Recruitment dynamics in a rainforest seedling community: context-independent impact of a keystone consumer

The influence of keystone consumers on community structure is frequently context-dependent; the same species plays a central organising role in some situations, but not others. On Christmas Island, in the Indian Ocean, a single species of omnivorous land crab, Gecarcoidea natalis, dominates the forest floor across intact rainforest. We hypothesised that this consumer plays a key role in regulating seedling recruitment and in controlling litter dynamics on the island, independent of the type of vegetation in which it occurred. To test this hypothesis, we conducted crab exclusion experiments in two forest types on the island and followed the dynamics of seedling recruitment and litter processing for six years. To determine if these effects were likely to be general across the island, we compared land crab densities and seedling abundance and diversity at ten sites across island rainforest. Surveys across island rainforest showed that seedlings of species susceptible to predation by land crabs are consistently rare. Abundance and diversity of these species were negatively correlated to red crab abundance. Although red land crabs may be important determinants of seedling recruitment to the overstorey, differences in overstorey and seedling composition at the sites suggests that recruitment of vulnerable trees still occurs at a temporal scale exceeding that of this study. These “windows” of recruitment may be related to infrequent events that reduce the effects of land crabs. Our results suggest that unlike the context dependence of most keystone consumers in continental systems, a single consumer, the red land crab, consistently controls the dynamics of seedling recruitment across this island rainforest.     

Supercolonies of the invasive yellow crazy ant, Anoplolepis gracilipes, on an oceanic island: Forager activity patterns, density and biomass

Summary. Ants have the capacity to reach unusually high densities, mostly in their introduced ranges. Numerical dominance is often cited as key to the ability of exotic ants to displace native ant species, reduce the abundance of invertebrates and negatively impact upon bird, land crab and other vertebrate populations. On Christmas Island, Indian Ocean, the yellow crazy ant, Anoplolepis gracilipes (Jerdon), forms supercolonies, where extremely high densities of foraging ants have contributed to ‘invasional meltdown’ in rainforest areas. Densities of up to 2254 foraging ants per m² and a biomass of 1.85 g per m² were recorded, and nest densities reached 10.5 nest entrances per m². Populations of A. gracilipes can overcome and kill red endemic land crabs (Gecarcoidea natalis) over 100 times their own biomass. This is the highest recorded density of foraging ants, and adds another element to the definition of ‘supercolony’ of unicolonial ants. This paper documents one extreme in a continuum of densities of unicolonial, invasive ant species and highlights the need to incorporate forager densities into invasive ant research.Received 17 November 2004; revised 14 February 2005, accepted 21 February 2005.     

Viable soil seed banks in disturbed lowland tropical rainforest sites in North Queensland

Four 0.125 m² soil samples, 50 mm deep, from each of eight tropical lowland rainforest sites ranging from primary and secondary forests to degraded Imperata grasslands were exposed to shade-house conditions. Germinations of soil seeds were monitored over 28 weeks in April-October. The viable seed banks in the primary forest (592 m^-2) and the secondary forest sites resulting from one cycle of clearing and abandonment (1300–1400 m^-2) were composed predominantly of secondary species. The viable seed banks were small in Imperata grassland sites that had been burnt regularly (200–400 m^-2) and very large (2000–3000 m^-2) in sites that had not been burnt for five years or more. The seed banks of the burnt and unburnt Imperata grassland sites were composed mostly of agricultural weeds whilst seeds of secondary rainforest trees were rare or absent. Similarities between the species composition of soil seed banks and regrowth vegetation types suggest that soil seed composition is important in determining the initial floristic composition following disturbance and that lack of seed of secondary rainforest trees is one factor which restricts re-growth and secondary succession on degraded Imperata grasslands. Species such as Eucalyptus intermedia R. T. Bak., E. pellita F. Muell. and Imperata cylindrica (L.)Beauv. which can establish and deflect the normal rainforest secondary succession were not present in the soil seed bank, and these species appear to rely on fresh seed inputs, lignotubers or rhizomes to colonize and regenerate after disturbance.     

