Establishment and growth of seedlings of Eucalyptus obliqua: Interactive effects of litter,water, and pathogens

We studied how leaf litter, water and fungal pathogens affect the establishment, survivorship, and growth of Eucalyptus obliqua using glasshouse and field experiments. In a glasshouse experiment, the presence of leaf litter of E. obliqua increased the establishment of seedlings when the seeds were placed beneath the litter, but dense litter layers reduced seedling emergence when seeds were placed on top. Results from a complementary experiment suggest that the effect of litter may be mediated by the high humidity microenvironment it creates, which may enhance imbibition of unburied seeds. In another glasshouse experiment, litter in combination with excessive water applications reduced seedling emergence and increased seedling mortality, but the addition of fungicide reverted these effects. Litter with lower water application or fungicide added increased seedling emergence. In all cases higher biomass in seedlings growing in litter treatments was associated with earlier emergence. In two recently burnt sites, the density of seedlings of E. obliqua was positively correlated to the presence of litter. Experimental addition of litter in the field reduced soil temperature during the summer, and together with watering, increased soil water content, and the water potential of established seedlings. However, neither treatment had any positive effect on their growth or survivorship. On the contrary, litter addition reduced the height and number of leaves, probably as a result of increased herbivory. We conclude that in Mediterranean-type systems litter may produce various effects on seedling establishment depending on their developmental stage, and the season.     

The role of seed depth,litter, and fire in the seedling establishment of amphicarpic peanutgrass (Amphicarpum purshii)

Summary Amphicarpum purshii is an annual grass which mostly grows in disturbed areas of the New Jersey Pine Barrens, USA. It is amphicarpic, producing spikelets (and seeds) both above and below the soil surface. Previous research has shown that subterranean seed production ensures reproduction in the event of a major disturbance such as fire and results in rapid post-burn colonization of these sandy habitats. The effects of fire, litter, and seed depth were further examined by planting subterranean seeds at four depths in 16 litter-covered flats buried at ground level and comparing plants arising from burned flats with those in undisturbed litter-covered flats. At 0 and 1 cm depth, rates of seedling emergence were lowest in burned flats. Surface-sown seeds produced seedlings more likely to desiccate. Sowing depth had a greater influence on most measured characters than burning treatments. The mean depth of subterranean seed placement by Amphicarpum is 3.5 cm and this coincides with the seed depth from which plants showed the greatest height growth, shoot biomass, and reproductive output. In a second experiment, subterranean seeds on the bare soil surface in clay pots were more likely to lose viability and less likely to germinate than seeds protected by litter or burial in soil. In addition to providing protection from fire, placement of seeds below ground in the sandy habitat of peanutgrass provides conditions more suitable for seed survival and subsequent seedling establishment.     

Clonal integration enhances survival and performance of Potentilla anserina, suffering from partial sand burial on Ordos plateau, China

On Ordos plateau, a semi-arid, desertified area in China, sand burial is a common stress factor for plants. The extent to which sand burial occurs is heterogeneous and unpredictable in space and in time. Therefore, clonal fragments (i.e., interconnected ramets of a clonal plant) often experience partial sand burial, with some ramets buried in sand while the rest may remain unburied. It was hypothesized that clonal fragments are able to benefit from clonal integration, in case they experience partial sand burial. A pot experiment was conducted with Potentilla anserina, a stoloniferous herb often found on Ordos plateau. We used clonal fragments consisting of four interconnected ramets. In the experiment, the two proximal (older) ramets were unburied while the two distal (younger) ramets were either unburied (control) or buried with a 2, 4 or 6 cm deep layer of sand (burial treatments). The stolon connection between the proximal and the distal ramets was either severed or left intact. Stolon severing dramatically decreased the survival of buried ramets. Stolon severing and sand burial had significant effects on plant performance in terms of biomass production, number of leaves and leaf area. A cost–benefit analysis based on performance measures shows that the proximal ramets supported their connected distal ramets and did not incur any cost from this resource export. These results suggest that clonal integration, which is one of the functionally most important consequences of clonal growth, contributes significantly to our test species' capacity to withstand partial sand burial on Ordos plateau, a semi-arid and desertified area of China.     

