Salt stress limitation of seedling recruitment in a salt marsh plant community

Summary Seedling recruitment in salt marsh plant communities is generally precluded in dense vegetation by competition from adults, but is also relatively rare in disturbance-generated bare space. We examined the constraints on seedling recruitment in New England salt marsh bare patches. Under typical bare patch conditions seed germination is severely limited by high substrate salinities. We examined the germination requirements of common high marsh plants and found that except for one notably patch-dependent fugitive species, the germination of high marsh plants is strongly inhibited by the high soil salinities routinely encountered in natural bare patches. Watering high marsh soil in the greenhouse to alleviate salt stress resulted in the emergence of up to 600 seedlings/225 cm². The vast majority of this seed bank consisted of Juncus gerardi, the only common high marsh plant with high seed set. We tested the hypothesis that salt stress limits seedling contributions to marsh patch secondary succession in the field. Watering bare patches with fresh water partially alleviated patch soil salinities and dramatically increased both the emergence and survival of seedlings. Our results show that seedling recruitment by high marsh perennial turfs is limited by high soil salinities and that consequently their population dynamics are determined primarily by clonal growth processes. In contrast, populations of patch-dependent fugitive marsh plants which cannot colonize vegetatively are likely governed by spatially and temporally unpredictable windows of low salinities in bare patches.     

Development of vegetation patterns in early primary succession

Question: We investigated colonisation filters in early plant community development on a glacial outwash plain. We asked if these were related to seed limitation or to a lack of safe sites, if topographical heterogeneity affected species patchiness and how species life cycles influence successional trajectories. Location: An outwash plain (Skeiðarársandur) in southeast Iceland. Methods: We identified surface heterogeneity at two different scales, ca. 10–15 cm (larger stones and established plants) and ca. 50 m (shallow depressions representing dry river beds) at two study sites. We quantified species cover, flowering plant density, seed production, seed rain, seed bank density, seedling emergence and seedling survival from June 2005 to June 2007 for the whole plant community, and measured seed production for five species. Results: Mean vegetation cover was <2.5% at the sites. Low emergence rates and high seedling mortality were the two main recruitment filters. Only 1.4% of seedlings emerging in 2005 survived into the 2007 growing season. Topographical heterogeneity had little effect on plant colonisation. High annual variation was recorded, and the two study sites (ca. 2 km apart) differed in their colonisation success. Of the five species, establishment of Cerastium alpinum and Silene uniflora was most limited by lack of seeds, whereas establishment of Luzula spicata, Poa glauca and Rumex acetosella was most limited by safe sites. Conclusions: We conclude that colonisation processes and patterns in early primary succession on Skeiðarársandur were largely influenced by stochastic factors.     

Mechanisms of plant regeneration during succession after shifting cultivation in eastern Amazonia

The role of seed bank, seed rain, and regeneration from seedlings and sprouts after swidden agriculture was compared in 5-, 10- and 20-year-old secondary forest and in a primary forest in Bragantina, Pará, Brazil. The seed bank (0–5 cm soil depth) was largest in the 5-year-old forest (1190 ± 284 seeds m⁻²) and decreased nearly ten-fold with age to 137 ± 19 seeds m⁻² in the primary forest. The highest seed rain was in the 5-year-old forest (883 ± 230 seeds m⁻² year⁻¹) and the least in the primary forest (220 ± 80 seeds m⁻² year⁻¹). Large plants (≥5 cm dbh) had more individuals and species that regenerated from sprouts than from seeds and the most abundant tree species in the secondary forest stands of all ages appear to be maintained by sprouting. The smaller individuals (≥1 m tall, <5 cm dbh) in the 5-year-old forest were mainly from sprouts, but those in the older secondary forests originated mainly from seeds. These results show that at the beginning of succession, although many species can be introduced to swidden fallow from seed bank and seed rain, it is the sprout that is the main source of recruits of primary forest species in secondary forests in Bragantina.     

Arrested succession in logging gaps: is tree seedling growth and survival limiting?

