Seed Dispersal of a High Quality Fruit by Specialized Frugivores: High Quality Dispersal?1

Dispersal quality, as estimated by the cumulative effects of dispersal, germination, seed predation, and seedling survival, was examined for Beilschmiedia pendula (Lauraceae) in Monteverde, Costa Rica. I determined the pattern of dispersal by finding seeds deposited by birds, protected the seeds from seed predators with cages to assess germination and seedling survival, and examined seed predation rates with marked seeds. Seed predation, germination, and seedling survival were compared between seeds naturally dispersed by birds and seeds placed at randomly located sites. Approximately 70 percent of seeds dispersed by birds (N= 244) were deposited <10 m from crown edges of fruiting B. pendula trees, although some seeds were dispersed at least 70 m away. Larger seeds were more likely to be dispersed under or close to the parent trees, and larger seeds produced larger seedlings. Seed size was not correlated directly with seedling survival, but larger seedlings at three months were most likely to survive one year. Seed predation by mammals and insects and seedling mortality due to fungal pathogens were concentrated beneath the crowns of parent trees. Seedlings and saplings were more abundant beneath fruiting B. pendula trees, but individuals farther away were taller on average. Thus, dispersal is beneficial for B. pendula, but such benefits appear most pronounced at a small spatial scale; seeds dispersed >30 m from the crown edges actually had a lower probability of survival than those dispersed 10–20 m. Only 10 percent of B. pendula. seeds received high‐quality dispersal in terms of landing in the zone with the highest per seed probability of seedling survival 10–20 m from parental crowns.     

Factors Influencing Seedling Establishment Rates in Zostera marina and Their Implications for Seagrass Restoration

Selection of strategies to efficiently utilize limited seed supplies in efforts to restore the seagrass Zostera marina (eelgrass) requires a better understanding of the processes that limit seedling establishment at potential restoration sites. We investigated the effect of seed distribution timing on seedling establishment and tested for interactive effects of seed burial and distribution timing. We also investigated the effect of habitat type on seedling establishment by distributing Z. marina seeds inside and outside of established Ruppia maritima (widgeongrass) patches and examined mechanisms causing habitat differences by manipulating seed position (buried or unburied) and vulnerability to seed predators (unprotected or protected in packets). Seeds distributed on the sediment surface in the summer (July or August) produced fewer seedlings than seeds distributed in fall (October) in five of six trials over 3 years. Seed burial increased success rates for seeds distributed in summer at one of two sites tested, eliminating the effect of season, but reduced success at the other site. Seeds placed in R. maritima generally produced fewer seedlings than seeds in bare sand, and although seed burial and protection in packets increased success in bare sand at three of four sites, the effect was less consistent in R. maritima. We conclude that seed predation and physical interactions were influential in reducing seedling establishment in R. maritima, contrary to hypotheses positing a nursery role for existing vegetation. Efficient restoration efforts with Z. marina seeds should target unvegetated areas after summertime sources of mortality have diminished. Direct seed burial may enhance seedling establishment rates.     

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.     

Differential seed and seedling predation by crabs: impacts on tropical coastal forest composition

Recently, the importance of seed predation by crabs on mangrove species distributions and densities has been established by several studies. In a tropical coastal terrestrial forest in Costa Rica, we investigated the relative importance of predation by land crabs, Gecarcinus quadratus, and hermit crabs, Coenobita compressus, on measured forest composition through a series of seed removal and seedling establishment experiments. We also used natural light-gaps and adjacent non-gap sites to test how canopy cover affects crab predation (seed removal) and seedling establishment. We found fewer tree species (S=18) and lower densities (seedlings, saplings, and adults) in the coastal zone within 100 m of coastline, than in the inland zone (S=59). Land crab densities were higher in the coastal zone (3.03±1.44 crabs m^–2) than in the inland zone (0.76±0.78 crabs m^–2), and hermit crabs were not present in the inland zone. Seed removal and seedling mortality also were higher in the coastal zone than in the inland zone, and in the open controls than in the crab exclosures. Mortality of seeds and seedlings was two to six times higher in the controls than exclosures for four of the five experiments. Crabs preferred seeds and younger seedlings over older seedlings but showed no species preferences in the seed (Anacardium excelsum, Enterolobium cyclocarpum, and Terminalia oblonga) and seedling (Pachira quinata and E. cyclocarpum) stages. We conclude that the observed differences in tree densities were caused by differential crab predation pressure along the coastal gradient, while the differences in species composition were due to predator escape (satiation) by seed quantity. Canopy cover did not affect seed removal rates, but did affect seedling survival with higher mortality in the non-gap versus gap environments. In summary, crab predation of seeds and seedlings, and secondarily canopy cover, are important factors affecting tree establishment in terrestrial coastal forests.     

