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.     

Comparative demography of co-occurring introduced and native tussock grasses: persistence and potential expansion

Summary Demographic characteristics associated with the maintenance and growth of populations, such as seed dynamics, seedling emergence, survival, and tiller dynamics were examined for two tussock grasses, the native Agropyron spicatum and the introduced Agropyron desertorum in a 30-month field study. The introduced grass was aerially sown onto a native grassland site. Seed production of the introduced grass was greater than the native grass in both above- and below-average precipitation years. Seeds of A. spicatum were dispersed when they mature, while A. desertorum retained some seeds in inflorescences, and dispersed them slowly throughout the year. This seed retention allowed some seeds of the introduced grass to escape peak periods of seed predation during the summer and allowed seeds to be deposited constantly into the seed bank. Carryover of seeds in the seed bank beyond one year occurred in the introduced grass but not in the native species. For both species, seedling emergence occurred in both autumn or spring. Survival rates for A. desertorum were higher than A. spicatum when seedlings emerged between November and March. Survival rates of cohorts emerging before November favored A. spicatum whereas survival rates did not differ between species for cohorts emerging after March. Individuals of both species emerging after April were unable to survive the summer drought. Demographic factors associated with seeds of A. desertorum seemed to favor the maintenance and spread of this introduced grass into native stands formerly dominated by A. spicatum.     

Burial of canopy-stored seeds in the annual psammophyte Agriophyllum squarrosum Moq. (Chenopodiaceae) and its ecological significance

Agriophyllum squarrosum Moq. is a dominant annual on sand dunes in the arid regions of central Asia. A high percentage of seeds is retained on dead plants which become covered by moving sand, but little is known about the ecological significance of burial of canopy-stored seeds. We investigated the size and dynamics of the buried canopy-stored seed bank and effects of burial on seed germination. In March (during the windy season), May (beginning of the germination season), and July (middle of the growing season), the number of seeds per square meter in sample plots in the dunes was 623, 223 and 22, respectively, with 54.6, 30.6 and 12.9% of the total seeds retained on buried plant canopies. In a controlled experiment, more seedlings emerged from released (dispersed) than from canopy-stored seeds when burial depth was the same. No viable ungerminated released seeds were found, but 45–80% of the ungerminated canopy-stored seeds were viable. In general, with an increase in applied water germination of released seeds buried at a depth of 1 or 2 cm and of canopy-stored seeds buried at 1 cm increased, but regardless of watering regime few or no released seeds at 4 cm or canopy-stored seeds at 2 or 4 cm germinated. Significantly more seedlings emerged from plants buried in a horizontal than in a vertical position. Seedlings originating from buried canopy-stored seeds on an active dune accounted for only 5.4% of the total seedlings emerging, and most of them emerged later than those from released seeds. Thus, seed release is more effectively postponed in buried than in exposed canopies, and burial of canopy-stored seeds is a mechanism that helps regulate seed germination and seedling emergence of A. squarrosum on active dunes.     

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.     

茂兰喀斯特森林中华蚊母树群落土壤种子库动态初探

 茂兰喀斯特森林中华蚊母树群落900m2样地内共有种子植物59种,其中19种主要植物已繁殖产生种子。群落每年以种子雨形式向种子库输入种子351.1粒·m-2,其中对群落更新有用的成熟有效种子150.8粒·m-2。土壤种子库中活力种子丰富,共有41种2510.5～2646.8粒·m-2。种子库中活力种子的种类和数量均随时间的变化而变化,不同时间种子库的差异主要在于现存植物活力种子的变化。土壤种子库每年输出活力种子171.9粒·m-2,其中萌发输出21.9粒·m-2,动物采食输出133.5粒·m-2,病菌危害引起的输出为3.2粒·m-2,种子自然衰老引起的输出为13.3粒·m-2。每年输出超过输入21.1粒·m-2。演替前期植物种子没有输入,只有缓慢输出,但没有萌发输出。     