Thirty years after selective timber harvest in the Kibale National Park, Uganda, many abandoned logging gaps are dominated by Acanthus pubescens, and show little forest recovery. To examine if this arrested successional state was caused by limited tree seedling growth and survival, we planted seedlings of four forest tree species (Albizia grandibracteata, Mimusops bagshawei, Prunus africana and Uvariopsis congensis) in A. pubescens‐dominated logging gaps and in control areas of adjacent forest. To assess if clearing A. pubescens facilitates forest regeneration, we planted seedlings of two species (A. grandibracteata and U. congensis) in small clearings cut within the logging gaps. We examined mortality, growth, herbivory and site characteristics among the treatments. Finally, we described the physical attributes of the A. pubescens‐dominated gaps. Seedlings of all the four species survived and grew equally well in A. pubescens and forest treatments, and most site characteristics were also similar. Seedlings planted in clearings grew more than in either forest or A. pubescens sites. Very few established trees were found in A. pubescens sites, and most of these were near the forest edges. We also discussed the role of elephants (Loxodonta africana) and collapsing A. pubescens canopies in the maintenance of an arrested successional state in these logging gaps.     

Rodent seed predation and seedling recruitment in mesic grassland

Seedling recruitment of two grasses (Arrhenatherum elatius and Festuca rubra) and two herbs (Centaurea nigra and Rumex acetosa) was measured in areas with and without rodents to which seeds of each species were sown at three seed densities (1000, 10,000 and 50,000 seeds m⁻²) in two seasons (spring and autumn 1995). Seed removal was measured for 10-day periods and the fate of seedlings was followed for 15 months after sowing. The proportion of seed removed ranged from 6 to 85% and increased with increasing seed density for each species. Rodents had no effect on seedling emergence or survival in the spring sowing. In the autumn sowing, rodents reduced seedling emergence of all four species sown at 1000 and 10,000 seeds m⁻² but had no impact at 50,000 seeds m⁻², presumably because of microsite limitation. We suggest the difference between spring and autumn arose because emergence was seed limited in autumn but microsite limited in spring; microsite availability was higher in autumn because a summer drought killed plants, reduced plant biomass and opened up the sward. Fifteen months after the autumn sowing, fewer A. elatius and C. nigra seedlings survived on plots exposed to rodents. This result reflected not only the reduced seedling emergence but also increased seedling mortality (seedling herbivory) in sites exposed to rodents. In contrast, F. rubra and R.acteosa showed density-dependent seedling survival which compensated for initial differences in seedling emergence, so that no effect of rodents remained after 15 months. The results suggest that rodent seed predation and seedling herbivory exert strong effects on seedling recruitment of A.elatius and C. nigra when recruitment conditions are favourable (conditions that lead to high microsite availability) and may contribute to both species being maintained at low densities in the grassland. The results also demonstrate that highly significant impacts of rodent seed predation at the seedling emergence stage can disappear by the time of plant maturation. Received: 2 March 1998 / Accepted: 28 September 1998     

The effects of drought and fire on seed and seedling dynamics in a tropical seasonal forest in Thailand

Seed production and seedling dynamics were studied for 6 years(1992–1997) with particular emphasis on the effects of the drought andfire in a natural tropical seasonal forest at Mae Klong Watershed ResearchStation, Kantchanaburi, western Thailand. The number of seed produced andemerged seedlings varied greatly among years and species. Most tree speciesproduced very few seeds in 1994 and many in 1995 and 1996, particularly,Shorea siamensis Miq. which displayed the most seedinghabit. Some seedlings, Shorea, Dipterocarpus alatus Roxb.,and Dipterocarpus turbinatus Gaertn. f., emerged at theendof the dry season (late April, after the first rain but when soils still have alow soil moisture content), while the majority of seedlings emerged at thebeginning of the rainy season. After a fire disturbed the plot in April 1996,many species increased their seedling emergences, especially Berryaammonilla. Seedlings of most species less than a year old showed arelatively low mortality in the rainy season, but those ofPterocarpusmacrocarpus Kurz had high mortality. The mean survival rate for allseedling species in this forest was quite low (24.1% y^–1), anddifferent values existed for the rainy and the dry season (11.5, and 6.1%month^–1, respectively). This suggests that different specieshave adaptations related to the season of seedling emergence and resistance todrought in this tropical seasonal forest community. The traits of tree speciesare classified in terms of their traits in the early stages of regeneration;seed and/or seedling bank, resprouting, and drought resistance. Most specieshave adapted to fire and/or drought by resprouting, seed bank, and/or seedlingbank, although the few species which occur mainly in mesic evergreen forestshave less adapted to these environments. The demographic variations in seed andseedling stages may contribute to the coexistence of these species in thisspecies-rich dry tropical forest.     