Feathermoss seedbeds facilitate black spruce seedling recruitment in the forest–tundra ecotone (Labrador,Canada)

As climate warms, conifers are expected to expand their ranges into alpine tundra where ecological factors such as seedbed availability, and post‐dispersal seed and seedling predation may control local recruitment. Seedbed composition may influence microhabitat, nutrients, physical structure, and predation level and, therefore, affect the success of conifer recruitment, thereby providing the template for future expansion. In the boreal forest, seedbed–seedling competition dominates such that seedbed removal increases black spruce recruitment. In the harsher climate of the Mealy Mountains boreal forest–tundra ecotone (Labrador, Canada) the Stress gradient hypothesis (SGH) predicts that facilitation may dominate seedbed–seedling interactions. This study investigated potential mechanisms of seedbed facilitation (temperature, water, nutrients, physical protection) in three conifer seedbeds (Pleurozium schreberi, Cladonia spp., bare soil) and examined whether seed predation and/or seedling herbivory varied among seedbeds over three years. Seed emergence was low overall (< 10% on all treatments), but highest on Pleurozium (6.3%), followed by bare ground (4.6%) and Cladonia (0.3%). Facilitation was observed between Pleurozium and black spruce as seedling height increase (31%) and survival (55%) were highest; herbivory, seed predation and overwinter mortality were lowest compared to both Cladonia and bare ground seedbeds. Unlike in the closed canopy boreal forest, seedlings recruited poorly on bare soil as seedling height increase and survival were 20.5% and 26%. Temperature and water availability were similar across seedbeds, while nutrient availability was higher on Pleurozium. The physical structure of Pleurozium likely protects first to third‐year seedlings from temperature extremes and predators. As climate warms and seed availability increases, Pleurozium may facilitate black spruce recruitment and treeline expansion.     

Effects of Dispersal Pattern and Mammalian Herbivores on Seedling Recruitment for Virola michelii (Myristicaceae) in French Guiana1

The effects of dispersal pattern (seeds in small clumps vs. seeds scattered in pairs) and distance to the nearest Carapa procera (Meliaceae; a tree that produces seeds preferred by terrestrial vertebrates) on survival of seeds and seedlings were examined for the animal–dispersed tree species Virola michelii (Myristicaceae) in a mature forest‘at Paracou, French Guiana, in 1992 and 1993. We assessed the putative role of ground–dwelling mammalian herbivores, rodents, and ungulates that filter the seed shadow, acting either as dispersers or predators and thus modifying the original pattern of seed dispersal made by frugivores. We measured the effects of simulated seed burial by rodents using marked seeds and quantified the effect of protecting seeds and seedlings from ground–dwelling vertebrates on seedling germination and survival with fence exclosures in 1992. Dispersal pattern had short–term but no long–term effects on the proportion of V. michelii seeds that survived one year later as seedlings. In the short term, within six weeks, clumped seeds survived better than scattered seeds in both years. Marked seeds that were removed from their site of dispersal were eaten; rodents only rarely buried seeds of V. michelii, and seed burial reduced seed and seedling survivorship. The combined effect of the factors year and Carapa proximity significantly affected seed survival within six weeks. Although six–week seed survival was greater in 1993 than in 1992, seedling establishment was lower in 1993 than in 1992 following a lower rainfall regime during the key period of seed germination (February). One–year seed and seedling survivorship was similar between treatments and years. Seed survival and seedling establishment in V. michelii was dependent on vertebrates in the short term and on climate in the long term. Overall, seed and seedling survivorship depended on a combination of these factors.     

