Natural regeneration of Pinus resinosa on burned and unburned sites in Newfoundland

Abstract. Natural regeneration of Pinus resinosa (red pine) seedlings around mature trees was studied in burned and unburned stands. Growth inhibitory effects of the forest organic matter on red pine seedlings was tested by a stair-step experiment using leachate of forest soil monoliths and also by a seed germination bio-assay using forest floor substrates. To test if higher burning temperatures can remove the allelopathic effects of red pine-Kalmia organic matter, a laboratory bio-assay was conducted by germinating red pine seeds on the organic matter burned at 200, 400, 600 and 800°C. Deposition of dry needles and a thick duff layer under red pine stands affected seedling establishment. Red pine seedling establishment increased with the decreasing thickness of duff layer away from the stump of the seed-bearing trees. Wildfire helped in removing the duff layer and increased seedling establishment. A high fuel load within a 0 - 1 m radius around the tree stump caused a deep burn of the organic matter including part of the soil seed reserve. On a burned-over surface, more seedlings established in a band between 1 and 2 m around the stump than inside and outside the band. Primary root growth of red pine was severely inhibited when the seedlings were grown in unburned forest floor organic matter where Kalmia was the principal understory species. Water leachate of a Pinus resinosa-Kalmia soil monolith was inhibitory to red pine seedling growth. In greenhouse conditions, the seedlings grew well in burned-over soil from a Pinus resinosa stand. Burned organic matter from a red pine forest showed an increase in pH with a burning temperature of 600°C. Primary root growth of red pine seedlings was similarly increased with increasing temperature up to 600°C; at higher temperatures the root length of seedlings did not increase any further.     

Species-specific Seedling Responses to Hurricane Disturbance in a Puerto Rican Rain Forest1

Seedling dynamics were followed in a Puerto Rican forest for 20 months following a severe hurricane to study the interactive effects of hurricane debris, nutrients, and light on seedling diversity, density, growth, and mortality. Three treatments (debris removal, an unaltered control with hurricane debris, and chemical fertilization added to hurricane debris) altered levels of forest debris and soil nutrients. Canopy openness was measured twice using hemispherical photographs of the canopy. We examined the demographic responses of six common species to treatments over time. Seedling densities increased for all six species but the only significant treatment effects were increased densities of the pioneer tree Cecropia and the shrub Palicourea in the debris removal treatment. Seedling growth declined with declining light levels for four species but not for the pioneer tree Alchornea or the non‐pioneer tree Dacryodes. Only Cecropia and the non‐pioneer tree Chionanthus had treatment effects on growth. Mortality also differed among species and tended to be highest in the fertilized plots for all but Cecropia and Dacryodes. We found only some of the expected differences between pioneer and non‐pioneer plants, as each species had a unique response to the patchy distributions of organic debris, nutrients, and light following the hurricane. High local species diversity was maintained through the individualistic responses of seedlings after a disturbance.     

Facilitation and competition influence succession in a subtropical old field

The effects of neighboring vegetation and soil fertility on the establishment, survival and growth of tree species were studied in a subtropical old-field area in south Brazil. Seed damage, germination and seedling establishment of four tree species plus growth and survival of two transplanted tree species were monitored under factorial combinations of the following treatments: (1) pioneer vegetation (presence and absence); (2) soil fertility (addition of NPK and control). Facilitation was the main process affecting plant performance. The presence of pioneer vegetation significantly improved germination, establishment, growth and survival of most study species. Around 90% of sown seeds were damaged and the removal of pioneer vegetation significantly increased seed damage for all species studied, decreased germination in three out of four species, and decreased establishment in one species. Moreover a significantly higher seedling growth rate of Inga virescens was found after the first year of the experiment in plots where vegetation was present. The presence of vegetation significantly increased seedling survival of I. virescens by protecting seedlings from leaf loss due to winter frosts. Competition was detected by the second year when a higher growth of transplanted seedlings of the species Araucaria angustifolia occurred in plots where vegetation was present and fertilizer were applied. A lower growth rate was detected in plots where vegetation was present but fertilizer was not applied. These results indicate a balance between competition for soil nutrients and protection by neighbor vegetation. Damage of seedlings by leaf cutter ants was an important barrier for plant survival. Damage occurred in 80% of the A. angustifolia seedlings and 58% of these damaged seedlings died. The presence of neighboring vegetation tended to protect seedlings from ant damage. Although competition occurred, facilitation seems to be the main process driving early successional changes in this subtropical old field. This was mainly due to the improvement of local microclimatic conditions and protection against herbivores by neighboring vegetation. Facilitation occurred during establishment and growth phases in a subtropical area that is considered a productive, low stress environment. Our results indicate that facilitation may be more frequent in productive environments than previously thought.     