The effect of a seed source on primary succession in a forest ecosystem

An analysis of the Relative Importance Value Index (RIVI) of later successional tree species and the Shannon diversity indexH of all tree and shrub species was undertaken on previously mined sites within a sparsely forested area in central Florida, USA. A comparative analysis of the distance to a seed source and the age of a site suggested that the distance to a seed source was the best predictor (R ²=0.85) of the regeneration of the later successional species and a good predictor of species diversity. Both theRIVI of the later successional species and the diversity index decreased with the distance from the seed source. The lack of a seed source containing climax species resulted in arrested succession at some sites. It is suggested that biological information is the causative means of succession and the dispersal of this information entails spatial dependency while its introduction and development are time dependent processes.Research was part of a project of the Center for Wetlands, University of Florida, Gainesville under contract with the Florida Institute of Phosphate Research. Interactions of Wetlands with Phosphate Mining, H. T. Odum and G. R. Best, principal investigators. B. T. Rushton, J. Butner, G. R. Best, R. S. Schnoes and S. R. Humphrey are thanked for the use of their data.     

Testing the assumptions of chronosequences in succession

Many introductory ecology textbooks illustrate succession, at least in part, by using certain classic studies (e.g. sand dunes, ponds/bogs, glacial till, and old fields) that substituted space for time (chronosequence) in determining the sequences of the succession. Despite past criticisms of this method, there is continued, often uncritical, use of chronosequences in current research on topics besides succession, including temporal changes in biodiversity, productivity, nutrient cycling, etc. To show the problem with chronosequence-based studies in general, we review evidence from studies that used non-chronosequence methods (such as long-term study of permanent plots, palynology, and stand reconstruction) to test the space-for-time substitution in four classic succession studies. In several cases, the tests have used the same locations and, in one case, the same plots as those in the original studies. We show that empirical evidence invalidates the chronosequence-based sequences inferred in these classic studies.     

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.     

Seed mass and seedling dimensions in relation to seedling establishment

Several experiments have shown that seedlings from larger-seeded species are better able to survive various hazards during establishment. Previous work has suggested a general mechanism might underpin this outcome. Larger-seeded species might tend to mobilize their metabolic resources over a longer period into the autotrophically functioning structures of the seedling. Consequently relatively more resources would remain uncommitted at any given time during the early period of the seedling’s growth, and available to support respiration during carbon deficit. An important aspect of this larger-seed-later-commitment mechanism would be that at a given time, larger-seeded species would hold more resources uncommitted not just absolutely, but relative to the functional seedling structures that needed to be supported. Here we quantify, across a wide range of phanerocotylar species, an allometric pattern that supports the generality of a larger-seed-later-commitment mechanism as an explanation for superior performance by larger-seeded species in face of the hazards of seedling establishment. Larger-seeded species allocate relatively less to cotyledon area, reflecting the initial functional size of the seedling, and relatively more to dry mass per unit area of cotyledon, reflecting stored metabolic reserves. The shift in relative allocation is progressive, rather than seedlings falling into discrete morphological types. The allometry is similar whether considered as correlated evolutionary divergences (phylogenetically independent contrasts) or as correlation across present-day species. Received: 7 April 1999 / Accepted: 29 March 2000     

Reproductive investment and seedling survival of the mast-fruiting rain forest tree,Microberlinia bisulcata A. chev.