Advantage of early seedling emergence in <Emphasis Type="Italic">Fagus crenata</Emphasis>: importance of cotyledon stage for predator escape and pathogen avoidance

To examine how early-emerging seedlings take advantage of establishment in a deciduous forest, we explored the relationships among the emergence date, growth stage and major causes of mortality (damping-off by fungi and predation by rodents) in seedlings of Fagus crenata. The emergence of current-year seedlings and their survival and damage were followed at short (3–7 days in early spring) intervals for a growing season. The growth stage was divided into two stages, with only cotyledons (CT stage) and with true leaves (TL stage). The survival rate was negatively correlated with the emergence date, indicating the advantage of early seedling emergence. This advantage was largely explained by the lower occurrence of damping-off. In contrast, seedling predation occurred independently of the emergence date, but depended strongly on the developmental stage. Rodents consumed a considerable number of seeds during the early period after emergence, and strongly preferred CT-stage over TL-stage seedlings throughout the growing season. Therefore, seedling predation was inferred to be concentrated in a relatively short period while the remaining seeds were depleted and CT-stage seedlings were abundant. The seedling stage synchronously shifted from the CT to TL stage. This growth-stage transition was independent of the emergence date and appeared to correspond with the timing of seed depletion. Delayed stage transition resulted in a disproportionately high risk of damping-off later in the season. Our results indicate that early seedling emergence is advantageous for F. crenata for resistance to pathogens and that the timing of growth-stage progression of seedlings appears important to escape predation because of the distinct food preference of predators.     

THE EFFECT OF SEED DIMORPHISM ON THE GERMINATION AND SURVIVAL OF SALICORNIA EUROPAEA L. POPULATIONS

Seed dimorphism influenced the germination behavior of Salicornia europaea L., with small seeds being more dormant and less salt tolerant than large seeds. All of the large less dormant seeds of S. europaea germinated prior to May, and all seedlings produced after this time were from small seeds. A persistent seed bank was maintained by the small dormant seeds. Survivorship was relatively constant during the normal germination season, but increasing salinity stress at any time during the spring reduces the chance of seedling survival. Little germination occurred from July through September because of the high surface soil salinities during this period.     

Rodent seed predation promotes differential recruitment among bird-dispersed trees in temperate secondary forests

We investigated the role of seed predation by rodents in the recruitment of the fleshy-fruited trees Taxus baccata, Ilex aquifolium and Crataegus monogyna in temperate secondary forests in NW Spain. We measured the densities of dispersed seeds, early emerged seedlings, established recruits and adults, at four sites over a period of 2 years. Seed predation among species was compared by seed removal experiments and analysis of rodent larder-hoards. The three species differed markedly in local regeneration patterns. The rank order in the seed rain following decreasing seed density was Ilex, Taxus and Crataegus. However, Crataegus established 3.3 times more seedlings than Taxus. For all species, there was a positive linear relationship between the density of emerged seedlings and seed density, suggesting that recruitment was seed- rather than microsite-limited. A consistent pattern of seed selection among species was exerted by rodents, which preferred Taxus and, secondarily, Ilex seeds to Crataegus seeds. Predation ranking was the inverse of that of seed protection against predators, measured as the mass of woody coat per mass unit of the edible fraction. Recruitment potential, evaluated as the ratio of seedlings to seeds, was negatively related to seed predation, with the rank order Crataegus > Ilex > Taxus. The selective early recruitment limitation exerted by predation may have a demographic effect in the long term, as judged by the positive relationship between early seedling emergence and the density of established recruits. By modulating the pre-emptive competition for seed safe sites, rodents may preclude the progressive exclusion of species that produce low numbers of seeds (i.e. Crataegus) by those dominant in seed number (i.e. Ilex, Taxus), or at least foster the evenness for site occupation among seedlings of different species.     