Seed dynamics during forest succession in Costa Rica

Soil seed banks and current seed inputs each play a role in tropical succession. We compared the abundance and floristic composition of seeds from these two sources at a Costa Rican site by germinating seeds from the soil, measuring seed inputs for 3 yr, and monitoring the earliest colonists in a forest clearing.There were an estimated 6800 viable seeds/m² in the soil of 3.3-yr-old vegetation, 9500 seeds/m² in 11-yr-old vegetation, and 7000 seeds/m² in a 75-yr-old forest. An estimated 10100 seeds/m² fell on the soil surface of the young successional vegetation during 3 yr and 3700 seeds/m² fell during that same time in the forest.Locally produced seeds accounted for about 75% of the seed input to the soil surface early in succession. Seeds dispersed out of young successional vegetation increased the quantity and species richness of the seed input and storage in an adjacent forest. Much of the species richness of the young successional vegetation resulted from seeds dispersed there from other communities by animals.Deforestation stimulated germination of most seeds in the surface soil of the old forest, including seeds of the dominant canopy tree. The recruitment of seedlings from the soil seed bank numerically overwhelmed that from post-disturbance seed rain and sprouts.We evaluated patterns of soil seed storage during succession and predicted the ability of vegetation of differing ages to respond to disturbance. Immediately after disturbance the number of seeds in the soil plummeted due to mortality, low inputs, and germination. As the vegetation regrew, the soil seed bank increased to a peak after 4 to 7 yr, then gradually decreased to its pre-disturbance size. High-frequency pulses of disturbance should result in reduced species richness, dominance by species with long-lived seeds, and fast recovery by seedling recruitment from the soil seed bank.Journal series number 6459 from the Institute of Food and Agricultural Sciences, University of Florida, Gainesville, Florida 32611, USA.Reprint requests to J. J. E. at Florida.     

Germination and seedling survival in fens undergoing succession

We studied mechanisms of vegetation change in fens subject to succession from open water to floating mats and finally herbaceous rich-fens. Earlier research showed that these systems are characterized by transient seed banks. Our main question was whether seedlings of later successional fen stages are already present in earlier stages, remaining subordinate in the vegetation until conditions become suitable for them. If, however, conditions during succession change in a way that only a limited set of species can survive as seedlings during each of the successional stages, no seedling bank will exist. The transient character of the seeds would then imply that seeds will not germinate and will subsequently die and that seeds that have germinated in the “wrong” stage will not become established. We hypothesized that: (1) germination and seedling survival of fen species are significantly better in the successional fen stage for which these species are characteristic, (2) as a consequence no seedling banks occur in these fens. In a field experiment, seeds of five characteristic fen species in the standing vegetation of three successional fen stages i.e. raft fen, quaking fen and rich fen were sown in each of these stages in a turf pond in the Tienhoven area, The Netherlands. Germination and seedling survival were measured over two growing seasons together with environmental variables. Germination was higher in the “own” stage for all species groups as was survival for quaking fen species and rich fen species. For both these stages, percentage of germination and survival of four out of five characteristic species were significantly higher in the “own” stage. Germination and survival can be considered stage-dependent and it was concluded that seedling banks do not exist in these fens. Site-specific environmental variables act as a sieve and differentiate on species presence already during early life history stages. We found clues that the environmental sieve acts at the level of nutrient availability, tolerance for high sulphide concentration and light climate. Because of the transient seed bank and absence of a seedling bank in these fen wetlands, successful establishment of species necessitates a continuous dispersal of characteristic species until the environmental conditions permit establishment. This also implies that species of the whole successional sere should be present within dispersal distance.     

黄土丘陵沟壑区主要物种植冠种子库动态及其生态策略

植冠种子库是植物适应环境并应对外界干扰的种子生态策略之一,研究了黄土丘陵沟壑区12种主要植物植冠种子库动态,结果表明:杠柳不具有植冠种子库,其他11种植物均具有植冠种子库;除了黄刺玫种子在翌年5月达到脱落高峰,其他植物大部分种子在冬季脱落,其中杠柳、达乌里胡枝子、茭蒿、黄柏刺和水栒子的大部分种子脱落集中偏早,铁杆蒿和土庄绣线菊的大部分种子脱落集中偏晚;植冠宿存对大部分植物种子的萌发特性表现为促进作用;但不同植物种子的萌发时滞对植冠宿存响应差异较大;9种植物种子在植冠上宿存至翌年2月底,其种子活力仍能维持达60%以上;该区植物表现出不同的植冠种子库策略,通过不同的方式来减少干扰的威胁,提高成功萌发与更新的几率,它们或具有较大规模的宿存量、或调控种子萌发特性、或提高种子维持活力的百分比。此外,全面了解该区植物形成植冠种子库的机理及对应的生态策略还有待于全面、深入的研究。     

Seed Ecology of the Invasive Tropical Tree <Emphasis Type="Italic">Parkinsonia aculeata</Emphasis>