In the southern part of Korup National Park, Cameroon, the mast fruitingtree Microberlinia bisulcata occurs as a codominant ingroves of ectomycorrhizal Caesalpiniaceae within a mosaic of otherwisespecies-rich lowland rain forest. To estimate the amount of carbon andnutrientsinvested in reproduction during a mast fruiting event, and the consequentialseed and seedling survival, three related field studies were made in 1995.Theseprovided a complete seed and seedling budget for the cohort. Seed productionwasestimated by counting woody pods on the forest floor. Trees produced on average26,000 (range 0–92,000) seeds/tree, with a dry mass of 16.6kg/tree. Seeds were contained in woody pods of mass 307kg/tree. Dry mass production of pods and seeds was 1034 kgha^–1, equivalent to over half (55%) of annual leaflitterfall for this species, and contained 13% of the nitrogen and 21% of thephosphorus in annual leaf litterfall. Seed and young-seedling mortality wasinvestigated with open quadratsand cages to exclude vertebrate predators, at two distances from the parenttree. The proportion of seeds on the forest floor which disappeared in thefirst6 wk after dispersal was 84%, of which 26.5% was due to likelyvertebrate removal, 36% to rotting, and 21.5% to other causes. Vertebratepredation was greater close to the stem than 5 m beyond the crown(41 vs 12% of seeds disappearing) where the seed shadow was less dense.Previousstudies have demonstrated an association between mast years at Korup and highdry-season radiation before flowering, and have shown lower leaf-litterfallphosphorus concentrations following mast fruiting. The emerging hypothesis isthat mast fruiting is primarily imposed by energy limitation for fruitproduction, but phosphorus supply and vertebrate predation are regulatingfactors. Recording the survival of naturally-regeneratingM. bisulcata seedlings (6-wk stage) showed that 21% ofseedlings survived to 31 mo. A simple three-stage recruitment model wasconstructed. Mortality rates were initially high and peaked again in each ofthenext two dry seasons, with smaller peaks in the two intervening wet seasons,these latter coinciding with annual troughs in radiation. The very poorrecruitment of M. bisulcata trees in Korup, demonstratedinprevious investigations, appears not to be due to a limitation in seed oryoung-seedling supply, but rather by factors operating at theestablished-seedling stage.     

Dispersal and establishment both limit colonization during primary succession on a glacier foreland

Plant colonization can be limited by lack of seeds or by factors that reduce establishment. The role of seed limitation in community assembly is being increasingly recognized, but in early primary succession, establishment failure is still considered more important. We studied the factors limiting colonization on the foreland of Coleman Glacier, Washington, USA, to determine the importance of seed and establishment limitation during primary succession. We also evaluated the effects of seed predation, drought, and existing vegetation on establishment. We planted seeds of seven species into plots of four different ages and found evidence that both seed and establishment limitation are strong in early succession. We also found that seed and establishment limitation both remained high in later stages of succession. Seed predation reduced establishment for most species and some evidence suggested that drought and existing vegetation also limit establishment. Because both dispersal and establishment failure restrict colonization in recently exposed habitat, late-seral forest species may have a difficult time migrating upward in response to global climate change. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Seed bank assembly follows vegetation succession in dune slacks

Question: Is the seed bank in dune slacks during the whole successional range mainly composed of early successional species or does it vary according to the changing vegetation? Location: Belgium, western part of the coast. Methods: We investigated the soil seed bank composition using a seedling germination method in a chronosequence of 20 dune slacks, ranging in age from five to 55 yr. Results: Both seed density and species richness in the seed bank were very low in the first successional stages and increased with age, mainly as a result of increasing seed production. The similarity between seed bank and vegetation was higher in older slacks. A comparison of characteristics between seed bank and vegetation showed that the seed bank was, to a large extent, composed of later successional species. Occurrence patterns of individual species also showed that seeds become incorporated in the soil after the species has established in the vegetation. Conclusion: The seed bank is not likely to be the driving force for successional changes in the vegetation, and successional changes rely on dispersal. Some early successional species persist in the seed bank, but generally only in low numbers. The results also confirm that most typical dune slack species do not form persistent seeds, so that re‐establishment from the seed bank is not to be expected when the species has disappeared from the vegetation.     