Effects of germination time on seed morph ratio in a seed-dimorphic species and possible ecological significance

Background and Aims Diaspores of heteromorphic species may germinate at different times due to distinct dormancy-breaking and germination requirements, and this difference can influence life history traits. The primary aim of this study was to determine the effect of germination time of the two seed morphs of Suaeda corniculata subsp. mongolica on life history traits of the offspring.Methods Germinated brown and black seeds were sown on the 20th of each month from April to September in a simulated but near-natural habitat of the species. Phenological and vegetative traits of the maternal plants, and number, size and germination percentage of the offspring were determined.Key Results Germinated seeds sown late in the year produced smaller plants that had a higher proportion of non-dormant brown than dormant black seeds, and these brown seeds were larger than those produced by germinated seeds sown early in the year. The length of the seedling stage for brown seeds was shorter than that for black seeds, and the root/shoot ratio and reproductive allocation of plants from brown seeds were more variable than they were for plants from black seeds. Late-germinating brown seeds produced larger plants than late-germinating black seeds.Conclusions Altering the proportion of the two seed types in response to germination timing can help alleviate the adverse effects of delayed germination. The flexible strategy of a species, such as S. corniculata, that produces different proportions of dimorphic seeds in response to variation in germination timing may favour the maintenance and regeneration of the population in its unpredictable environment.     

Wild flower seed predation by Pterostichus madidus (Carabidae: Coleoptera)

The objective of the study was to investigate the ability of Pterostichus madidus, one of the most abundant carabid beetles in the United Kingdom, to consume wildflower seeds. The plant species chosen are commonly included in wildflower seed mixes sown to enhance the biodiversity of arable field margins, and hence any seed predation could potentially reduce successful seedling establishment. The study combined cafeteria style laboratory feeding experiments with predation studies in an established field margin. In laboratory studies the Pterostichus madidus consumed both unimbibed and imbibed seeds of Centaurea scabiosa, Galium verum, Leucanthemum vulgare and Primula veris. Seed consumption was still evident when Drosophila pupae were also supplied, although there was significant preference for the pupae. Placement of seeds in the field showed high levels of seed consumption, although part of this was attributed to slugs. Carabid beetles may have a significant role as wild flower seed predators and thus may affect establishment of re‐seeded flower‐rich meadows.     

Cryptic seed heteromorphism in Packera tomentosa (Asteraceae): differences in mass and germination

Seed mass variation and heteromorphism may afford plant species differential germination behavior and ultimately seedling success, particularly in disturbed habitats. We asked whether such variation occurs in Packera tomentosa (Michx.) C. Jeffrey (Asteraceae), a clonal species of the southeastern USA. Seed mass was compared within and among genetic individuals differentiated using amplified fragment length polymorphisms. We compared central and peripheral seeds produced by disc and ray florets, respectively, for their morphology, mass, and germination behavior, including response to water availability, aging, and cold stratification. Seed mass was highly variable both within and among individuals and influenced germination behavior. We found cryptic seed heteromorphism in P. tomentosa. Central and peripheral seeds had similar morphologies but dissimilar mass and biomass allocation. We used failure time analysis to detect different germination behavior. Central seeds were heavier, contained larger embryos, and germinated faster and at a higher proportion in most germination studies. Highly variable mass and heteromorphism of seeds may allow persistence of P. tomentosa in its disturbed habitats. Based on our results, some future studies of Asteraceae species with disc and ray florets may need to account for possible differences between seed types, even when morphological differences are not apparent. Evaluation of individual seed mass and maternal differences in germination studies may assist in the detection of cryptic seed heteromorphism, a phenomenon thought to be common, yet rarely documented.     

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     

Regeneration of Salix arbuscula and Salix lapponum within a Large Mammal Exclosure: The Impacts of Microsite and Herbivory

Over‐grazing or browsing by large herbivores may result in the loss of individual plant species or entire plant communities. Restoration schemes often involve exclusion of large mammals, but the resulting changes in vegetation may alter other important ecological processes such as regeneration, via changes in microsite availability for seed germination or increases in populations of seedling predators. Working within a large fenced area from which large mammals were excluded, we experimentally tested the effects of microsite, small herbivores, and their interactions on post‐dispersal seed and early seedling mortality of one nationally scarce (Salix arbuscula) and one nationally rare (S. lapponum) species of montane willow. Seeds were sown in three different microsites: natural vegetation, mown vegetation (mimicking grazed sward), and bare ground. Small exclosures and slug pellets were used to examine the effects of small mammal and slug predation, respectively. Survival of seedlings was monitored during the summer following planting. The presence of bare ground, rather than the absence of herbivores, was of over‐riding importance for early seedling survival and establishment. Protecting seedlings from small mammals made no difference to the levels of survival; however, protecting seedlings from slugs (Arion spp.) resulted in approximately 45% of seedlings surviving until the end of the summer compared to only 30% when seedlings were available to slugs. Although excluding large herbivores may increase seed production of existing individuals, the impacts of changes to plant communities on processes such as regeneration need to be considered if restoration projects are to be fully successful.     

