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 regeneration of pioneer tree species under browsing pressure of Sika deer in an evergreen oak forest

The effects of Sika deer (Cervus nippon) browsing on the regeneration of pioneer species were studied in relation to canopy gaps in a warm temperate evergreen oak forest in Kasugayama, Nara City. Four study sites, three in canopy gaps and one under a closed canopy, were selected and each divided into fenced and unfenced plots. Under the closed canopy, seedlings of all the pioneer species died irrespective of browsing pressure. However, in the canopy gap sites, seedlings of the pioneer species could establish and grow well. The seedling survival ratio in the fenced plots in the canopy gaps was >60% 1 year after germination. However, in the unfenced plots, only <20% of the seedlings survived 1 year, with all dying within 3 years after germination. Thus, the regeneration of pioneer trees in this forest was strongly inhibited by deer browsing. Successful regeneration of a pioneer,Zanthoxylum ailanthoides, occurred for several years even after two major wind disturbances during the past 90 years. This may be due to less browsing pressure from the deer.     

Survival and growth of Virola surinamensis yearlings: Water augmentation in gap and understory

Summary Factors affecting seedling Virola surinamensis (Myristicaceae) survival and growth were investigated on Barro Colorado Island, Panama. Seedlings planted 3 months after germination were monitored in treefall gaps and understory using 2.25 ha irrigated and control plots through the first dry season. During the dry season, irrigated plants in gaps increased total leaf area significantly more than did irrigated plants in the shaded understory. Over the same dry season, control plants in gaps and in the shaded understory lost similar amounts of leaf area. Seedlings in understory were suppressed in stem height and biomass in both irrigated and control plots; these measures were greater in gaps and greatest in irrigated gaps (height). Roots were similar in length in all treatments, but greater in biomass in gaps than understory due to greater proliferation of secondary roots in control and irrigated gaps than in control and irrigated understory. This experiment demonstrates both water and light limitation during the first dry season after germination. V. surinamensis seedlings are capable of survival and modest growth of leaf area in the deep shade of the understory in moist locations; they are severely disadvantaged in shaded understory subject to drought, where most seeds fall and most seedlings establish. The broken canopy of a gap allows shoot and consequently root growth that permits seedlings to survive seasonal drought.     

An interactive effect of simultaneous death of dwarf bamboo, canopy gap, and predatory rodents on beech regeneration

To clarify the interactive effect of the simultaneous death of dwarf bamboo (Sasa kurilensis), forest canopy gap formation, and seed predators on beech (Fagus crenata) regeneration, we analyzed beech demography from seed fall until the end of the first growing season of seedlings in an old-growth forest near Lake Towada, northern Japan. The simultaneous death of S. kurilensis took place in 1995. We established four types of sampling site differing in forest canopy conditions (closed or gap) and Sasa status (dead or alive). Beech seed survival and emergence ratio were both highest in gaps with dead Sasa (gap-dead), because rate of predation was lowest. Seedling survival during the first growing season was also highest in the gap-dead treatment, because of less predation and less damping off. As a result, even though density of seed fall was lowest in the gap-dead treatment, the living seedling density there was highest at the end of the first growing season. Predation, which caused the greatest mortality during the seed and seedling stages, was significantly lower at both sites in gaps and sites with dead Sasa. This was probably due to changes in the behavior of rodents in response to the structure of the forest canopy and undergrowth. Both the death of Sasa and canopy gap formation allowed seedlings to avoid damping off because of the high light availability. The indirect effect of the simultaneous death of Sasa and canopy gap formation in reducing predation contributed more to beech regeneration than their direct effect in increasing light for the seedlings.     

