Vegetation and seed bank in a calcareous grassland restored from a Pinus forest

Question: What is the importance of the seed bank in the maintenance of the restoration potential of a 60‐year‐old abandoned calcareous grassland overgrown by Pinus trees? Location: ‘Les Pairées’, province of Luxembourg, Belgium. Methods: The seed bank and the above‐ground vegetation were surveyed in three adjacent stands, previously forming a unique calcareous grassland: a 60‐year‐old Pinus forest, a four‐year‐old clear‐cutting and a typical calcareous grassland. Floristic diversity was compared among stands and between vegetation and seed bank. Results: The species richness of the vegetation and the seed bank was significantly lower in the Pinus forest. More floristic similarities were found between the clear‐cutting and the calcareous grassland. Seed bank was essentially transient, dominated by annual species. Its correspondence with the above‐ground vegetation was weak. Conclusion: Very few calcareous grassland species have persisted in the Pinus stand. Four years after clear‐cutting, the stand was nearing restoration towards a calcareous grassland. Seed longevity in the soil was not the most explicative factor. Dispersal of propagules from adjacent sources was also important.     

阿拉善干旱荒漠区不同植被类型土壤种子库研究

[1]兰州大学草地农业科技学院,兰州730020 [2]甘肃草原生态研究所,兰州730020 [3]内蒙古阿拉善盟草原站,巴彦浩特750360     

Abundance and habitat segregation in Mediterranean grassland species: the importance of seed weight

Abstract. We analysed the relationship between seed traits (weight, shape and dispersal structures) and the abundance and habitat segregation of Mediterranean grassland species. To take into account possible correlations with other plant traits, the study also includes 5 vegetative traits (growth form, plant longevity, clonality, onset of flowering and plant size) of commonly accepted functional importance. Data were recorded for 85 species from dehesa grasslands in central Spain. Species abundance was measured in upper (dry and less productive, high stress) and lower (moist and more productive, low stress) slope zones in the same area. Habitat segregation was estimated using an index based on the relative frequencies of species in upper and lower slope zones. Multiple regression models were fitted using species, as well as phylogenetically independent contrasts, as data points. Annual small‐seeded species without specialised dispersal structures are over‐represented in dehesa grasslands. Abundance was negatively related to seed weight in upper slope zones. None of the recorded plant traits were related to abundance in the lower slope zones. Habitat segregation was mainly related to seed weight, but also to some vegetative traits. Annual, early flowering and small‐seeded species were relatively more abundant in the upper than the lower slope zones. This pattern is independent of phylogeny. Our results suggest that in dry Mediterranean grasslands, abundance of many species is determined by dispersal (production of numerous small seeds) rather than by competitive ability.     

Variation in exotic and native seed arrival and recruitment of bird dispersed species in subtropical forest restoration and regrowth

Invasive bird-dispersed plants often share the same suite of dispersers as co-occurring native species, resulting in a complex management issue. Integrated management strategies could incorporate manipulation of dispersal or establishment processes. To improve our understanding of these processes, we quantified seed rain, recruit and seed bank density, and species richness for bird-dispersed invasive and native species in three early successional subtropical habitats in eastern Australia: tree regrowth, shrub regrowth and native restoration plantings. We investigated the effects of environmental factors (leaf area index (LAI), distance to edge, herbaceous ground cover and distance to nearest neighbour) on seed rain, seed bank and recruit abundance. Propagule availability was not always a good predictor of recruitment. For instance, although native tree seed rain density was similar, and species richness was higher, in native plantings, compared with tree regrowth, recruit density and species richness were lower. Native plantings also received lower densities of invasive tree seed rain than did tree regrowth habitats, but supported a similar density of invasive tree recruits. Invasive shrub seed rain was recorded in highest densities in shrub regrowth sites, but recruit density was similar between habitats. We discuss the role of microsite characteristics in influencing post-dispersal processes and recruit composition, and suggest ways of manipulating these processes as part of an integrated management strategy for bird-dispersed weeds in natural areas.     