Polyploidy affects the seed,dormancy and seedling characteristics of a perennial grass,conferring an advantage in stressful climates

• Polyploidy (the state of having more than two genome copies) is widely distributed in flowering plants and can vary within species, with polyploid races often associated with broad ecological tolerances. Polyploidy may influence within‐species variation in seed development, germination and establishment. We hypothesized that interactions between polyploidy and the seed developmental environment would affect subsequent dormancy, germination and early growth traits, particularly in stressful environments.
• Using seeds developed in a common garden under ambient and warmed conditions, we conducted germination trials under drought and temperature stress, and monitored the subsequent growth of seedlings. The study species, Themeda triandra, is a widespread, keystone, Australian native grass and a known polyploid complex.
• Tetraploid plants produced heavier, more viable seeds than diploids. Tetraploids were significantly more dormant than diploids, regardless of seed developmental environment. Non‐dormant tetraploids were more sensitive to germination stress compared to non‐dormant diploids. Finally, tetraploid seedlings were larger and grew faster than diploids, usually when maternal plants were exposed to developmental temperatures atypical to the source environment.
• Seed and seedling traits suggest tetraploids are generally better adapted to stressful environments than diploids. Because tetraploid seeds of T. triandra are more dormant they are less likely to germinate under stress, and when they do germinate, seedling growth is rapid and independent of seed developmental environment. These novel results demonstrate that polyploidy, sometimes in interaction with developmental environment and possibly also asexuality, can have within‐species variation in seed and seedling traits that increase fitness in stressful environments.

     

Seed dispersal, plant recruitment and spatial distribution of Bactris acanthocarpa Martius (Arecaceae) in a remnant of Atlantic forest in northeast Brazil

Seed dispersal ecology of Bactris acanthocarpa Mart. (Arecaceae), an Atlantic forest understory palm, was investigated during two years as an attempt to test the following predictions: (i) seeds of Bactris are dispersed by mammals and large-gaped birds; (ii) Bactris benefits from seed dispersal in terms of seed predation avoidance, improvement of seed germination and seedling survival; and (iii) spatial distribution of adults is related to patterns of seed dispersal. The study was conducted at Dois Irmãos Reserve, a 387.4-ha reserve of Atlantic forest in northeastern Brazil (8º S–35º W). Black–rumped agoutis (Dasyprocta prymnolopha) and Guianan squirrels (Sciurus aestuans) were identified as the seed dispersers/predators, moving seeds short distances (< 4 m from parents) and at low rates (0.04-0.05 diaspore/palm/day). Pyrene burial prevented seed predation by vertebrates and reduced by half seed infestation by Scolytidae beetles. Only buried pyrenes germinated. Pyrene predation was not correlated with distance from conspecific adults. In contrast, early seedling mortality was higher near conspecific adults. Most adults (64%) had their nearest conspecific adult neighbour > 4 m away in contrast to 96% of seedlings that occurred concentrated within 4 m from adults (77% under the palm crowns). Here, we present evidence that spatial distribution of B. acanthocarpa is partly due to low rates of seed removal, short-distance seed dispersal by agoutis and squirrels, and early seedling mortality associated with presence of seedlings under palm crowns.     