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.     

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.     

Resistance of Island Rainforest to Invasion by Alien Plants: Influence of Microhabitat and Herbivory on Seedling Performance

Experiments have promise in determining mechanisms by which communities resist invasion. Growth and survivorship of transplanted seedlings of introduced tree species (Leucaena leucocephala, Muntingia calabura, Adenanthera pavonia, and Clausena excavata) were used to assess abiotic (light regime) and biotic resistance (herbivory) to invasion of rainforest on Christmas Island (Indian Ocean). At four sites, seedlings were transplanted into the forest edge along roadside verges and into adjacent intact forest; half were caged to prevent access by the dominant seedling consumer, the red land crab (Gecarcoidea natalis). Red crab densities did not differ between roadside and forest plots. Red crabs initially reduced survivorship of Leucaena in both edge and interior plots but virtually all seedlings in the forest interior were dead after 41 weeks. Survivorship of Muntingia was also initially reduced by crabs in the forest edge, but again, all but one seedling died by the end of the experiment. Seedlings of Adenanthera and Clausena fared much better overall, surviving well in both locations. Red crabs had no overall effect on Adenanthera survival but significantly reduced survival of Clausena in forest plots. For both species, seedling performance was greater in the forest edge than in the forest interior. Red crabs had no effect on height increment for either species; however, for Clausena, red crabs reduced seedling mass in the forest interior. Both Adenanthera and Clausena were able to persist in the intact forest. Clausena is now actively invading intact rainforest, but Adenanthera appears dispersal-limited. Resistance factors in intact forest appear hierarchical: biotic resistance afforded by land crabs can impede establishment of some plant invaders but seedling responses to abiotic factors (e.g., the light regime) largely overwhelm its effect. Together, these two community attributes are likely to restrict the range of plant invaders to a small suite of species that can successfully establish in intact rainforest on the island. However, increased propagule pressure from a variety of shade-tolerant species and further declines in the abundance of the dominant seedling consumer may lead to increasing invasion success in this island rainforest. This revised version was published online in July 2006 with corrections to the Cover Date.     

The secondary invasion of giant African land snail has little impact on litter or seedling dynamics in rainforest

In the absence of empirical evidence, invasive species are often assumed to have negative impacts because of their conspicuously high abundance. The giant African land snail Achatina (Lissachatina) fulica is one such invader where its impact in natural ecosystems remains completely untested. On Christmas Island (Indian Ocean), A. fulica has become established across large tracts of rainforest following the impacts of invasive yellow crazy ant (Anoplolepis gracilipes) in mutualism with non‐native scale insects. Yellow crazy ants facilitate the secondary invasion of A. fulica by extirpating native red land crabs (Gecarcoidea natalis) that are normally effective predators of A. fulica. We used a multifaceted approach to investigate some potential impacts of abundant A. fulica in invaded rainforest. Over the course of a wet season, diel activity transects showed that A. fulica consumed detrital material almost exclusively. However, stable isotope analysis did not confidently identify A. fulica as a predominantly detritivorous species. We found no statistically significant treatment effects of A. fulica exclusion on standing leaf litter and seedling recruitment processes during a 6‐month manipulative field study. However, litter cover and biomass did remain slightly higher where A. fulica were excluded, albeit with overlapping confidence intervals with control plots. Our study constitutes the first empirical test for impact of A. fulica in a natural ecosystem and suggests that for Christmas Island rainforest, this species is not a damaging invader. Other studies will need to assess the impacts of A. fulica in other natural areas before these findings could be considered broadly applicable.     