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.     

Ignorant seed predators and factors affecting the seed survival of a tropical palm

In addition to acting as seed predators, some terrestrial mammals bury seeds via scatter hoarding. This study system used two permanent plots in examining the interaction between small rodents and the seeds of the palm Astrocaryum mexicanum. We tested how experimental burial, and fruiting status of the parent, distance to the parent, seed size, and microsite characteristics affect the survival of these seeds. Up to 34% of the buried seeds that were exposed only to ignorant rodent foragers (individuals not responsible for burial) survived. In comparison, less than 1% of seeds buried by scatter hoarding rodents survived in previous studies, a percentage that is comparable to the low survival of unburied seeds in this study (<2%). Although unburied seeds had very low survival, increasing distance and/or seed density positively affected survival of unburied seeds. Distance to parent had no effect on buried seed survival.
Buried seed survival was most strongly and significantly determined by the fruiting status of the trees under which they occurred. Seeds experienced significantly greater predation if buried under “parent” trees that fruited during the experiment. Buried seed survival was also negatively affected by germination, as germination may signal the presence of a seed to foraging rodents. There was some indication of a positive effect of tree density on seed survival between the two plots, whereas differences in rodent abundance appear to have no effect on seed survival. Seed size and microsite characteristics had no significant effect on buried seed survival, likely due to the greater proportional effects of other factors and the longevity of A. mexicanum seeds. The results of this study were used to generate a hypothetical causal network showing how comparatively low recovery of buried seeds by ignorant foragers – combined with processes determining the removal of scatter hoarding foragers from their scattered seed caches – may affect seedling recruitment in A. mexicanum.     

沙埋对半灌木羊柴幼苗的存活、生长和生物量分配的影响

羊柴(Hedysarum laeve Maxim.)是中国北方沙地飞播进行植被恢复与重建的主要植物种.在晚春与夏初的出苗期 ,羊柴幼苗经常遭受不同深度的沙埋.研究旨在探明沙埋对羊柴幼苗存活、生长和生物量分配格局的影响.6周的沙埋试验结果显示:当沙埋深度达到其株高时,约有70%的幼苗死亡 ;沙埋深度达到其株高的133%时,可使羊柴幼苗全部死亡;沙埋深度分别为株高的33%和67% 时,羊柴幼苗的整株生物量、叶片生物量、根系生物量以及相对生长率都相应地高于非沙埋的对照(即:0%沙埋).与非沙埋的对照相比,羊柴幼苗在33%和67%沙埋条件下并不显著地改变其生物量分配格局;并且羊柴幼苗的叶数和株高与对照无明显差异,但试验期内的新生叶数是沙埋处理(即:33%、67%和100%沙埋)的幼苗明显地高于非沙埋的对照.     

沙埋对半灌木羊柴幼苗的存活、生长和生物量分配的影响

羊柴 (HedysarumlaeveMaxim .)是中国北方沙地飞播进行植被恢复与重建的主要植物种。在晚春与夏初的出苗期 ,羊柴幼苗经常遭受不同深度的沙埋。研究旨在探明沙埋对羊柴幼苗存活、生长和生物量分配格局的影响。6周的沙埋试验结果显示 :当沙埋深度达到其株高时 ,约有 70 %的幼苗死亡 ;沙埋深度达到其株高的 133%时 ,可使羊柴幼苗全部死亡 ;沙埋深度分别为株高的 33%和 6 7%时 ,羊柴幼苗的整株生物量、叶片生物量、根系生物量以及相对生长率都相应地高于非沙埋的对照 (即 :0 %沙埋 )。与非沙埋的对照相比 ,羊柴幼苗在 33%和 6 7%沙埋条件下并不显著地改变其生物量分配格局 ;并且羊柴幼苗的叶数和株高与对照无明显差异 ,但试验期内的新生叶数是沙埋处理 (即 :33%、6 7%和 10 0 %沙埋 )的幼苗明显地高于非沙埋的对照。     

Effects of dung and seed size on secondary dispersal,seed predation,and seedling establishment of rain forest trees