Soil factors predict initial plant colonization on Puerto Rican landslides

Tropical storms are the principal cause of landslides in montane rainforests, such as the Luquillo Experimental Forest (LEF) of Puerto Rico. A storm in 2003 caused 30 new landslides in the LEF that we used to examine prior hypotheses that slope stability and organically enriched soils are prerequisites for plant colonization. We measured slope stability and litterfall 8–13 months following landslide formation. At 13 months we also measured microtopography, soil characteristics (organic matter, particle size, total nitrogen, and water-holding capacity), elevation, distance to forest edge, and canopy cover. When all landslides were analyzed together, plant biomass and cover at 13 months were not correlated with slope stability or organic matter, but instead with soil nitrogen, clay content, water-holding capacity, and elevation. When landslides were analyzed after separating by soil type, the distance from the forest edge and slope stability combined with soil factors (excluding organic matter) predicted initial plant colonization on volcaniclastic landslides, whereas on diorite landslides none of the measured characteristics affected initial plant colonization. The life forms of the colonizing plants reflected these differences in landslide soils, as trees, shrubs, and vines colonized high clay, high nitrogen, and low elevation volcaniclastic soils, whereas herbs were the dominant colonists on high sand, low nitrogen, and high elevation diorite soils. Therefore, the predictability of the initial stage of plant succession on LEF landslides is primarily determined by soil characteristics that are related to soil type.     

Seed predation,disease and germination on landslides in Neotropical lower montane wet forest

Abstract. Seed mortality (caused by predators and pathogens) and germination were compared between Puerto Rico and Costa Rica on landslides in lower montane wet forest. Seeds of six common species on five Puerto Rican landslides and four common species on two Costa Rican landslides were used with a Cecropia species and a Gonzalagunia species included at both sites. In the Puerto Rican experiments Cecropia schreberiana was the only species to show significant seed predation (which was due to insects), pathogens grew from all species and fewer seeds were lost to predators than pathogens. Also in Puerto Rico mean germination across all species was 57 % before dispersal (filled seeds collected while still on the tree) and 71 % after, with Phytolacca rivinoides seeds germinating most abundantly, followed in descending order by Ocotea leucoxylon, Cecropia spec, Miconia racemosa, Palicourea riparia and Gonzalagunia spicata. In the Costa Rican experiments three species had significant predation: Cecropia polyphlebia and Urera caracasana (both due to insects) and Witheringia coccoloboides (due to mammals); pathogenic disease caused more seed loss than predation, and germination was high (61 % pre-dispersal, 69 % post-dispersal). Similarities between these island and mainland sites included (1) percentage of seeds lost to predation and percentage lost to pathogens (all in the 5–15 % range), (2) generalist pathogens which claimed more seeds than predators and (3) majority germination with a general increase after dispersal. Finally sites were dissimilar only in the number of species with significant predation loss and whether it was by insects or mammals, casting doubt on the traditional island/mainland dichotomy.     