Monopolization of litter processing by a dominant land crab on a tropical oceanic island

Litter processing by macroinvertebrates typically involves suites of species that act together to determine rates of breakdown and decomposition. However, tropical oceanic islands and coastal fringes on continents are often dominated by one or a few species of omnivorous land crabs that consume leaf litter. We used an exclusion experiment, together with other leaf removal and litter decomposition studies, to assess the role of a single dominant species, the red crab (Gecarcoidea natalis), in litter dynamics in rain forest on Christmas Island, Indian Ocean. In the presence of red crabs, litter cover and biomass varied seasonally, from almost complete cover and high biomass at the end of the dry season to almost total absence of litter at the end of each wet season. When crabs were excluded from both the shaded understory and light gaps in rain forest, litter increased rapidly to almost complete cover, which was then maintained year round. Leaf tether experiments, and measures of litter input and standing crops, indicated that red crabs monopolize litter processing, removing between 39 and 87% of the annual leaf fall from the forest floor. Rates of litter turnover were over twice as high in the presence of land crabs: the decomposition constant, k, was 2.6 in the understory exclusion plots, but rose to 6.0 in the presence of crabs. Red crabs occur at biomass densities (114 g m⁻²) far greater than those reported elsewhere for entire litter faunas. They significantly reduced the abundance of other litter invertebrates, but we did not detect any change in the relative frequencies of the major invertebrate groups (mites, collembolans, pulmonate snails, ants, psocopterans, and spiders). Wherever omnivorous land crabs are abundant, their activities may be paramount in litter decomposition and in regulating the rate of nutrient cycling. In monopolizing litter processing, they may also be important physical “ecosystem engineers”, translocating organic matter and nutrients into the soil and reducing available habitat for other animals. Received: 19 August 1998 / Accepted: 11 January 1999     

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.     

Persistent soil seed banks in old-growth Hyrcanian Box tree (Buxus hyrcana) stands in northern Iran

Little is known about the soil seed bank and the influence of plant communities on the interaction between the seed bank and aboveground vegetation in the Hyrcanian temperate deciduous forest. We surveyed species composition and diversity of the persistent soil seed bank and the aboveground vegetation in six community types in old-growth Hyrcanian Box tree (Buxus hyrcana) stands in northern Iran. Fifty-two species with an average of 3,808 seeds/spores m⁻² germinated; forbs accounted for 64% of the seed bank flora. Thirty-four species in the aboveground vegetation were not presented in the seed bank, 32 species in the seed bank were not found in the vegetation, and 20 species were in both. The dominant tree species were Diospyros lotus and Alnus subcordata with an average of 17 and 4.6 seeds m⁻², respectively. Our results suggest that (1) vernal geophytes and shade-tolerant perennials are not incorporated in the seed bank, (2) early successional species are well represented in the seed bank, (3) plant community type has significant impacts on seed bank densities, and seed bank richness and diversity were significantly related to presence/absence of Box tree in the aboveground vegetation. The persistent seed bank contained species that potentially have a negative impact on the regeneration of forests, thus forest managers should retain old-growth Hyrcanian Box tree stands to conserve disturbance-sensitive indicator forest species.     

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.     

Relationships among the seed rain,seed bank and vegetation of a Hawaiian forest

Abstract. Hawaiian ecosystems are prone to invasion by alien plant species. I compared the seed rain, seed bank, and vegetation of a native Hawaiian forest to examine the potential role that seed ecology plays in allowing alien species to invade native forest. Absolute cover of seed plants in the forest was 126 %, annual seed rain was 5 713 seeds m^-2 yr^-1, and the mean density of seedlings emerging from the seed bank averaged across four seasons was 1 020/m². The endemic tree Metrosideros polymorpha was the most abundant species in the vegetation, seed rain and winter seed bank. Overall, native seed plants comprised 95 % of the relative cover in the vegetation and 99 % of the seeds in the seed rain, but alien species comprised 67 % of the seeds in the seed bank. Alien species tended to form persistent seed banks while native species formed transient or pseudo-persistent seed banks. Dominance of the seed bank by alien species with persistent seed banks suggests that aliens are favorably placed to increase in abundance in the vegetation if the forest is disturbed.     