Seeds dispersed by tropical, arboreal mammals are usually deposited singly and without dung or in clumps of fecal material. After dispersal through defecation by mammals, most seeds are secondarily dispersed by dung beetles or consumed by rodents. These post-dispersal, plant-animal interactions are likely to interact themselves, as seeds buried by dung beetles are less likely to be found by rodents than unburied seeds. In a series of three experiments with seeds of 15 species in central Amazonia (Brazil), we determined (1) how presence and amount of dung associated with seeds influences long-term seed fate and seedling establishment, (2) how deeply dung beetles bury seeds and how burial depth affects seedling establishment, and (3) how seed size affects the interaction between seeds, dung beetles, and rodents. Our overall goal was to understand how post-dispersal plant-animal interactions determine the link between primary seed dispersal and seedling establishment. On average, 43% of seeds surrounded by dung were buried by dung beetles, compared to 0% of seeds not surrounded by dung (n=2,156). Seeds in dung, however, tended to be more prone than bare seeds to predation by rodents. Of seeds in dung, probability of burial was negatively related to seed size and positively related to amount of dung. Burial of seeds decreased the probability of seed predation by rodents three-fold, and increased the probability of seedling establishment two-fold. Mean burial depth was 4 cm (0.5–20 cm) and was not related to seed size, contrary to previous studies. Probability of seedling establishment was negatively correlated with burial depth and not related to seed size at 5 or 10 cm depths. These results illustrate a complex web of interactions among dung beetles, rodents, and dispersed seeds. These interactions affect the probability of seedling establishment and are themselves strongly tied to how seeds are deposited by primary dispersers. More generally, our results emphasize the importance of looking beyond a single type of plant-animal interaction (e.g., seed dispersal or seed predation) to incorporate potential effects of interacting interactions.     

Effects of sand burial on purple sandgrass (Triplasis purpurea): the significance of seed heteromorphism

The effects of sand burial on seedling emergence, growth, and reproduction of the annual dune grass Triplasis purpurea (Walt.) Chapm. were examined. This species matures heteromorphic seeds on panicles enclosed by leaf sheaths along tiller nodes in a characteristic pattern: seeds at the lower nodes are heaviest, whilst those at the upper nodes are lightest. Field excavation of seedlings revealed that seedlings can emerge from seeds buried to 4 cm.In one experiment in an incubator, seeds collected from the upper and lower tiller nodes of glasshouse-grown parental plants were buried in sand at depths of 0, 3, and 6 cm. As expected, the effects of seed depth on seedling growth were due to delayed emergence for seedlings from buried seeds. However, seedlings from the larger seeds from the lower nodes of parental plants had significantly greater dry mass than those from the lighter seeds from upper nodes. Because lower node seeds are the most likely to become buried by sand in nature, the seed heteromorphism may be adaptive in the coastal environment.A second experiment involved the burial of seedlings to 0, 50, or 100% of their height. Although most seedlings did not survive complete burial (100% height), for partially buried seedlings (50% height) there was high survival and a marked stimulation of growth and eventual reproduction compared to unburied seedlings. This stimulation could be linked to increased root growth for seedlings that had been buried. The ability to respond positively to sand accumulation may represent an adaptation to the dynamic dune environment.     

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

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

Seed Dispersal by Monkeys and the Fate of Dispersed Seeds in a Peruvian Rain Forest1

Primary seed dispersal by two species of monkeys and the effects of rodents and dung beetles on the fate of dispersed seeds are described for a rain forest in southeastern Perú. During the six-month study period (June–November 1992) spider monkeys (Ateles paniscus) dispersed the seeds of 71 plant species, whereas howler monkeys (Alouatta seniculus) dispersed seeds of 14 species. Spider and howler monkeys also differed greatly in their ranging behavior and defecation patterns, and as a consequence, produced different seed rain patterns. Monkey defecations were visited by 27 species of dung beetles (Scarabaeidae). Dung beetles buried 41 percent of the seeds in the dung, but the number of seeds buried varied greatly, according to seed size. Removal rates of unburied seeds by rodents varied between 63–97 percent after 30 d for 8 plant species. The presence of fecal material increased the percentage of seeds removed by seed predators, but this effect became insignificant with time. Although seed predators found some seeds buried in dung balls (mimicking burial by dung beetles), depth of burial significantly affected the fate of these seeds. Less than 35 percent of Brosimum lactescens seeds buried inside dung balls at a depth of 1 cm remained undiscovered by rodents, whereas at least 75 percent of the seeds escaped rodent detection at a depth of 3 cm and 96 percent escaped at 5 cm. Both dung beetles and rodents greatly affected the fate of seeds dispersed by monkeys. It is thus important to consider postdispersal factors affecting the fate of seeds when assessing the effectiveness of frugivores as seed dispersers.     