Reproductive potential and seedling establishment of the invasive alien tree Schinus molle (Anacardiaceae) in South Africa

Schinus molle (Peruvian pepper tree) was introduced to South Africa more than 150 years ago and was widely planted, mainly along roads. Only in the last two decades has the species become naturalized and invasive in some parts of its new range, notably in semi‐arid savannas. Research is being undertaken to predict its potential for further invasion in South Africa. We studied production, dispersal and predation of seeds, seed banks, and seedling establishment in relation to land uses at three sites, namely ungrazed savanna once used as a military training ground; a savanna grazed by native game; and an ungrazed mine dump. We found that seed production and seed rain density of S. molle varied greatly between study sites, but was high at all sites (384 864–1 233 690 seeds per tree per year; 3877–9477 seeds per square metre per year). We found seeds dispersed to distances of up to 320 m from female trees, and most seeds were deposited within 50 m of putative source trees. Annual seed rain density below canopies of Acacia tortillis, the dominant native tree at all sites, was significantly lower in grazed savanna. The quality of seed rain was much reduced by endophagous predators. Seed survival in the soil was low, with no survival recorded beyond 1 year. Propagule pressure to drive the rate of recruitment: densities of seedlings and sapling densities were higher in ungrazed savanna and the ungrazed mine dump than in grazed savanna, as reflected by large numbers of young individuals, but adult : seedling ratios did not differ between savanna sites. Frequent and abundant seed production, together with effective dispersal of viable S. molle seed by birds to suitable establishment sites below trees of other species to overcome predation effects, facilitates invasion. Disturbance enhances invasion, probably by reducing competition from native plants.     

The role of the seed coat in adaptation of dimorphic seeds of the euhalophyte Suaeda salsa to salinity

The role of the seed coat in adaptation of dimorphic seeds of the euhalophyte Suaeda salsa to salinity was investigated during germination and early seedling growth. Black and brown seeds were treated with chloroform for 1 min before the extract was used to analyze waxes and the seeds to investigate the protective role of the seed coat under saline conditions. Waxes in black seed coats were more abundant than those in brown seed coats. Salinity (500 mM NaCl) increased the concentration of Na⁺ and decreased the concentration of K⁺ in both black and brown seeds regardless of chloroform treatment. Chloroform treatment alone (in the absence of NaCl) had no effect on the concentration of Na⁺ or K⁺ in black or brown seeds and in the presence of 500 mM NaCl had no effect on the concentration of Na⁺ or K⁺ in brown seeds. However, chloroform treatment increased Na⁺ and decreased K⁺ in black seeds with 500 mM NaCl. A change of MDA (malondialdehyde) concentration in black and brown seeds treated with or without chloroform was similar to the change of Na⁺ concentration. High salinity (1500 mM NaCl) pretreatment for 40 days had a less adverse effect on germination of black seeds compared with brown seeds after they were transferred to fresh water regardless of chloroform treatment. Similar results were found for seedling emergence. In conclusion, a black seed coat may be more protective than a brown seed coat, probably by shielding the embryo from ion toxicity, because of its higher content of waxes. Thus black seeds can better maintain seed viability than brown seeds for extended periods under hypersaline conditions.     

Assessing context-dependent survival of <Emphasis Type="Italic">Quercus glauca</Emphasis> seeds in a humid maritime woodland

Whilst consumption by rodents is often invoked as a major mortality factor for large-seed species of trees, its relative importance compared with other mortality factors is poorly known. We investigated experimentally the fate of post-dispersal seeds of Quercus glauca under different understorey environments (areas covered by (i) a pteridophyte Pyrrosia lingua, (ii) a ground-vine, Trachelospermum asiaticum and (iii) no vegetation) from the germination stage to seedling emergence and establishment stages in humid maritime woodland. We employed a pair of caged and uncaged treatments to evaluate the impact of seed removal/predation by rodents, which allowed us to separate seed removal/predation mortality from mortality due to other factors. Effects by rodents were greater in the no-understorey habitat than in the Pyrrosia and Trachelospermum habitats at early stages of development, whilst non-rodent-associated mortality was relatively more important towards the seedling establishment stages in all habitats. In the absence of predation/removal by rodents (i.e. the caged treatment), more seedlings survived in the no-understorey habitat whilst seedlings were significantly taller in the Pyrrosia habitat. In contrast, no significant difference was observed in either seed/seedling survivorship or seedling height amongst habitats where seeds/seedlings were exposed to rodent predation/removal. Overall, this study in a humid maritime woodland has revealed the temporally variable influence of mortality factors and the context-dependent survival of oak seeds/seedlings, making a contrast to observations in drier woodlands; in the no-understorey environment predation/removal effect was heavier but later survivorship was higher, whilst in vegetated environments, predation/removal was reduced but survivorship was not high.     

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.