confronted with different combinations of seeds depending on the microsite where they arrived, which could promote forest heterogeneity in the combination of plant species.     

Importance of Seed and Microsite Limitation: Native Wildflower Establishment in Non‐native Pasture

Barriers to establishing native plant communities on former pasture include dominance by a single planted species, hydrologic and edaphic alteration, and native species propagule limitation. Establishment may be dispersal‐limited (propagules do not arrive at the site), microsite‐limited (areas suitable for seedling emergence and survival do not exist), or both. Successful restoration strategies hinge on identifying and addressing critical limitations. We examined seed and microsite limitation to establishment of a native wildflower (Coreopsis lanceolata ) in a former pasture dominated by Paspalum notatum (bahiagrass). We determined the relative and interactive effects of microsite (irrigation and disturbance) and seed limitation on C. lanceolata establishment. We tested (1) irrigation (none, pre‐seeding, and pre‐ and post‐seeding), (2) disturbance (none, sethoxydim, glyphosate, and topsoil removal), and (3) C. lanceolata seeding rate (three seeding densities). Applying glyphosate before seeding increased C. lanceolata establishment compared to other disturbance treatments. Ultimately, C. lanceolata establishment was not affected by irrigation. Coreopsis lanceolata establishment was limited when seeded at 100 live seeds/m² but not at 600 or 1100 live seeds/m². Seed and microsite availability interactively affected C. lanceolata establishment, in that microsite limitation was biologically relevant only when a minimum number of seeds were present. In practice, both seed and microsite requirements must be met for successful establishment, and increasing the availability of seeds or microsites does not compensate for limitations of the other. Here, it is the relative importance of seed and microsite limitations that drives plant establishment; these limitations do not represent a simple dichotomy.     

Linking seed dispersal,germination and seedling recruitment in the invasive species <Emphasis Type="Italic">Berberis darwinii</Emphasis> (Darwin’s barberry)

Seedling recruitment is a multi-phased process involving seed production, dispersal, germination, seedling establishment and subsequent survival. Understanding the factors that determine success at each stage of this process is of particular interest to scientists and managers seeking to understand how invasive species spread and persist, and identify critical stages for management. To understand the factors and processes influencing recruitment of the invasive species Berberis darwinii Hook. (Darwin’s barberry), temporal and spatial patterns of seed dispersal, germination and seedling establishment were examined. Seed dispersal from a large source population was measured over two fruiting seasons, and subsequent patterns of seedling emergence and survival within each cohort were measured. Seed longevity was tested under both natural and artificial conditions. Seeds were widely dispersed by birds, up to 450 m from the source population. Dispersal was essential to seedling establishment, as few seedlings survived beneath the parent canopy. Seeds were relatively short-lived in the soil under both field and glasshouse conditions, with few surviving for more than 1 year. Patterns of newly emerged seedlings largely reflected patterns of seed rain, but seedling survival was significantly affected by distance from source population, seedling density and light environment. These results suggest that recruitment of B. darwinii is dependent on dispersal of seeds to favourable microsites. Management priorities should include the removal of fruiting plants, and seedling control in highlight areas.     

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.     

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.     

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.     

Spatial Segregation of the Sexes and Nutrients affect Reproductive Success in a Dioecious Wind-pollinated Grass

In many dioecious plant species in which spatial distributions of males and females have been examined, the sexes are spatially segregated – usually along an environmental gradient. Unless pollen is uniformly distributed in a population, spatial segregation of the sexes should reduce the average mating success of individuals. In three Californian populations of Distichlis spicata – a wind-pollinated grass species that exhibits spatial segregation of the sexes – I examined patterns of pollen movement and the effects of pollen load and nutrient availability on seed set to determine whether spatial segregation of the sexes actually reduces mating success for both males and females. In two of the populations, pollen dispersal was restricted, and pollen augmentation consistently, significantly increased seed set. However, in the third population – which had the lowest seed set – I found that although there were some indications of pollen limitation, pollen dispersal was not restricted, and seed production was limited primarily by nutrient availability. These results imply that in some populations of D. spicata nutrient limitation on the production of seeds by females may be sufficiently strong that spatial segregation of the sexes causes a fairly low cost to reproductive success compared with a more random distribution of the sexes. However, in other populations, pollen does limit mating success, and the spatial segregation of males and females in these populations is reducing the fecundity of both males and females.     