Restoration Ecology of an Endangered Plant Species: Establishment of New Populations of Cirsium pitcheri

We determined the effects of shade, burial by sand, simulated herbivory, and fertilizers on the survival and growth of artificially planted population of Cirsium pitcheri—an endangered plant species of the sand dunes along Lake Huron. Sand burial experiments showed that greenhouse grown plants should optimally be transplanted into areas receiving 5 cm of sand deposition: burial at this depth maximized emergence, survivorship, and below‐ground biomass. Under field conditions, simulated herbivory of up to 50% of the plant height produced a slight increase in biomass after one year of growth. Field observations showed that when white‐tailed deer removed more than 50% of the transplant's leaf tissue, the plant died. The application of a 20:20:20 (N:P:K) water‐soluble fertilizer produced a significant increase in the dry leaf biomass, total leaf area, and total dry biomass relative to control plants. We also tested for the presence or absence of a persistent seed bank. Few seeds were recovered from soil samples collected from Pinery Provincial Park and Providence Bay. However, C. pitcheri has the ability to form a persistent seed bank under field conditions but only at soil depths of 15 cm. Cirsium pitcheri seeds are able to germinate and seedlings can emerge from a burial depth of up to 6 cm. Thus, seeds planted in open, sunny areas will probably maximize emergence, growth, and survivorship of seedlings. Populations of C. pitcheri can be restored by planting seeds at shallow depths, transplanting greenhouse‐grown plants, applying water soluble fertilizers, and protecting plants from herbivores.     

啮齿类取食的物种偏好与时空格局

通过强烈消耗土壤种子库,动物取食种子对植物种群更新和群落动态产生深远的影响。一般认为种子被食概率的空间格局取决于种子密度和离母树的距离,而环境（如地形）异质性的影响则一直没有得到足够的关注,与此相关的机制及其影响程度亦不清楚。研究设计在野外埋放种子以模拟种子扩散后的情形,监测啮齿类对种子的取食,以检验种子取食受埋藏生境、时间及动物对种子种类的偏好等因素的影响。结果表明,经过1a实验,8种落叶阔叶树种子的累计取食概率为0～48．25％,平均值为20％;山顶部位的取食概率大致是其它部位概率的3倍;埋放在凋落物层中的种子被食概率大约为埋放在土壤层中概率的2倍。利用logistic回归模型进行统计分析表明,种子被食概率变化的45％可以被上述因素解释。其中,物种偏好是影响种子被取食概率的首要因素,其后依次是地形、埋藏时间和深度。啮齿类明显喜好较大的种子;其取食行为在山脊部位明显较其它部位更频繁和剧烈;对埋藏种子的取食从3月份开始加剧,到7月份以后平息下来。种子埋放深度对啮齿类的取食概率有显著影响。     

Vertical structure of seed banks and the impact of depth of burial on recruitment in two temporary marshes