Parkinsonia aculeata is an invasive tree native to tropical America, but introduced to Australia. Propagation and stand regeneration is mainly by seed. To gain baseline knowledge for management decisions, seed bank dynamics were monitored for two months during the fruit dispersal period at a coastal wetland in Costa Rica (native habitat), and at a coastal wetland and two semi-arid rangeland sites in Northern Queensland, Australia (introduced habitats). Seed bank densities underneath dense, uniform Parkinsonia stands were found to be lowest in the Australian wetland but highest in the Costa Rican wetland. Post-dispersal seed losses were highest in the Australian wetland, primarily due to seed germination and/or death. At the other sites, seed losses were minor during the study period, and predation was the most important cause of losses. At the two rangeland sites bruchid beetles accounted for more than 95% of the seed losses by predation. Total predation was lowest in the Costa Rican wetland. In order to test for intrinsic differences of seed characteristics, germination trials were conducted using both canopy seeds and seeds from the soil seed bank. Dormancy release and germination rate were studied under four temperature treatments. In all populations, dormancy release increased with increasing temperature, but averaged responses were significantly different between Costa Rican and Australian seed populations, and between seeds collected from the soil and from trees. Germination rate of scarified seeds was fastest at 35 °C in all tested seed populations. While high seed germination levels seem to explain low seed bank densities in the Australian wetland, the large seed banks at the rangeland sites reflect the lower incidence of favourable conditions for germination. In the Australian wetland biocontrol with bruchids is unlikely to be successful, while control by conventional methods, such as killing stands by basal bark spraying, seems feasible, due to a lower long-term risk of re-infestation from the soil seed bank. At the rangeland sites conventional control will be difficult and costly. Parkinsonia stands may be better left to their own, while bruchid populations are monitored and management efforts are concentrated on preventing further invasion.     

Seed development and germination ecophysiology of the invasive tree <Emphasis Type="Italic">Prunus serotina</Emphasis> (Rosaceae) in a temperate forest in Western Europe

Seed development, dormancy and germination of the American invasive tree species, Prunus serotina, are described for plants growing in a large forest in Belgium. Seeds of P. serotina were collected following anthesis in the first week of July and thereafter at fortnightly intervals. Seed dormancy, temperature requirements for germination and the soil seed bank were investigated. At maturation (about 105 days after anthesis), seed moisture content had decreased to around 13.7%, and 44% of the seeds had attained the capacity to germinate. Mature seeds of P. serotina exhibited physiological dormancy, germinating only after a long cold, moist stratification period. Highest germination percentage occurred in seeds treated with gibberellic acid (GA₃), at 10°C. We found no evidence that P. serotina forms a persistent seed bank but noticed a persistent seedling bank in the field.     

六盘山区辽东栎的实生苗更新及其影响因子

研究了六盘山区辽东栎(Quercus liaotungensis)灌丛的种子产量、土壤种子库组成、湿沙和风干贮藏对种子寿命的影响以及动物取食子叶对种子萌发和幼苗建立的影响。结果表明, 辽东栎灌丛种子的完好率为27.51%, 被动物取食或搬运种子的比例(41.51%)显著高于其他类型种子(p < 0.01); 辽东栎次生林土壤种子库中萌发和虫蛀种子分别占35.16%和38.29%, 完好种子仅占13.65%, 捕食动物主要通过贮藏或搬运而影响土壤种子库中的种子密度。湿沙贮藏60天的辽东栎种子自动萌发率高达96.67%, 短期贮藏可加快种子的萌发进程, 提高萌发率和萌发指数, 但随着贮藏时间的延长, 种子萌发进程延迟, 萌发率、萌发指数和活力指数均不同程度地降低。排除动物取食处理的幼苗在林窗和林下生境的存活率分别为80%和83%, 而不排除动物取食幼苗在2种生境中分别仅有25%和31%能够存活, 表明子叶在幼苗建立中具有重要作用。林窗中幼苗子叶的动物取食率(85.00%)高于林下(71.00%), 子叶留存的幼苗在林窗中的存活率(6.00%)低于林下(15.50%), 而子叶被取食幼苗的存活率在两种环境中基本相等(分别为18.50%和18.00%)。     

The impact of flooding and drought on seeds of Cnidium dubium, Gratiola officinalis, and Juncus atratus, three endangered perennial river corridor plants of Central European lowlands