氮磷添加与不同栽植密度交互对樟树幼苗土壤化学性质的短期影响

研究氮磷添加对不同密度樟树(Cinnamomum camphora)幼苗土壤化学性质的影响,以期为全球化背景下樟树人工林生态系统的土壤养分管理提供依据。以1年生樟树幼苗为试验材料,选择氯化铵(NH4Cl)作为氮肥模拟大气氮沉降,以二水合磷酸二氢钠(NaH_2PO_4·2H_2O)模拟磷添加。氮磷处理设置CK、施N、施P和施N+P 4个水平,其中N、P和N+P施肥量分别为40 g m~(-2)a~(-1)(NH_4Cl)、20 g m-2a-1(NaH_2PO_4·2H_2O)和40g m~(-2)a~(-1)(NH_4Cl)+20 g m~(-2)a~(-1)(NaH_2PO_4·2H_2O)。种植密度设置4个水平:10、20、40和80株/m~2,试验时间为2017年6月至9月。研究结果表明,在各密度幼苗土壤中,N和N+P处理引起pH值的显著下降,N、P和N+P处理的土壤有机质和碱解N含量的变化规律不明显,P处理的幼苗土壤全P含量上升,P和N+P处理的土壤有效P含量增加,N+P处理的土壤全K含量以及N、P和N+P处理的土壤速效K含量均下降。在10、20和40株/m~2幼苗的土壤中,P处理的土壤全N含量高于N和N+P处理的,而80株/m~2幼苗的土壤全N含量低于其他密度幼苗。随着种植密度的增加,各施肥处理的土壤pH、全P、有效P、全K和速效K含量均呈现上升趋势,而施N和施P处理的土壤有机质呈现下降趋势,各施肥处理的土壤碱解N含量变化规律不明显。施肥和密度处理对樟树幼苗土壤有机质、碱解氮和速效钾含量有显著的交互作用。     

深挖垦复对锥栗重要农艺性状及土壤理化性质的影响

为探索锥栗林地高效的土壤管理方式,提升我国锥栗栽培管理技术水平,进而提高其产量、品质以及经济效益。该研究采用全垦和环垦两种方式对锥栗林地进行连续4年的深挖垦复,通过测定垦复前后土壤理化性质变化以及锥栗树体生长、叶片表型和生理特征、结果性状、产量及品质等重要农艺性状,统计数据并进行对比分析。结果表明:(1)深挖垦复对锥栗林地土壤理化性质改善效果显著,两种垦复方式土壤容重较垦复前降低31.21%及以上(0～30 cm处),土壤含水率、土壤孔隙度、有机质含量以及各种大量元素含量较垦复前和对照均有不同程度的增加,土壤肥力及其保水保肥能力显著增强。(2)环垦区土壤有机质含量、有效磷含量以及交换性镁含量高于全垦区,其土壤有机质含量较垦复前增加40.59%,远高于全垦增加幅度(17.76%),从土壤保肥能力的角度来看,环垦效果优于全垦。(3)土壤肥力的提升增强了其对锥栗叶片的供肥能力,使得叶片含水率、叶绿素含量以及各种矿质元素含量显著增加,从而提升其光合作用能力。(4)深挖垦复对锥栗树体生长、结实能力、产量及品质同样具有显著的提升效果,其中全垦和环垦区单位面积产量分别是对照的1.75倍和1.33倍,且栗苞总重、单果质量、出籽率、可溶性糖含量以及磷、钾元素含量显著高于对照,而空苞率显著低于对照。综上,深挖垦复是改良林地土壤和提高锥栗生产力的有效举措。     

Effects of light,moisture, temperature,and litter on the regeneration of five tree species in the tropical montane wet forest of Puerto Rico

Field experiments were conducted to investigate the effects of light, moisture, temperature, and litter on the regeneration of two early-, one mid-, and two late-successional tropical tree species. High light and litter seem to be universally good cues for regeneration, increasing seed/seedling survival for all species except for Cecropia (an early-successional species) whose small seeds may not be able to penetrate the litter layer. In addition, the high temperature environment in both artificially shaded and nonshaded areas of a natural gap exhibits less seed loss, an increase in the percent and rate of germination, and an increase in seedling survival for Dacryodes (a late-successional species), than the lower temperature environment under an intact canopy. Low soil water is also a good cue for Dacryodes germination as it is for Prestoea and Cecropia. Finally, the lower temperature environment found under the forest canopy (compared to the natural gap) leads to less seed loss and more germination for Guarea (a mid-successional species). Our results suggest that a good patch for regeneration of many species in this forest, early- as well as late-successional species, would have high light and a litter layer that moderates temperature and moisture extremes. The substantial variation in suitability among regeneration filters and species could: (1) contribute to low establishment success, i.e., most dispersed propagules do not become trees, (2) make it difficult to group species into germination strategies, and (3) make it hard to generalize about a net effect of any specific environmental variable on establishment. We suggest that tropical disturbances should be viewed in terms of their impact on a variety of environmental cues, which may signal germination and impact subsequent growth and survival.     