The structure of the seed bank (including Chara oospores), in relation to depth within the sediment and disturbance, was studied in two Rhône delta temporary marshes for two years. The seeds of all species were concentrated in the top 2 cm of sediment with very low numbers beeing found below 4 cm. When an exclosure eliminated disturbances of the sediment by animals, the vertical repartition of seeds at site 2 was more pronounced than outside the exclosure.In experiment 1, the emergence capacity of seeds from different depths and buried under layers of sterile equivalent to those in the field was measured. Depending of the species, 22 to 98% of the seeds germinated from unburied seeds in the top 2 cm. Only 1% of the oospores of Chara (from site 2) at 2 to 4 cm depth in the sediment emerged.In experiment 2, surface seed bank samples were placed under 0, 2 or 4 cm sterile sediment depth. The samples contained numerous recent seeds and the emergence percentage reached 41% (for Ruppia maritima). Only the seeds of Zannichellia spp failed to germinate from a depth of 2 cm or more. The emergence percentage from 2 cm depth or more was always lower than at the surface. These experiments showed that both burial and ageing of seeds decrease germination capacity.The majority of the active seeds located at the surface germinate when the marsh is flooded. Seeds located between 2 and 4 cm can be brought back to the surface by disturbances and play the role of a reserve involved in maintenance of populations that go without seed production for one or some years.     

Germination responses ofRubus palmatus var.coptophyllus andRubus parvifolius seeds with different burial durations to a variable light and temperature regime

To clarify the adaptive significance of seed dormancy, the effects of burial duration were examined for two deciduousRubus species:Rubus palmatus var.coptophyllus andRubus parvifolius, which are found mainly in relatively stable, shaded sites and disturbed sites, respectively. In early summer, newly ripened seeds were buried under litter on the soil surface in a pine forest, and germination tests were carried out for the seeds retrieved from the soil litter after 0 (not buried), 1, 2, 3, 5 and 8 or 9 months of burial. In general, the germination percentages increased and light requirements for germination decreased with increased burial duration. The percentage of seeds germinated with alternating temperatures in darkness also increased with increasing burial duration for both species. After 8 or 9 months of burial (corresponding to the next germination season in the field), the percentage of non-dormant seeds (including germination under alternating temperatures in the dark) was about 80% and 40% forR. palmatus var.coptophyllus andR. parvifolius, respectively. These seed dormancy traits of the twoRubus species may explain the differences in germination strategy in their habitats:R. palmatus var.coptophyllus seems to have adapted to the seasonal occurrence of favorable growing conditions after the dormancy breakage, whileR. parvifolius seems to have adapted to favorable conditions created by temporally unpredictable disturbances.     

Burial disturbance leads to facilitation among coastal dune plants

There is growing evidence that interactions among plants can be facilitative as well as competitive, but knowledge of how disturbances influence these interactions and how they vary with species diversity is lacking. We manipulated plant density, species diversity (richness), and a burial disturbance in a controlled, complete factorial experiment to test theories about the relationships among species interactions, disturbance, and richness. The hypotheses tested were 1) burial disturbance reduces plant performance at all levels of density and richness, 2) burial disturbance can cause net plant interactions to become more facilitative, and 3) facilitation increases with species richness. Burial decreased plant survival by 60% and biomass by 50%, supporting the hypothesis that burial reduces plant performance. In the control (unburied) treatment, there was no difference in proportion survival or per plant biomass between low and high density plots, meaning that neither competition nor facilitation was detected. In the buried treatment, however, high density plots had significantly greater survival and greater per plant biomass than the low density plots, indicating net facilitative interactions. Thus facilitation occurred in the buried treatment and not in the unburied control plots, supporting the hypothesis that facilitation increases with increasing disturbance severity. The hypothesis that facilitation increases with increasing species richness was not supported. Richness did not affect survival or biomass, and there was no richness by burial treatment interaction, indicating that richness did not influence the response of the community to burial. The influence of the disturbance on plant interactions was thus consistent across levels of richness, increasing the generality of the relationship between disturbance and facilitation.     