Invasive ants compete with and modify the trophic ecology of hermit crabs on tropical islands

Invasive species can dramatically alter trophic interactions. Predation is the predominant trophic interaction generally considered to be responsible for ecological change after invasion. In contrast, how frequently competition from invasive species contributes to the decline of native species remains controversial. Here, we demonstrate how the trophic ecology of the remote atoll nation of Tokelau is changing due to competition between invasive ants (Anoplolepis gracilipes) and native terrestrial hermit crabs (Coenobita spp.) for carrion. A significant negative correlation was observed between A. gracilipes and hermit crab abundance. On islands with A. gracilipes, crabs were generally restricted to the periphery of invaded islands. Very few hermit crabs were found in central areas of these islands where A. gracilipes abundances were highest. Ant exclusion experiments demonstrated that changes in the abundance and distribution of hermit crabs on Tokelau are a result of competition. The ants did not kill the hermit crabs. Rather, when highly abundant, A. gracilipes attacked crabs by spraying acid and drove crabs away from carrion resources. Analysis of naturally occurring N and C isotopes suggests that the ants are effectively lowering the trophic level of crabs. According to δ¹⁵ N values, hermit crabs have a relatively high trophic level on islands where A. gracilipes have not invaded. In contrast, where these ants have invaded we observed a significant decrease in δ¹⁵ N for all crab species. This result concurs with our experiment in suggesting long-term exclusion from carrion resources, driving co-occurring crabs towards a more herbivorous diet. Changes in hermit crab abundance or distribution may have major ramifications for the stability of plant communities. Because A. gracilipes have invaded many tropical islands where the predominant scavengers are hermit crabs, we consider that their competitive effects are likely to be more prominent in structuring communities than predation. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

天山中部天山云杉林土壤种子库年际变化

通过定点采样,采用萌发法对天山中部天山云杉(Picea schrenkiana Fisch.et Mey.)近熟林(101-120a)和成熟林(121-160a) 2004-2011年(8a)土壤种子库物种组成、种子密度的年际变化和不同间隔年限土壤种子库物种组成的相似性进行了分析.结果表明:(1)土壤种子库中共萌发鉴定出种子植物87种,隶属29科70属,其中乔木种子植物2种,灌木种子植物2种,草本种子植物83种,土壤种子库中草本植物种子密度远远大于木本植物种子密度;8个采样年份土壤种子库恒有种仅有6种;(2)土壤种子库种子密度及其中天山云杉种子密度存在巨大的年际变动,且不具有同步性;土壤种子库种子密度最大(2009年)值为(953.75±66.12)粒/m2,最小(2008年)值为(186.50±20.37)粒/m2,其中天山云杉种子密度最高(2006年)达到(584.50±53.58)粒/m2,最低(2005年)仅有(0.25±0.26)粒/m2;(3)天山云杉林土壤种子库年际间物种组成的相似性不高,Czekanowski相似系数均值仅为0.344,并随间隔年限的增加呈现减小—增大—减小的变化趋势.天山云杉林土壤种子库物种组成和种子密度稳定性差,年际间相差悬殊,物种组成的相似性不高,种子库中天山云杉种子密度主要受其种子库采样前一年天山云杉结实丰歉的影响,属间断型.土壤种子库年际变化特征可为天山森林的更新恢复和可持续经营提供科学依据.     