Biodiversity and spatial distribution of vascular epiphytes in two biotopes of the Cameroonian semi-deciduous rain forest

The diversity and spatial distribution of vascular epiphytes were surveyed in two biotopes (dryland forest and swamp-inundated forest) of the semi-deciduous rain forest area in Cameroon. Eight sites in each biotope were selected, which included 530 individuals of phorophytes in dryland forest and 460 in swamp-inundated forest. A total of 148 epiphyte species were recorded, which showed that semi-deciduous rain forests represent a major source of African-epiphyte diversity. Dryland forest hosted 110 epiphyte species, while the swamp-inundated forest harboured 108. A total of 56 species were found only in the dryland forest and 60 were restricted to the swamp-inundated forest. At family level, Orchidaceae exhibited the highest-species richness within both biotopes. Pteridophytes were more abundant in the swamp-inundated forest. A TWINSPAN analysis of the floristic similarities separated the two investigated forest types very clearly. No significant difference existed between the two biotopes regarding vertical distribution of epiphytes within single trees. The swamp-inundated forests may serve as a refuge for many epiphytes that occur in the outer canopy of the dryland forests, both because they are inherently less vulnerable to timber extraction, and enjoy better protection by legislation.     

Fourteen months of seed rain in three Australian semi‐arid communities

The abundance and composition of seed rain was measured over 14 months (February 2004 to March 2005) in Currawinya National Park, western Queensland. The experimental design included four measurement periods, three vegetation communities and two grazing regimes. A total of 12 586 seeds from 104 species were captured. There were significantly more seeds and species captured during the measurement period with the least rainfall, although no significant correlation was found between the amount of rainfall and the number of seeds or species captured. More seeds and species were captured where native and feral grazing pressure was removed, but this was only significant for the number of species. The above‐ground vegetation showed no significant difference between grazing treatments over the study period and exhibited far fewer species than the seed rain. However, the majority of species found in the above‐ground vegetation were represented in the seed rain. Hypotheses are explored as an attempt to understand the apparent lack of a relationship between seed rain and rainfall. The effects of grazing and seed movement and storage are also discussed.     

Seed dispersal patterns in a temperate forest during a mast event: performance of alternative dispersal kernels

Seed dispersal patterns were studied in a north-western Spain temperate forest community to assess the performances of alternative dispersal kernels during two years with ecologically contrasting scenarios; a non-mast year, and a mast year of the dominant canopy species, beech Fagus sylvatica. Dispersal kernels were fitted under a Bayesian modeling framework. Both simple and mixture kernels were considered for the five more abundant tree species (Corylus avellana, Crataegus monogyna, F. sylvatica, Ilex aquifolium and Taxus baccata). Mixture kernels provided a better fit for almost all species, and the log-normal performed best for T. baccata. No relationship between dispersal syndromes and the best dispersal kernel function emerged. However, we found temporal changes in the shape of the dispersal kernels that seemed to be related to variation in relative fruit production among species and the resulting changes in the responses of dispersal vectors. This reveals a potential role for disperser-mediated indirect effects in terms of introducing temporal variation in species spread. In this sense, our results highlight the need to consider single species seed dispersal as a community process. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Stabilization of metapopulation cycles: toward a classification scheme

The stability of population oscillations in ecological systems is considered. Experiments suggest that in many cases the single patch dynamics of predator-prey or host-parasite systems is extinction prone, and stability is achieved only when the spatial structure of the population is expressed via desynchronization between patches. A few mechanisms have been suggested so far to explain the inability of dispersal to synchronize the system. Here we compare a recently discovered mechanism, based on the dependence of the angular velocity on the oscillation amplitude, with other, already known conditions for desynchronization. Using a toy model composed of diffusively coupled oscillators we suggest a classification scheme for stability mechanisms, a scheme that allows for either a priori (based on the system parameters) or a posteriori (based on local measurements) identification of the dominant process that yields desynchronization.     