Could poor seed dispersal contribute to predation by introduced rodents in a Hawaiian dry forest?

Dry forests are among the most diverse, yet threatened, communities in Hawai’i. Dry forests throughout the archipelago suffer from a lack of natural regeneration of trees. Two factors that may limit tree recruitment include poor seed dispersal and seed predation by rodents. Poor or limited dispersal of fleshy-fruited species results in seeds and fruits falling directly under parents. Dispersed and non-dispersed seeds may differ in their vulnerability to predation. We tested effects of seed location (under/away from parent trees) and pulp (presence/absence) on predation of four native species that suffer from limited dispersal and one readily-dispersed alien species in Kanaio Natural Area Reserve, Maui. Three natives (Diospyros sandwicensis, Pleomele auwahiensis, Santalum ellipticum), had significantly more seeds removed under parent trees than in exposed sites away from trees. For the one alien (Bocconia frutescens) and two native trees (D. sandwicensis, P. auwahiensis) that were evaluated, significantly more intact fruits were removed than were cleaned seeds. Presence of teeth marks and gnawed seed husk fragments indicate introduced rodents are destroying many of the seeds they remove. These results suggest that seed predation is disproportionately concentrated among poorly-dispersed seeds and may contribute to recruitment failure.     

Post-dispersal seed predation and the establishment of vertebrate dispersed plants in Mediterranean scrublands

The post-dispersal fate of seeds and fruit (diaspores) of three vertebrate-dispersed trees, Crataegus monogyna, Prunus mahaleb and Taxus baccata, was studied in the Andalusian highlands, south-eastern Spain. Exclosures were used to quantify separately the impact of vertebrates and invertebrates on seed removal in relation to diaspore density and microhabitat. The three plant species showed marked differences in the percentage of diaspores removed, ranging from only 5% for C. monogyna to 87% for T. baccata. Although chaffinches (Fringilla coelebs) fed on diaspores, rodents (Apodemus sylvaticus) were the main vertebrate removers of seed and fruit. Two species of ant (Cataglyphis velox and Aphaenogaster iberica) were the only invertebrates observed to remove diaspores. However, the impact of ants was strongly seasonal and they only removed P. mahaleb fruit to any significant extent. While removal of seed by rodents was equivalent to predation, ants were responsible for secondary dispersal. However, their role was limited to infrequent, small-scale redistribution of fruit in the vicinity of parent trees. Rodents and ants differed in their use of different microhabitats. Rodents foraged mostly beneath trees and low shrubs and avoided open areas while the reverse was true of ants. Thus, patterns of post-dispersal seed removal will be contigent on the relative abundance and distribution of ants and rodents. Studies which neglect to quantify separately the impacts of these two guilds of seed removers may fail to elucidate the mechanisms underlying patterns of post-dispersal seed removal. The coincidence of both increased seed deposition by the main avian dispersers (Turdus spp.) and increased seed predation with increasing vegetation height suggested that selection pressures other than post-dispersal seed predation shape the spatial pattern of seed dispersal. Rather than providing a means of escaping post-dispersal seed predators, dispersal appears to direct seeds to microhabitats most suitable for seedling survival. Nevertheless, the reliance of most vertebrate-dispersed trees on regeneration by seed and the absence of persistent soil seed banks imply that post-dispersal seed predators may exert a strong influence on the demography of the plants whose seeds they consume. Even where microsites are limited, the coincidence of the most suitable microhabitats for seedling establishment with those where seed predation is highest provide a means by which selective seed predators can influence community composition. Received: 19 August 1996 / Accepted: 25 January 1997     

A Comparison of Frugivory and Scatter-Hoarding Seed-Dispersal Syndromes

Frugivory (animals ingest all or part of fleshy-fruits and defecate or regurgitate seeds) and scatter-hoarding (animals store seeds in small caches in soil) are two important vertebrate-mediated seed dispersal syndromes. In both instances, there is an exchange of food for seed transport, but there are many important differences. For example, the seeds of frugivore-dispersed plants are often clumped in feces or under roosts and the microsite of depostion is often unsuitable for seedling establishment. However, frugivores can transport seeds long distances allowing plants to colonize new areas and promoting gene flow between populations. Scatter-hoarding animals, on the other hand, actively space seeds in microsites that often favor seedling establishment, but don’t move seeds very far. Both modes of seed dispersal can be very effective, but in different way. Some species of plants have evolved propagules that take advantage of both modes of dispersal in two separate phases (i.e., diplochory), apparently to obtain both types of benefits.     