Fire and regeneration: the role of seed banks in the dynamics of northern heathlands

Questions: How do species composition and abundance of soil seed bank and standing vegetation vary over the course of a post‐fire succession in northern heathlands? What is the role of seed banks – do they act as a refuge for early successional species or can they simply be seen as a spillover from the extant local vegetation? Location: Coastal Calluna heathlands, Western Norway. Methods: We analysed vegetation and seed bank along a 24‐year post‐fire chronosequence. Patterns in community composition, similarity and abundances were tested using multivariate analyses, Sørensen's index of similarity, vegetation cover (%) and seedling counts. Results: The total diversity of vegetation and seed bank were 60 and 54 vascular plant taxa, respectively, with 39 shared species, resulting in 68% similarity overall. Over 24 years, the heathland community progressed from open newly burned ground via species rich graminoid‐ and herb‐dominated vegetation to mature Calluna heath. Post‐fire succession was not reflected in the seed bank. The 10 most abundant species constituted 98% of the germinated seeds. The most abundant were Calluna vulgaris (49%; 12 018 seeds m^?2) and Erica tetralix (34%; 8 414 seeds m^?2). Calluna showed significantly higher germination the first 2 years following fire. Conclusions: Vegetation species richness, ranging from 23 to 46 species yr^?1, showed a unimodal pattern over the post‐fire succession. In contrast, the seed bank species richness, ranging from 21 to 31 species yr^?1, showed no trend. This suggests that the seed bank act as a refuge; providing a constant source of recruits for species that colonise newly burned areas. The traditional management regime has not depleted or destroyed the seed banks and continued management is needed to ensure sustainability of northern heathlands.     

Disturbance Effects on the Seed Bank of Mexican Cloud Forest Fragments1

The density and floristic composition of the soil seed bank was assessed in six cloud forest fragments with different levels of human disturbance in central Veracruz, Mexico. A total of 8416 seeds germinated in 60 soil samples, at 5‐cm depth, corresponding to 107 species, 85 genera, and 48 families. Significant differences were found among study sites in seed densities with values ranging from 873 to 3632/m². Tree species contributed 20 percent of the total soil seed bank in four sites and herbs accounted for the majority of the species in each site. Among tree species, Trema micrantha displayed the highest seed density, accounting for 84 percent of the germinated seeds. In general, the tree species composition of the soil seed bank did not closely reflect the composition of the tree community. Results suggest that disturbance produced by human activities (trail use, selective cutting of trees, livestock) may influence the size and composition of the soil seed bank in forest fragments. Sites where human activity has been reduced showed the highest proportion of dormant seeds.     

Supercolony mosaics: two different invasions by the yellow crazy ant, Anoplolepis gracilipes, on Christmas Island, Indian Ocean

Invasive species are one of the main reasons for the ongoing global loss of biodiversity. Anoplolepis gracilipes is an invasive ant that has recently received significant attention due to its negative effect on the native fauna and flora of Christmas Island, Indian Ocean. This species has contributed to a drastic change in the structure of the Christmas Island rainforest through its negative impact on the island’s endemic red land crab, the dominant consumer on the islands forest floor. In this study, we investigate the population structure of A. gracilipes on Christmas Island in order to determine whether multiple introductions occurred on the island and how they correspond to known infestations. We genotyped 578 individuals collected from 50 nests across the Island. We identify two distinct subgroups in the population that represent two different supercolonies. These supercolonies are interspersed across the island, however both nuclear (microsatellites) and mitochondrial markers strongly suggest that there is no gene flow between the two colonies. Significant heterozygote excess within the entire sampling area, with all but one worker examined being heterozygous for all seven microsatellite loci, suggests an unusual reproductive system in these ants. Our results are consistent with recent sociogenetic findings in a population of A. gracilipes in Northern Borneo.     

Long-Term Effects of Forest Regrowth and Selective Logging on the Seed Bank of Tropical Forests in NE Costa Rica1.