Seed bank versus seed rain in the regeneration of a tropical pioneer tree

Summary We used the tropical pioneer tree, Cecropia obtusifolia to evaluate the relative importance of different sources of seeds in the regeneration of species that depend on ephemeral sites. We studied seed production in a population established in a 5 ha plot, and dispersal, dormancy and seed predation in two recent treefall gaps (<1 year-old), two building or successional forest patches (10–15 since disturbed), and two mature forest patches (>35 years since disturbed) for a one year period at Los Tuxtlas (Mexico). Flowers and fruits were counted at monthly intervals. Annual fecundity per tree ranged from 1.4×10⁴ to 1.4×10⁷ seeds. Seeds were continuously available on the trees and on the ground. Average annual seed rain per m² (as measured by 0.5×0.5 m seed traps) varied from 184 to 1925 among the six sites. Distance to nearest seed source and patch type explained more than 60% of the seed rain variation among sites. Soil seed density, estimated by counting seeds from ten samples (78.5 cm²×10 cm deep) collected from each site in October and January, ranged among the six sites from 269 to 4485 seeds per m² in January and from 204 to 5073 in October. Soil seed viabilities were much lower (17.1% in October and 5.1% in January) than those of rain seeds (48.26%). Annual survivorships of 2.2% were estimated for seeds artificially sown on the soil surface of a gap and a mature patch, and 3.75% in a building patch. In two other experiments seed removal rates ranged from 27% to 98% in 4 days. Removal rates were significantly higher in gap and mature patches than in building patches. Ants (Paratrechina vividula) and grasshopper nymphs (Hygronemobius. sp.) were the main predators. We draw three main conclusions from our data: (1) Pathogens and predators determine low survivorship of C. obtusifolia's seeds in the soil and a rapid turnover rate (1.07 to 1.02 years) of its seed bank; (2) a continuous and copious seed production and an abundant and extensive seed rain replenish the soil seed pool in patches with different disturbance ages at least up to 86 m from nearest source; (3) more than 90% of the seeds contributing to C. obtusifolia seedling recruitment in gaps are less than one year-old. We discuss our results in the context of previous similar studies for tropical forests.     

Effect of three mound-building ant species on the formation of soil seed bank in mountain grassland

The effect of three ant species (Lasius flavus, Formica spp., Tetramorium caespitum) on soil seed bank formation was studied in temperate mountain grassland. Seed removal experiments, analysis of soil seed content and seed survival experiments were carried out to evaluate the influence of ground ants on the seed fate. In the seed removal experiment seeds of 16 species, including 5 species with elaiosome-bearing seeds (myrmecochores), were exposed and their removal followed for 39 h. On average, ants removed 63.8% of myrmecochorous seeds and 10.9% of seeds without adaptation to ant dispersal. Analysis of soil seed content revealed that myrmecochores, in spite of expectations that they would accumulate in nests of seed dispersing ants, were most abundant in the soil of control plots. Evidence on seed relocation to the ant nests was obtained from a comparison of mounds of seed dispersing and seed non-dispersing ant species, as more seeds were found in the mounds of Formica spp. and Tetramorium caespitum (seed dispersers) in comparison with the mounds of Lasius favus (non-disperser).The soil seed bank of the compared microhabitats (control plots and mounds of 3 ant species) differed in their species composition, seed abundance and vertical distribution. The most distinct qualitative differences were between seed flora of control plots and mounds of Tetramorium caespitum. Control plots had approximately 30,000 propagules per m², which was double the number of seeds found in the ant mounds. In control plots, abundance and diversity of seeds steeply declined with depth; this trend was not observed in the mounds probably due to bioturbation. In the seed survival experiment, more seeds (2 out of 3 species) survived in control plots, which may also contribute to the higher seed abundance in this microhabitat.This study showed that seed relocation by ants does not contribute significantly to seed bank build-up at this study site. Ants may, however, increase the regeneration success of myrmecochores, mainly by dispersal for distance and placement in a larger spectrum of microsites, in contrast to species not adapted for myrmecochory.     

Biodiversity of settled material in a sediment trap in the Gulf of Trieste (northern Adriatic Sea)