The effects of flooding and drought on the post-flooding survival and germinability of seeds in the soil were investigated for three endangered river corridor angiosperms Cnidium dubium (Schkuhr) Thell., Gratiola officinalis L. and Juncus atratus Krocker. Freshly matured seeds were placed in small nylon bags and were buried in pots filled with soil collected from a wetland along the Lower Havel River. The pots were subjected to different winter/spring flooding treatments (30, 60, 120, and 180 days) and to drought for up to 3 years. Every year in spring, after drawdown, the seeds were exhumed, counted and tested for germination. Seed mortality of flooded seeds was low, except for C. dubium. Flooding did not affect germination of seeds of J. atratus. Flooding of seeds of C. dubium and G. officinalis had inconsistent, but no beneficial effects on germination of either. The year of harvest and the interannual variability of weather are found to be the main factors affecting germinability of C. dubium and G. officinalis in the course of the three consecutive years.The experiment showed that the capacity of the flooded seed bank to maintain germination ability and viability for extended periods of time could be directly related to the flooding duration of natural habitats of the three species. The three species are resilient to variable flooding. Seeds appeared desiccation tolerant, and single flooding events do not deplete their seed bank.     

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.     

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     

A STUDY OF THE SEED BANK OF A SALT MARSH IN NORTHERN SAN FRANCISCO BAY

Abundance, species composition, and distribution of buried seeds in a San Francisco Bay salt marsh were studied by collecting soil samples in October and February and observing seedling emergence in the greenhouse. Results were compared with existing vegetation patterns and field germination. Average numbers of buried viable seed down to a 5-cm depth were 380/ m² in October and 700/m² in February, with field germination averaging 118/m². Salicornia virginica dominated the seed bank in the greenhouse and in field germination. Most other marsh species were present in the seed bank but numbers of seeds were low. A significant correlation was found between highest species diversity and proximity to channels. The nature of this low-diversity, low-density seed bank reflects dominance of long-lived perennial species, seed dispersal patterns and selective environmental pressures.     

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.     

植冠与土壤种子库储存种子的萌发特性及策略

为明确黄土丘陵沟壑区植物种子库如何调控种子萌发来提高个体适合度,选择研究区7种具有种子库的主要物种为研究对象,以刚成熟和室内储存种子为对照,比较植冠宿存(5个宿存期)和土壤埋藏(5a埋藏期)对植物种子萌发特性的影响,探讨植冠种子库与土壤种子库储存下的种子萌发策略。结果表明:7种植物种子经过不同种子库储存后萌发特性表现出明显的种间差异,黄刺玫(Rosa xanthina)和水栒子(Cotoneaster multiflorus)种子萌发力表现为植冠宿存不变型、土壤储存增强型,土壤储存明显提高水栒子种子萌发速率;达乌里胡枝子(Lespedeza davurica)和狼牙刺(Sophora davidii)种子萌发力表现为植冠宿存增强型、土壤储存减弱型,种子萌发历时表现为植冠宿存延长型,土壤种子库储存还可加快达乌里胡枝子萌发速率、缩短萌发历时;茭蒿(Artemisia giralaii)和铁杆蒿(Artemisia gmelinii),种子萌发率随植冠宿存时间先升高后降低,随土壤储存时间先降低后升高,土壤储存可推迟其萌发,铁杆蒿种子萌发速率在植冠与土壤储存后均加快;紫丁香(Syringa oblata)种子萌发率随植冠宿存先升高后降低,土壤储存明显加快其种子启动萌发与速率。在黄土丘陵沟壑区,植物种子经过植冠或土壤种子库储存,或增加、加快、提早萌发充分利用有利条件提高占据性,或减少、减缓、推迟萌发分摊不利条件的风险;而且该区植物植冠与土壤储存后种子萌发特性间的关系,体现各自适应环境与应对干扰的分工与协作策略,主要表现为:单一主导型和相辅相成型。     

Seed bank patterns in Restionaceae and Epacridaceae after wildfire in kwongan in southwestern Australia

Abstract. Post-fire seed germination, seedling mortality and seed banks were investigated in scrub-heath (kwongan) in SW Australia. Study species included herbaceous and woody, obligate seeders and resprouters in two non-bradysporous but significant plant families (Restionaceae and Epacridaceae). In all species, seedlings were recruited only in the first autumn-spring after fire and occurred in similar densities as the estimated germinable annual seed input. Seedlings were absent from unburnt vegetation. Although most species retained some residual seeds after fire, tests (excised embryo culture) indicated that a negligible number of seeds were germinable. Regardless of fire response or species, there appeared to be a large loss of seeds each year and in most cases, only a small proportion of the annual seed production was used in post-fire recovery of plants. Based on seedling: parent ratios, all species had the capacity to reconstitute parent densities from germinants in the first year after fire, but high seedling mortality and no further recruitment resulted in less seedlings than replacements for four resprouter Restionaceae and three Epacridaceae (all obligate seeders) at the end of the third year after fire.     