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.     

Microsite variation in light availability and seedling growth ofQuercus serrata in a temperate pine forest

Spatial heterogeneity in light availability for tree seedlings under the canopy of a temperate pine forest was studied. Six-day measurements at 10-s intervals revealed a great variety in the temporal patterns of photosynthetic photon flux density (PPFD) and histograms among observation days and microsites; mean daily total PPFD relative to full sun varied from 1.5% to 10.4% depending on the microsites. The occurrence and duration of PPFD above 80 μmol m⁻² s⁻¹, which might reflect sunfleck activity, varied greatly among the microsites. However, several simple empirical relationships were found between some parameters characterizing microsite light availability and sunfleck activity; the diffuse site factor was correlated well with other parameters, including daily total PPFD, daily totals and daily summed durations of high PPFD above any examined threshold level, and its contribution to daily total PPFD. Diffuse site factors which were obtained for 700 microsites within an area of 28 m² on three different occasions during the growing season showed high correlations within the microsite. Based on the regressed relationship between the relative growth rate of current-year seedlings ofQuercus serrata and the microsite diffuse site factor and the results of area-survey measurement of the diffuse site factor, an estimation was made of the abundance of potential ‘safe-sites’ for seedling growth of the species; the ‘safe sites’ were estimated to cover 40% and 0% of the total area of the sunny and shady sites of the forest, respectively.     

Spatial and seasonal patterns of microsite light availability in a remnant fragment of deciduous riparian forest and their implication in the conservation ofArisaema heterophyllum, a threatened plant species

Seasonal and spatial patterns of light availability were investigated in the understory of a small fragment (approximately 1300 m²) of a riparian deciduous forest of the Kokai River in central Japan dominated byQuercus acutissima Carruth., with the aim to understand the characteristics of microsite light availability forArisaema heterophyllum Blume, a threatened plant species uniquely associated with the riparian habitat. Diffuse site factor, which is the ratio of PFD (photon flux density, 400–700 nm) at a microsite to the open sky reference under diffuse light condition, was shown to be a satisfactory index for the evaluation of light availability for the understory plants in the habitat. Diffuse site factor 1,0 1,000 understory microsites along a 20 m transect from the edge to the interior of the forest, showed conspicuous seasonal changes in both mean and variation. Light availability decreased with seasonal tree canopy regeneration, with the highest spatial heterogeneity being recorded during the time of canopy closure. Auto-correlations of microsite light availability between different seasons were considerably high, suggesting the stability of relative light availability for individual microsites throughout the growing season ofA. heterophyllum. Fairly high light availability during summer season, which surpassed 20% of the open sky reference in most microsites, would be important for the growth and persistence of summer herbaceous plants likeA. heterophyllum.     

A spatially explicit analysis of seedling recruitment in the terrestrial orchid Orchis purpurea

Seed dispersal and the subsequent recruitment of new individuals into a population are important processes affecting the population dynamics, genetic diversity and spatial genetic structure of plant populations. Spatial patterns of seedling recruitment were investigated in two populations of the terrestrial orchid Orchis purpurea using both univariate and bivariate point pattern analysis, parentage analysis and seed germination experiments. Both adults and recruits showed a clustered spatial distribution with cluster radii of c. 4-5 m. The parentage analysis resulted in offspring-dispersal distances that were slightly larger than distances obtained from the point pattern analyses. The suitability of microsites for germination differed among sites, with strong constraints in one site and almost no constraints in the other. These results provide a clear and coherent picture of recruitment patterns in a tuberous, perennial orchid. Seed dispersal is limited to a few metres from the mother plant, whereas the availability of suitable germination conditions may vary strongly from one site to the next. Because of a time lag of 3-4 yr between seed dispersal and actual recruitment, and irregular flowering and fruiting patterns of adult plants, interpretation of recruitment patterns using point patterns analyses ideally should take into account the demographic properties of orchid populations.