Phytoplankton succession and sinking rates were studied from January to December 2003 at a coastal station in the Gulf of Trieste (northern Adriatic Sea), 200 m offshore, in a relatively undisturbed area. A conical sediment trap, moored at 15 m depth (water depth 17 m), was used. The hypothesis if the presence of benthic and epiphytic diatoms can lead to an overestimation of the vertical fluxes was tested. To evaluate primary and secondary sedimentation contributions, planktonic, benthic and epiphytic diatoms were distinguished. Benthic species abundance varied throughout the year and it was related to resuspension that strongly influenced sinking rates. All over the year, diatoms were the prevailing class in the trap material accounting for 75.32% of the settled cells, while flagellates represented 24.11%. Dinophyceae and resting cells constituted minor components, accounting for 0.43% and 0.14%, respectively. The gross sedimentation rates ranged from 0.006 × 10⁸ cell m⁻² d⁻¹ in the second week of May to 6.30 × 10⁸ cell m⁻² d⁻¹ in the third week of January with a mean annual value of 1.09 ± 1.43 × 10⁸ cell m⁻² d⁻¹. To the primary sedimentation rate Pseudo-nitzschia seriata of the group “Nitzschia seriata complex” contributed for 49.77% followed by Chaetoceros spp. (23.88%). The major contributor to the secondary sedimentation rate was the diatom Paralia sulcata, accounting for 24.76%. Epiphytic diatoms contributed for 11.19% and 12.27% on annual average gross abundance and biomass, respectively, reaching even 72.04% of gross abundance and 56.06% of gross biomass in the second week of August. The correlation between temperature and the logarithm of the epiphytic biomass was statistically significant, with r = 0.66 and P < 0.001. Both in the cluster analysis and in the PCA four main groups were formed, where benthic and epiphytic species were separately gathered. Planktonic, benthic and epiphytic forms accounted for 50.78%, 36.95% and 12.27%, respectively, calculated on the annual average biomass. Therefore, vertical fluxes can be overestimated of 50% or more if benthic and epiphytic species are not rejected.     

浙江天童常绿阔叶林栲树种子雨的时空格局

在浙江天童常绿阔叶林中选择5株栲树,通过种子筐和地表取样,分析了栲树种子雨的时空格局特征.结果表明,栲树种子雨从11月3日到12月16日,持续45d,存在散落的高峰期(11月18日～27日);散落在地表的种子雨主要集中在离母株树干1.5～5.5m区段内,占总数的85.6%,最远不超过8m,空间格局符合二次分布;整个种子雨期间,单株平均落果数为112.8个·m^-2(70个·m^-2～239.5个·m^-2),东南向较多(135.6个·m^-2)、西北向较少(86.8个·m^-2),但无显著差异;种子雨中有活力的坚果占总数的27.3%、未成熟的占25.7%,昆虫幼虫侵害的占42%,松鼠啃食的占5%.     

The role of seed dispersal in the natural regeneration of rain forest after strip-cutting in the Peruvian Amazon

Seed dispersal and forest regeneration were studied on a 30×150 m strip cleared by strip-cutting, a system of forest management designed for sustained yield (Hartshorn 1989), in high terrace rain forest in the Department of Loreto, Peru. After one year the strip was dominated by seedlings of a few bat- and bird-dispersed pioneer tree species (Cecropia spp., Melastomataceae, and Alchornea triplinervia); stump sprouts from cut trees and saplings that survived the clearing were less numerous. The density of saplings (>2 m in height) surviving the clearing was 903 per hectare; 94% of these survived the subsequent 18 months. About 30% of 417 stumps (>7.5 cm diameter at breast height) resprouted within 3 months, with an additional 10% sprouting in the subsequent 10 months. Sprouting frequency was greater for small stumps than large and varied greatly among plant families. Seed deposition over this year was much lower in the interior of the strip, both in species richness and numbers of seeds, than within the forest; strip edges were intermediate in richness and number. The decline in seed input from forest to edge to strip, both in species and in numbers of seeds, was most pronounced for bird-dispersed taxa (primarily Melastomataceae); bat- and wind- dispersed taxa were more evenly distributed. The similarity in bat species composition between the strip and nearby primary forest was higher than the similarity in bird species composition between these habitats, reflecting a failure of many forest bird species to venture into the strip. The predominance of Cecropia spp. and other pioneers of minimal commercial value in the regeneration question the sustainability of strip-cutting. Subsequent succession and future tree species composition on the cleared strip will depend not only on the survivorship and growth of sprouts, survivors, and seedlings, but also on responses of different seed-dispersing animal taxa to changes in the species composition and structure of the vegetation in the strip.     