Seed dispersal in fens

Question: How does seed dispersal reduce fen isolation and contribute to biodiversity? Location: European and North American fens. Methods: This paper reviews the literature on seed dispersal to fens. Results: Landscape fragmentation may reduce dispersal opportunities thereby isolating fens and reducing genetic exchange. Species in fragmented wetlands may have lower reproductive success, which can lead to biodiversity loss. While fens may have always been relatively isolated from each other, they have become increasingly fragmented in modern times within agricultural and urban landscapes in both Europe and North America. Dispersal by water, animals and wind has been hampered by changes related to development in landscapes surrounding fens. Because the seeds of certain species are long‐lived in the seed bank, frequent episodes of dispersal are not always necessary to maintain the biodiversity of fens. However, of particular concern to restoration is that some dominant species, such as the tussock sedge Carex stricta, may not disperse readily between fens. Conclusions: Knowledge of seed dispersal can be used to maintain and restore the biodiversity of fens in fragmented landscapes. Given that development has fragmented landscapes and that this situation is not likely to change, the dispersal of seeds might be enhanced by moving hay or cattle from fens to damaged sites, or by reestablishing lost hydrological connections.     

常绿阔叶林内和林窗中栲树的种子萌发和幼苗生长

对浙江天童地区常绿阔叶林林窗和林内两种生境下人工播埋的栲树（Castanopsis fargesii）种子的萌发和当年幼苗的生长进行了研究。在林窗和林内两种生境的栲树种子（单粒重约0.5-0.6g）的萌发率没有明显的差异,分别为31.7±7.1％和33.3±2.l％,一年龄幼苗的平均株高在生长季末分别为 6.56±1.2cm和 5.88±1.09cm,两者有明显差异,而平均叶长、平均叶宽以及平均根长等指标没有差异;幼苗的生物量也没有明显差异。但一年龄幼苗在生长过程中,其生物量分配有所变化,在林窗条件下,幼苗逐渐将较多的生物量分配于根,而林内幼苗则逐渐将生物量较多地分配于叶,生物量的分配反映出幼苗在不同生境下的适应性特征。     

Seed demography of three co-occurring Acer species in a Japanese temperate deciduous forest

Abstract. Seed demography of three co-occurring Acer species in an old-growth mixed deciduous forest in Japan was studied. Almost all of the seeds of A. mono germinated in the first spring, while those of A. palmatum var. amoenum showed a delay in germination of almost one year. A. rufinerve showed a rather opportunistic germination habit. Both A. palmatum var. amoenum and A. rufinerve form short-term persistent seed banks, but without input of newly dispersed seeds they may become extinct in about one year. The seed bank for these two species is not as significant as for a typical pioneer species, and the seedling bank is important for all three species. Only a small proportion of the dispersed propagules contained viable embryos, mainly due to pollination failure or abortion (A. mono and A. palmatum var. amoenum), and invertebrate predation (A. rufinerve). For all three species, larger seed crops had a higher percentage of viable seeds. Even for these relatively small, wind-dispersed seeds, the predation pressure was very high. A large part of the dispersed seeds was eaten by wood mice during the first winter (30–80 %). Estimation from the 5-yr average of seed dispersal and seedling emergence showed that only 7–16 % of the dispersed viable seeds succeeded in germinating.     

Germination responses to temperature responsible for the seedling emergence seasonality ofPrimula sieboldii E. Morren in its natural habitat

Temperature requirements for the breaking of seed dormancy and germination inPrimula sieboldii E. Morren and the annual surface-soil temperature regime in one of its natural habitats were investigated in order to clarify the germination responses determining the seedling emergence seasonality of the species. In a grassland nature reserve in an abandoned flood plain of the Arakawa River, natural seedling emergence of the species was shown to be restricted to mid- to late-spring before the closure of seasonal vegetational gaps, when the daily mean soil surface temperature reached about 15°C, accompanied by large daily fluctuations of about 10°C. Mature seeds collected in late June were never able to germinate at any constant temperature in the range of 8–40°C unless they had been previously subjected to moist-chilling treatment. The proportion of seeds which were released from dormancy increased with increasing duration of the moist-chilling treatment at 2°C, 70–85% of seeds becoming germinable at 16–28°C after 12 weeks of pretreatment at 2°C. The thermal time required for the germination of the thus-pretreated seed population was 905–1690 Kh with a base temperature of around 5°C. Fluctuating temperatures between 24°C and 16 or 12°C had a remarkable dormancy-breaking effect, inducing considerably quick germination in most of the seeds previously subjected to 2°C moist-chilling for 8 weeks.