Short‐term changes affecting regeneration of Fagus crenata after the simultaneous death of Sasa kurilensis

Abstract. Question: The aim of the present study is to determine whether seed/seedling predation will increase and Fagus survival will decline with the recovery of the Sasa cover. Methods: We examined Fagus crenata regeneration for seven years in an old‐growth Fagus‐Sasa forest near Lake Towada, northern Japan, by examining the effects of simultaneous death of Sasa, tree canopy gap formation, mast seeding of Fagus and seed and seedling predation by rodents on the survival of Fagus seeds and current year seedlings. We established four types of sites differing in forest canopy (closed or gap) and Sasa status (dead or alive) following the simultaneous flowering and death of Sasa kurilensis (dwarf bamboo) in 1995. Results: Fallen Fagus seed was abundant in 1997 and 2000 (mast years). In sites with alive Sasa, survival from the first growing season was low due to high seed and seedling predation. In sites with dead Sasa, seed survival under the canopy was high for both mast years, but in gaps it varied between years. Seedling survival was highest in canopy gaps with dead Sasa (gap‐dead) in 1998, because of higher light levels and lower predation by rodents. However, seedling survival in these plots was low in 2001, apparently because rapid Sasa recovery favoured rodent predation. In both mast years, Sasa die‐back had significant positive effects on seed and seedling survival under closed canopies because the seedlings there were more successful in escaping predation. Conclusion: The change in successful sites for the early stage of regeneration of Fagus appears to reflect the combined effects of canopy gap, seed/seedling predation and revegetation of Sasa.     

Functional anatomy of five endangered tropical timber wood species of the family Dipterocarpaceae

Wood anatomy of five dipterocarp species endemic to the Philippines was studied with the goal to explore functional wood traits of ecological significance. Stem wood of 6-year-old trees grown under similar environmental conditions in a plantation (Leyte, Philippines) was used. Wood densities decreased in the following order Hopea plagata > Dipterocarpus kerrii > Parashorea malaanoman > Shorea almon ≈ Shorea contorta. This was mainly caused by significantly thicker fiber cell walls of H. plagata and D. kerrii than those of the other three species. Wood density was negatively correlated with the abundance of axial parenchyma cells. Predicted conductance was independent from wood density and lowest in H. plataga and highest in D. kerrii and S. contorta. These results indicate that H. plagata and D. kerrii is woods have higher constructions costs in term of carbon per unit of biomass, and that H. plagata is probably better acclimated to varying soil moisture than the other species.     

Fire behaviour and the invasion of Acacia sieberiana into savanna grassland openings

Following the recent decline in browsing and grazing pressures and changed fire regimes in Queen Elizabeth National Park, Uganda, Acacia thickets have encroached on grassland habitats important for grazing mammals. The objective of this research was to test experimentally the effects of fire behaviour, using simulated and natural fuel loading conditions, on A. sieberiana seedling and sapling regeneration. A high fire intensity (3200 kW m^-1) in natural fuels stimulated high seedling emergence (172 seedlings m^-2) compared to 6 seedlings m^-2 without fire. Also a highly significant linear regression was established between percentage top-kill of seedlings and fire intensities. High fire intensities during late dry season fires were more effective in controlling sapling height growth than early dry season fires of low intensities. A conceptual diagram was developed to show the major factors and possible pathways leading to successful invasion by A. sieberiana into grassland openings of the savanna.     

林窗对川西周公山柳杉人工林林下物种多样性的影响

采用典型样地法,以川西周公山柳杉人工林5种不同大小的林窗为研究对象,以林下非林窗为对照,研究了不同大小的林窗对柳杉人工林物种多样性的影响,同时分析了不同梯度林窗下林窗中心、林窗边缘、及林下群落的物种组成、物种多样性的变化情况。结果表明:(1)在所调查的18个样地231个样方中共记录到维管束植物141种,隶属于76科113属;随着林窗面积的增大,群落各层次的物种数呈现出先升高后降低的趋势,灌木层物种数在各林窗梯度上表现为林缘林下林窗中心,草本层物种数在各林窗梯度上表现为林缘林窗中心林下。(2)不同林窗优势种及其重要值不同,即在小林窗内,优势种为柳杉和野桐,其重要值之和高达0.292 3;在大林窗内,杉木及亮叶桦为群落优势物种,群落内出现大量其更新幼苗。(3)不同大小的林窗表现为灌木层物种丰富度指数(D)、Shannon-Wienner指数(H)、和Pielou均匀度指数(Jsw)值在400~450 m2面积的大林窗内达到一个均优水平,草本层物种的多样性在面积为100~150m2的小林窗内达到较高水平;不同梯度的林窗各层次群落D、H值整体表现为林缘林窗中心林下。研究认为:林窗的存在会改变群落物种组成,提高群落物种多样性水平,并且大林窗(400~450m2)更利于柳杉人工林林下树种更新及物种多样性的提高。     