To investigate long-term effects of land use on the soil seed bank, we compared the abundance/density, species richness, life form distribution, and species composition of seeds stored in the soil of four 15–20 yr-old second-growth stands, two old-growth stands, and two previously selectively-logged stands in the Caribbean lowlands of Costa Rica. Surface soil (10 cm deep, 4.7 cm diameter) was collected at 10 m intervals along three 120–160 m long transects in each stand (44–48 soil cores, 22–24 combined seed bank samples per site). Seed density was highest but variable in second-growth stands (8331–14535 seeds/m²), low and homogeneous in old-growth stands (2258–2659 seeds/m²), and intermediate and highly variable in selectively-logged stands (1165–6854 seeds/m²), which also had contrasting logging intensities. Species richness was strongly dependent on seed density, but showed less variation. Life form distribution did not differ statistically among or within land-use categories. In each stand, herbs-forbs, shrubs, and vines dominated the seed bank (> 75% of the species richness and abundance), whereas trees were a minor component (< 20% of the species richness and < 5% of the abundance) and were predominandy early successional. Shrubs and vines were most abundant in second-growth stands where regrowth vegetation was repeatedly cut before abandonment, whereas grasses and sedges were most abundant in the only forest stand that was completely surrounded by pastures. In terms of species composition, old-growth stands were more similar to selectively-logged stands than to second-growth stands, but across stands, selectively-logged forests were most distinct from the other two forest types. An inventory of the standing woody vegetation in each site showed little representation of the woody taxa found in the seed bank. We discuss these results in the context of the main factors that have been postulated to influence the abundance, life form, and species composition of tropical forest seed banks, and explore the role of the latter during intermediate phases of tropical forest succession and regeneration.     

Comparative seed germination ecology of Austrostipa compressa and Ehrharta calycina (Poaceae) in a Western Australian Banksia woodland

Austrostipa compressa, a native ephemeral of southwest Western Australia was stimulated to germinate under a range of temperatures, in the presence of light, and exposure to smoke-water. This combination of environmental cues results in winter-maximum germination in immediate postfire and disturbed-soil environments of this Mediterranean-type climate. In contrast, Ehrharta calycina, an introduced perennial grass from southern Africa that has invaded Banksia woodlands, germinated under a wide range of temperature and light conditions, but showed no promotive response to smoke-water. Although A. compressa seeds tolerated heat shock better than E. calycina, the self-burial mechanism of A. compressa seeds ensures protection from fire. High-intensity fire could have a greater impact on E. calycina, as the seeds of this species tend to accumulate in the top of the soil profile where they are more susceptible to high temperatures. Although seeds of E. calycina are more susceptible to high temperatures, survival of mature individuals by postfire resprouting ensures continued survival in native woodlands. Estimates of soil seed bank densities showed extreme variability, but some recently burnt areas of the Yule Brook Botany Reserve contained up to 8000 seed m^?2 of A. compressa and nearly 75 000 seeds m^?2 of E. calycina. Viable soil seed bank densities of A. compressa are reduced with time-since-last fire, but areas of greater than 45 years since the last fire, still contained up to 119 seeds m^?2. In both species, only about half their soil seed bank germinates following fire, thus ensuring the potential for later recruitment. Massive soil seed populations of E. calycina in native Banksia woodlands pose a major problem to management of this plant community type.     

Soil seed bank and regeneration potential on eroded hill slopes in the Kondoa Irangi Hills,central Tanzania

Abstract. The soil seed bank of the severely eroded Kondoa Irangi Hills, Tanzania was studied in order to determine the seed density and composition and to establish the relationship between seed bank and standing vegetation. The area had not been grazed for 15 yr prior to the study. A recently grazed area was used for comparison. The seed bank density (at 0 - 5 cm depth) ranged from 344 to 915 8 seeds/m² in the dry season and 172 to 5107 seeds/m² in the wet season. The seed bank was very heterogeneous, both spatially and temporally, and it showed significant variation in size and composition in both sampling periods. The species similarity between the seed bank and the above-ground vegetation in all plots was low (Sørensen's index = 0.00–0.44). The highest similarity was found in the recently grazed area. The seed bank was dominated by annuals and by early successional species. It is concluded that re-vegetating the hill slopes with woody vegetation by using the seed bank will be difficult because seeds of woody species were not found in the soil.