Litter and seed burying alter food availability and foraging efficiency of granivorous birds in the Monte desert

We assessed experimentally if the main granivorous bird species that feed on the ground in the central Monte desert are able to detect and consume seeds buried in the soil or trapped within litter. Understanding seed vulnerability to birds allows 1) a better understanding of how seed abundance translates into seed availability, a necessary step to assess seed limitation scenarios, and 2) whether birds alter the distribution of soil seeds through their consumption. Rufous‐collared sparrows found and consumed high proportions of buried seeds, though less seeds were eaten at increasing depths. In contrast, many‐colored chaco‐finches, common diuca‐finches and cinnamon warbling‐finches did not find buried seeds. All bird species fed on every substrate offered but, as a whole, birds reduced by 50% their seed consumption in Prosopis litter, and by 30% in Larrea litter, compared to consumption in bare soil. This effect was less notable for rufous‐collared sparrows, whose ‘double scratch’ foraging method would contribute to its great diet breath and abundance in the Monte desert. As birds do not reach a fraction of seeds buried and trapped by litter, seeds readily available for them may be scarcer than previously estimated through soil seed bank studies. Furthermore, since the four bird species detect and consume seeds from littered microhabitats, seed consumption by them surely affects the seasonal dynamics of the soil seed bank in all microhabitat types of the Monte desert.     

The viability and germination characteristics of exhumed Solanum mauritianum seeds buried for different periods of time

The site, depth and duration of burial significantly influenced the viability and state of dormancy of Solanum mauritianum seeds. Burial at a depth of 15 cm was most effective in reducing the level of conditional dormancy. Secondary dormancy was not induced at any of the environmental (burial) sites when seeds were maintained at 15 cm, where light and temperature fluctuations were minimal. When buried at 4 cm or maintained on the soil surface secondary dormancy was induced, particularly at the inland sites where environmental conditions such as temperature and moisture were more extreme. Conditional dormancy could generally be overcome by incubating seeds at 15/30 °C in the light, even after prolonged burial at unfavourable germination conditions. Gibberellic acid (500 mg l^–1) was very effective in breaking secondary dormancy of seeds induced by storage under unfavourable conditions after burial. These results have important implications for the control of this week in commercial forests.     

The effects of foraging by the superb lyrebird (Menura novae-hollandiae) in Eucalyptus regnans forests at Beenak,Victoria

Abstract In an early spar-stage stand of Eucalyptus regnans at Beenak, Victoria, foraging by lyrebirds in bare floor areas on steep slopes results in a complex microtopography of excavations, accumulations and terracettes. About 200 t ha^?1 of litter and top soil may be displaced an average of 70 cm downhill per year. Magnetic ferruginous pisolite was used as a marker to monitor progressive soil movement over 3 years. Very little disturbance occurred in areas of dense ground fern, but in bare areas the whole forest floor may be turned over every 20 months. In the site studied, foraging activity by lyrebirds varied seasonally and topographically. Disturbance by other biotic agents was minimal. The mean depth of soil cultivation was about 10 cm and litter was frequently buried or mixed intimately with soil. Since buried leaf litter decays more quickly than that on the surface, lyrebird foraging is likely to increase the rate of nutrient cycling. The small, steep clifflets left at the uphill limits of each scratch microsite provide litter-free niches for the establishment of tree fern prothalli and shade-tolerant herbs. All stages in the growth of the rough tree fern, Cyathea australis, were present in bare floor areas, but in dense ground fern patches, young stages were confined to rotten logs and upturned root balls. Since dense tree fern development tends to diminish the cover of dense ground fern, lyrebird foraging activity may maintain an accessible food resource which would otherwise diminish with increased ground fern cover in these forests in the course of secondary succession after fire.     

Direct seeding of dry forest tree species in abandoned pastures: effects of grass canopy and seed burial on germination

Natural tree regeneration in abandoned pastures can be hampered by various biotic and abiotic filters, including seed removal and predation. We tested the effects of maintenance and removal of grass and seed deposition (buried and unburied) on seed germination of 12 tree species in dry forest pastures. We obtained evidence supporting the hypothesis that seeds attain higher germination under a grass canopy than on bare ground. For most species, grass cover provides safety from seed predators and facilitates germination by providing a suitable microclimate with soil humidity similar to the forest. The hypothesis that buried seeds attain higher germination was not supported by our data. Predation and removal of unburied seeds ranged from 0 to 77 % and, alone or together, were the major causes of non-germination. Direct seeding is a promising technique for revegetation of recently abandoned pastures in areas originally covered by tropical dry forests. The high germination rate of seeds deposited on the ground and under grass reduces costs during initial restoration stages, potentially facilitating the spread and use of this technique.