Factors limiting early establishment of the Mediterranean grassland species Brachypodium retusum at disturbed sites

In Mediterranean dry grasslands, water availability and grazing strongly influence plant establishment. Although higher water availability and lower grazing pressure usually increase seedling recruitment in the beginning, higher competition may reduce seedling survival. In our study, we analysed this trade-off for the dominant species of Western Mediterranean steppes Brachypodium retusum which hardly recolonises restoration sites degraded by soil disturbance such as arable use.A split-plot experiment was set up on an ex-arable site in the “La Crau” steppe in order to test the effects of initial watering and sheep grazing on seedling recruitment, survival and early growth during two years. The watering treatment was applied in the first autumn and spring doubling the usual precipitation during this period and grazing was tested using exclosures. Survival and growth of seedlings established from field-sown seeds were compared to those of transplanted seedlings pre-grown in a growth chamber.Watering did not affect germination whereas the grazing effect was negative. Initial watering had a positive effect on survival only in grazed plots suggesting a compensatory effect under grazing disturbance. Significant main effects of watering on early growth disappeared in the second season. Grazing significantly reduced most measured growth traits whereas the effect on survival was only marginally significant. Planted seedlings (49%) survived better than seedlings emerged from field-sown seeds (36%). In conclusion, initial fencing increases establishment success of B. retusum in ecological restoration. Initial watering may help to mitigate negative grazing effects when fencing is not possible.     

Legacy microsite effect on the survival of bitterbrush outplantings after prescribed fire: capitalizing on spatial variability to improve restoration

Restoration of shrubs in arid and semi‐arid rangelands is hampered by low success rates. Planting shrub seedlings is a method used to improve success in these rangelands; however, it is expensive and labor intensive. The efficiency of shrub restoration could be improved by identifying microsites where shrub survival is greater. Bitterbrush (Purshia tridentata Pursh DC) is an important shrub to wildlife that has declined because of conifer encroachment, excessive defoliation, wildfires, and low recruitment. We investigated planting bitterbrush seedlings in western juniper (Juniperus occidentalis ssp. occidentalis Hook) encroached shrublands after prescribed fire was used to control trees. Bitterbrush seedlings were planted in under (canopy) and between (interspace) former juniper canopies at five blocks and evaluated for three growing seasons. Bitterbrush survival was greater than 50% in the former canopy, but only 5% in the interspace microsite by the third growing season. Growth of bitterbrush was also greater in the former canopy compared with the interspace, potentially due to markedly less perennial vegetation in this microsite. Exotic annual grasses and annual forbs became prevalent in the former canopy in the second and third growing season, suggesting that soil resource availability was greater in this microsite. These results suggest that restoration success will vary by specific locations within a burned landscape and that this variability can be used to improve restoration efficiency. In this situation, bitterbrush restoration can be improved by planting seedlings in former canopy compared with interspace microsites.     