Hydrochory,seed banks,and regeneration dynamics along the landscape boundaries of a forested wetland

Following the environmental sieve concept, the setting in which the recruitment of Taxodium distichum occurs in, becomes increasingly restrictive from the seed to seedling stage in an impounded forested wetland. Although a wide elevational band of dispersing seed moves across the boundary of a swamp-field in the water sheet, the zone of germination is relegated to that portion of the forested wetland that draws down during the growing season. Seedling recruitment is further restricted to the uppermost zone of the winter water sheet. These patterns are likely applicable to other species of dominant swamp species, e.g., Cephalanthus occidentalis crossed the boundary of a forested wetland and abandonded field in winter flooding (November–December and November–March, respectively) in Buttonland Swamp. The elevation of the boundary was 101.3 m NGVD. While the seeds of at least 40 swamp species were dispersed across the boundary, few viable seeds were dispersed after the winter season. Kriged maps showed seeds of T. distichum and C. occidentalis dispersed in patches in the water depending on the position of the water sheet. Most species of both water- and gravity-dispersed species had a localized pattern of seed distribution (either spherical or exponential) and this indicated that seeds may not be dispersed for great distances in the swamp. Water-dispersed T. distichum and C. occidentalis had larger dispersal ranges (A ₀=225 and 195 m, respectively) than Bidens frondosa and B. discoidea (A ₀=14 and 16 m, respectively). Seed dispersal varied with season depending on the availability of seeds. In Buttonland Swamp, viable seeds typically were dispersed for T. distichum in November–June, and for C. occidentalis in November-July. Low water occurred in August 1993 and high in February 1994 (99.8 and 101.6 m NGVD, respectively). The seed banks along the landscape boundary varied in species composition according to elevation (r ² = 0.996). While the similarity of species richness between water-dispersed seeds and the seed bank at elevations that flooded (during June 1993 through May 1995) was high (10–17%), it was low between water-dispersed seeds and the seed bank at elevations that did not flood (5%). T. distichum seeds had a short germination window in that seeds germinated within a year following their production in zones that were flooded in the winter followed by drawdown during the next growing season. After 1 year, less than 5% of the T. distichum seeds remained viable on the surface of the soil. Germination of T. distichum was confined to specific elevations (above 99.3 but below 101.6 m NGVD) during this study with 4.1% of the seedlings surviving for more than 2 years at a mean of 101.4 m NGVD. All seedlings below this elevation died. To maximize natural regeneration along the boundaries of swamps in abandoned farm fields targeted for restoration, this study suggests a flood pulse regime consisting of high water in the winter to maximize dispersal of live seeds followed by low water in the summer to facilitate seed germination and seedling recruitment. Hydrologic restoration could assist in the natural recovery of damaged wetlands if a seed source exists nearby.     

Regeneration filters in post-agricultural fields of Puerto Rico and Ecuador

To better understand how rainforest regenerates after agriculture I sampled the seed rain and seed bank, and set out seeds and seedlings, on microsites defined by distance to the forest in fields both in Puerto Rico and Ecuador. I found that (1) total seeds, species richness and life-form richness were twice as great in the Ecuador seed rain compared to Puerto Rico but Puerto Rico seeds were more evenly distributed among species and (2) total seedlings from the seed bank were similar between Puerto Rico and Ecuador, (3) the majority of seeds were lost to predation among all species and study sites, (4) seed disease was absent in P. aduncum and Miconia prasina, and no seeds germinated for Gonzalagunia spicata and P. riparia, (5) in Ecuador pathogenic disease claimed more seeds than germinated for all species, and Solanum ovalifolium was the only species that had seeds germinate but did not lose seeds to disease, (6) also in Ecuador, insect predation was significantly lower in the forest border for P. aduncum, and seed disease was significantly greater at the 10 m micro site for S. ovalifolium, (7) distance has a significant effect on seedling height and basal diameter, (8) losses of leaf area due to herbivory and pathogens were always low and (9) biomass and leaf specific mass were significantly reduced in the border and forest microsites. I conclude that Ecuador fields had more seeds, species, and life-forms than Puerto Rico fields, predation was the most severe post-dispersal seed filter in all fields, seeds that survived predation on Puerto Rico were lost either to disease or germination but to both mechanisms in Ecuador, all three seed mechanisms in Ecuador fields showed distance effects of seedling growth but not survivorship.