Factors influencing the seed source and sink functions of a floodplain nature reserve in the Netherlands

Question: How do species traits and abiotic factors influence the extent of hydrochorous dispersal into and out of a small floodplain area along a free‐flowing river in The Netherlands? Location: The Kappersbult nature reserve (53°07′28″N, 6°37′14″E), which is a floodplain along the Dutch River Drentsche Aa. Methods: Seeds transported by the river were collected in fine mesh nets for 24 consecutive hours once or twice a week for 1 year, upstream and downstream of the studied floodplain. Data on the captured seeds were related to species traits and abiotic factors and species composition in the floodplain. Results: The floodplain functioned both as a seed source and sink. High levels of river water seemed to promote seed transport to or from the floodplain. Seeds of riverbank species occurred significantly more often in the river water than expected. Net source species had significantly higher seed production, taller stature and higher seed buoyancy, but lower site elevation than net sink species. Seed weight was significantly higher for sink species than for other species. Conclusion: Our study found that inundation, and therefore more natural river water management, is a prerequisite for seed transport to and from a floodplain. The restoration of target floodplain vegetation may be successful for common species that produce many seeds and grow in proximity to the river. Consequently, it is expected that the probability of restoring vegetation types that occur further from the river, such as wet grasslands, by hydrochorous dispersal is low.     

The role of soil seed banks in the restoration of dry acidic dune grassland after burning of Ulex europaeus scrub

Question: Can the seed bank play a significant role in the restoration of plant communities of dry acidic dune grassland where fire has destroyed Ulex europaeus scrub? Location: Northern French Atlantic coast. Methods: One year after the fire, the seed bank and vegetation were sampled in 1 m × 1 m plots along three transects from the oldest scrub vegetation towards the grassland. Differences in species richness, seed density and contribution of ecological groups in the seed bank and vegetation along the transects were analysed. Results: Seed density and species richness in the seed bank decreased significantly from the grassland towards the centre of the scrub vegetation; 50% of the seed bank consisted of core species of the target plant community, such as Carex arenaria, Aira praecox, Rumex acetosella and Agrostis capillaris. Seeds of these species were also found in the deeper soil layers beneath the oldest scrub vegetation, indicating that they can be considered to be long‐term persistent. Beneath the youngest scrub vegetation, seeds of rare satellite target species also occurred. However, no target species were established on the burned site after one year, resulting in a large discrepancy between seed bank and vegetation. Conclusions: Although the seeds present in the soil indicate that restoration of the acidic grassland based on the seed bank is possible, additional management actions such as mowing and soil disturbance may be necessary to restrict resprouting of Ulex and to stimulate the germination of seeds of target species in the deeper soil layers.     

Habitat degradation,topography and rainfall variability interact to determine seed distribution and recruitment in a sand dune grassland

Question: Understanding the mechanisms underlying how habitat degradation, topography and rainfall variability interactively affect seed distribution and seedling recruitment is crucial for explaining plant community patterns and dynamics. Interactions between these major factors were studied together in a semiarid sand dune grassland. Location: Eastern Inner Mongolia, China. Methods: The study system used four sites of fixed, semifixed, semishifting and shifting sand dune grasslands, representing a gradient of habitat degradation. We investigated the density of germinable seeds deposited in the top 5 cm of soil and in situ seedling emergence (number of seedlings emerging early in the growing season) and establishment (number of plants recruited at the end of the growing season) at three topographic positions (dune top, windward and leeward sides) within each site over 2 years that differed in rainfall. Habitat characteristics (i.e. vegetation cover, plant species composition and diversity, soil moisture and nutrient availability and soil erodibility) of the four sites were also measured. Results: Habitat degradation (i.e. decreased vegetation cover and enhanced wind erosion rate) significantly reduced the size of the germinable soil seed bank. On average, germinable seed number from the high‐vegetation cover fixed dune was 36‐fold larger than the low‐vegetation cover shifting dune, and eight‐ and two‐fold larger, respectively, than the semishifting and semifixed dunes with intermediate vegetation cover. We observed within‐habitat variability in seed distribution, but among‐topographic position variation differed among habitats. Seedling recruitment showed large between‐year, and among‐ and within‐habitat variability, but these variations varied significantly depending on the response variables evaluated (i.e. initial seedling density, final plant density, emergence rate and recruitment rate). Path analysis revealed complex density‐dependent positive and negative, direct and indirect effects of germinable seed density and initial seedling density on recruitment, but the relative importance of these density‐dependent effects varied depending on habitat type and rainfall availability. Conclusion: Our results suggest that habitat degradation, microtopography and rainfall availability interact in shaping sand dune seed bank and plant community recruitment patterns and dynamics. Their effects were mainly mediated through changes in both the biotic and abiotic environment during the process of habitat deterioration.