Abundance of arbuscular mycorrhizal fungi spores in different environments in a tropical rain forest, Veracruz, Mexico

 The aim of this study was to compare mycorrhizal abundance and diversity in sites with different regimes of disturbance in a tropical rain forest at Los Tuxtlas, Veracruz, Mexico. Arbuscular mycorrhizal spores were quantified at two sites: closed canopy and gaps in the forest. Data were recorded during dry, rainy, and windy ("nortes") seasons. Spores of eight Glomus species, sporocarps of three Sclerocystis species, three species of Acaulospora and two of Gigaspora were found. Significant differences in the number of species and spores were found among seasons. The highest numbers of species and spores were observed during the dry season, with a marked decrease during the rainy season. Our results show that disturbance does not but seasonality does affect abundance and richness of mycorrhizal spores in this tropical wet forest. Accepted: 11 October 1998     

Relationships between intra‐specific variation in seed size and recruitment in four species in two contrasting habitats

Large seeds contain more stored resources, and seedlings germinating from large seeds generally cope better with environmental stresses such as shading, competition and thick litter layers, than seedlings germinating from small seeds. A pattern with small‐seeded species being associated with open habitats and large‐seeded species being associated with closed (shaded) habitats has been suggested and supported by comparative studies. However, few studies have assessed the intra‐specific relationship between seed size and recruitment, comparing plant communities differing in canopy cover. Here, seeds from four plant species commonly occurring in ecotones between open and closed habitats (Convallaria majalis, Frangula alnus, Prunus padus and Prunus spinosa) were weighed and sown individually (3200 seeds per species) in open and closed‐canopy sites, and seedling emergence and survival recorded over 3 years. Our results show a generally positive, albeit weak, relationship between seed size and recruitment. In only one of the species, C. majalis, was there an association between closed canopy habitat and a positive seed size effect on recruitment. We conclude that there is a weak selection gradient favouring larger seeds, but that this selection gradient is not clearly related to habitat.     

Exposure to herbivores increases seedling growth and survival of American chestnut (Castanea dentata) through decreased interspecific competition in canopy gaps

The successful development of early stages of blight‐resistant hybrid stock has increased hopes for restoration of American chestnut (Castanea dentata) to eastern North American forests. However, these forests have undergone substantial ecological change in the century since the functional extirpation of American chestnut, and it remains unknown to what extent American chestnut will be able to recolonize contemporary forests. In particular, high densities of white‐tailed deer (Odocoileus virginianus) and competition with mesophytic tree species such as maple (Acer) may impede chestnut regeneration, much as they affect oak (Quercus). We used a split‐plot analysis of variance (ANOVA) design to examine the effects of canopy gaps and herbivory on survival and growth of third generation backcrossed (BC₃) hybrid chestnut seedlings over two growing seasons in central Indiana, U.S.A. Only 4 of 588 (0.7%) seedlings in closed‐canopy plots survived to the end of the study, as opposed to 264 of 589 (45%) seedlings in gap plots. Within the gap treatment, fencing was associated with reduced chestnut survival as well as reduced herbivory and increased cover of non‐chestnut vegetation. Our results indicate that herbivory may indirectly benefit chestnut regeneration by suppressing competition. However, this beneficial effect is likely context‐dependent and additional work is needed to establish the conditions under which it occurs.     

Species Identity and Water Availability Determine Establishment Success Under the Canopy of Retama sphaerocarpa Shrubs in a Dry Environment

Seedling establishment in harsh environments is often enhanced by the proximity of adult shrubs. This information has been used in restoration work by placing seedlings of species being restored under the canopy of some shrubs. However, monitoring this process is often restricted to a single species, and comparisons with practices that protect planted seedlings against harsh conditions are scant. Similarly, few studies have supplied seedlings with water in the summer to observe the effects of water availability on the interaction. We compared sapling survival of three woody species (Olea europaea, Pistacia lentiscus, and Ziziphus lotus) under the leguminous shrub Retama sphaerocarpa and in gaps covered with piled branches that mimicked a shrub canopy. After 3 years, survival of saplings planted under Retama differed depending on species identity and water supply. Survival of Olea saplings placed under Retama shrubs was twice that under piled branches if not watered (35 ± 8 vs. 17 ± 2 %, respectively), whereas survival of saplings under Retama, if watered, was less (48 ± 11 vs. 68 ± 8%, respectively). Retama shrubs had a negative effect on Ziziphus; most saplings died under its canopy, whereas survival in piled branches ranged 10–54%. Pistacia was neither facilitated nor outcompeted by Retama. Facilitation of Olea by Retama shrubs was more apparent under dry conditions where watering increased competition and decreased facilitation. Overall, we conclude that Retama shrubs can help dry land restoration to a greater extent than artificial shade for Olea when not watered. The critical role played by water supply in determining nursing success rates warrants further study.     

Effect of Sowing Season,Plant Cover,and Climatic Variability on Seedling Emergence and Survival in Burned Austrocedrus chilensis Forests

Fire is the most important disturbance factor in Cypress (Austrocedrus chilensis) forests in Patagonia, Argentina. This ecosystem recovers poorly after fire, and direct sowing could be a potentially useful restoration practice. To evaluate the effect of season of sowing, post‐fire plant cover (PC), and climatic variability on seedling emergence and survival, three direct sowing studies were established in two burned cypress stands: Trevelin (xeric conditions) and El Bolsón (mesic conditions). Two studies were conducted in winter (2000 and 2001) and one in spring (2001). Precipitation was higher than the mean during the 2000–2001 growing season and lower during 2001–2002. At both sites, emergence and survival were much higher for winter‐ than for spring‐sown seedlings. In the xeric stand, emergence and survival of winter‐sown seedlings increased with medium and high PC values, after the humid and dry summers, respectively. However, most spring‐sown seedlings did not emerge, and those that did were short‐lived. Because of the more favorable growing conditions in the mesic stand, PC had no effect on emergence and only favored first year survival of winter‐sown seedlings after the dry summer. Spring‐sown seedlings showed no association with PC in the mesic site, probably because the first summer was exceptionally humid. We speculate that shading plants exert a positive effect on cypress seedling establishment, likely by reducing the stress from high temperatures and low water availability. Sowing of small patches under the protection of understory vegetation could be useful in restoring burned cypress stands.     

Effects of fire regimen on two species of epiphytic orchids in tropical savannas of the Northern Territory

The effect of four fire regimens (early dry season annual, late dry season annual, early dry season biennial, unburnt) on the numbers of epiphytic orchids in a savanna community in Kakadu National Park was investigated. These fire regimens had been maintained on I ha plots for 16 years. Two species of epiphytic orchid were present — Cymbidium canaliculatum and Dendrobium affine. The numbers of Cymbidium plants were insufficient to enable statistical analysis, but plants were present in the unburnt, early annual and early biennial fire treatments. Dendrobium occurred in substantial numbers irrespective of fire treatment, but the extent of colonization of all available host trees was greater in the unburnt than the burnt treatments. The lower numbers of Dendrobium plants in burnt treatments was due to a lower proportion of potential host trees being colonized and a lower number of orchids on each colonized tree. The proportion of small trees colonized was markedly lower in burnt treatments. Frequent fires late in the dry season may prevent recruitment of the main host species. Eucalyptus tectifica. The survival of orchids despite frequent fires depended on the availability of relatively protected sites for colonization and the ability of the orchids to withstand some fire damage.     

Improving restoration success through microsite selection: an example with planting sagebrush seedlings after wildfire

Post‐fire restoration of foundation plant species, particularly non‐sprouting shrubs, is critically needed in arid and semi‐arid rangeland, but is hampered by low success. Expensive and labor‐intensive methods, including planting seedlings, can improve restoration success. Prioritizing where these more intensive methods are applied may improve restoration efficiency. Shrubs in arid and semi‐arid environments can create resource islands under their canopies that may remain after fire. Seedlings planted post‐fire in former canopy and between canopies (interspace) may have different survival and growth. We compared planting Wyoming big sagebrush (Artemisia tridentata Nutt. ssp. wyomingensis Beetle & Young) seedlings post‐fire in former sagebrush canopy and interspace microsites at five locations. Four growing seasons after planting, seedling survival was 46 and 7% in canopy and interspace microsites, respectively. Sagebrush cover was 5.8 times greater in canopy compared to interspace microsites. Sagebrush survival and cover were likely greater because of less competition from herbaceous vegetation as well as benefiting from resource island effects in canopy microsites. Initially, post‐fire abundance of exotic annual grasses was less in canopy microsites, but by the third year post‐fire it was substantially greater in canopy microsites, indicating that resource availability to seedlings was greater, at least initially, in canopy microsites. These results suggest microsites with greater likelihood of success should be identified and then utilized to improve restoration success and efficiency. This is important as the need for restoration greatly exceeds resources available for restoration.     

Episodic seedling growth in Allosyncarpia ternata,a lignotuberous,monsoon rainforest tree in northern Australia

On the western Arnhem Land Plateau, Northern Territory, Australia, seedlings of the canopy tree Allosyncarpia ternata S.T. Blake typically spend many years (perhaps decades) as small (<1 m), multistemmed plants on the forest floor. In this establishment phase, long periods of apparent inactivity are interrupted by episodes of rapid growth. This paper describes a 5‐year field‐monitoring program to examine the pattern of seedling growth and survival in allosyncarpia forest, and field and shadehouse measurements of lignotuber size. Individual seedlings may produce, each wet season, a number of fast‐growing stems, which then die back in the following dry season. As a result, mean annual above‐ground growth during this life stage is negligible. With each wet season, however, the seedling extends its below ground parts – a large lignotuber and a deep root system. After a number of years, when the lignotuber has grown large enough to sustain massive shoot growth, when a suitable light gap becomes available, and presumably when roots reach reliable dry‐season water supplies, the seedling grows rapidly. Thus, the shortage of saplings in allosyncarpia forest is due to the short time that individual plants spend at that particular growth‐stage, rather than to any dysfunction in recruitment.     

Potential Risk of Hybridization in Ex Situ Collections of Two Endangered Species of Sinojackia Hu （Styracaceae）

Spontaneous hybridization in ex situ facilities can undermine the genetic integrity of ex situ collections and potentially contaminate open-pollinated seeds or seedlings destined for the reintroduction of endangered plant species into the wild. In the present study, the potential risk of hybridization between two endangered Chinese endemic species, namely Sinojackia xylocarpa Hu and S. rehderiana Hu, which are naturally allopatric species but were conserved ex situ in Wuhan Botanical Garden （WBG）, Wuhan, China, were investigated over three consecutive years from 2003 to 2005. The entire overlapping flowering period of the two species was 14-20 d and the two species shared the same pollinator insects during the entire flowering season in WBG. The floral isolation between the two species was not an issue in the ex sltu collection at WBG. The results suggest an opportunity for pollen transfer between species and a potential risk of genetic Introgression and loss of genetic identity of open-pollinated seeds produced in the ex sltu Collection of these two endangered species. An artificial reciprocal cross between S xylocarpa and S. rehderlana confirmed that the two congener species could readily set seeds, indicating no post-pollination barriers to hybridization and the importance of spatial isolation as a barrier to inter-specific crossing. Therefore, to manage these crossable species with overlapping flowering times and shared pollination vectors in ex situ facilities, spatial isolation should be carefully considered to minimize the possibility of spontaneous hybridization.     

The Fecundity,Seed, and Seedling Ecology of the Edible Palm Euterpe edulis in Southeastern Brazil1

Euterpe edulis flowers at the beginning of the wet season and fruits over an extended period during the end of the wet season and the dry season. Reproductive plants produced an average of ca 1500 fruits within a season. Five species of birds belonging to the family Turdidae were observed to feed on and disperse the fruits. Most seeds fell directly to the ground, where there was some subsequent local dispersal by water. The dispersion of seeds was highly clumped with most seeds being found in close proximity to adult plants; the distance of dispersal could be described by a negative power curve. Germination was higher on wetter soils. The probability of seedlings advancing to the next size class within a year was inversely related to the density of seedlings, and occurred optimally at ca 4 m from adult plants. The latter observation indicates that the survival and growth of seedlings are suppressed in the immediate vicinity of conspecific adults.