Effects of climate warming on plant autotoxicity in forest evolution: a case simulation analysis for Picea schrenkiana regeneration

In order to explore how plant autotoxicity changes with climate warming, the autotoxicity of P. schrenkiana needles' water extract, organic extract fractions, and key allelochemical DHAP was systemically investigated at the temperature rising 2 and 4°C based on the data‐monitored soil temperature during the last decade in the stage of Schrenk spruce regeneration (seed germination and seedling growth). The results showed that the criterion day and night temperatures were 12°C and 4°C for seed germination, and 14°C and 6°C for seedling growth, respectively. In the presence of water extract, the temperature rise of 2°C significantly inhibited the germination vigor and rate of P. Schrenkiana seed, and a temperature rise of 4°C significantly increased the inhibition to the seedling growth (P < 0.05). Among the three organic fractions, the low‐polar fraction showed to be more phytotoxic than the other two fractions, causing significant inhibitory effects on the seed germination and growth even at low concentration of 0.1 mg/mL, and the inhibition effect was enhanced as temperature increased. The temperature rise significantly enhanced the promotion effect of DHAP, while the inhibition effect of temperature rise became less important with increasing concentration of DHAP. This investigation revealed that autotoxicity of P. schrenkiana was affected by the climate warming. As expected, it provided an insight into the mechanism and effectiveness of allelopathy in bridging the causal relationship between forest evolution and climate warming.     

凋落物物理阻隔对格氏栲种子萌发及胚根生长的影响

为了探讨凋落物物理阻隔对格氏栲天然林自然更新状况的影响,通过模拟野外凋落物覆盖,设置格氏栲种子上层覆盖厚0 cm(CK)、2 cm(D2)、4 cm(D4)、6 cm(D6)、8 cm(D8)及种子下层铺垫厚2 cm(U2)、4 cm(U4)凋落物等7个处理,分析凋落物覆盖方式及厚度对格氏栲种子萌发及胚根生长的影响。结果表明:(1)凋落物覆盖方式及厚度对种子萌发进程存在显著影响。CK萌发持续时间最长,上层覆盖处理(D)次之,下层铺垫处理(U)的种子起始萌发时间显著滞后。(2)CK种子萌发率最高,其次D6处理发芽速度较快,发芽整齐;U处理较D处理的发芽率及发芽势均显著降低,且萌发抑制率显著增加。(3)D处理的种子胚根生长速度快,胚根长度大于CK;U处理的种子胚根生长速度呈先慢后快趋势。可见,凋落物是影响格氏栲种子萌发及胚根生长的重要因素,主要通过阻碍种子与土壤接触而抑制萌发,影响格氏栲林更新。     

Seed bed treatment effects on vegetation and seedling establishment in a New Zealand pasture one year after seeding with native woody species

Summary Efforts to re‐establish indigenous forests in pastoral New Zealand have increased as the value of native biodiversity has been realized. Direct seeding of woody species is preferable to transplanting, as labour and material costs are less. However, the success rate of direct seeding in pasture has been variable due to intense competition from adventive species. We initiated an experiment in pasture plots adjacent to a forest fragment where seed bed treatments (increasing in degree of disturbance from herbicide application to turf removal and topsoil removal) in combination with mulch treatments (wood chip shavings with and without forest floor organic material) were seeded with a mixture of New Zealand lowland forest species. The objective of the study was to determine if early successional plant communities, and ultimately seedling establishment, differed as a result of seed bed preparation after 1 year. Coprosma robusta (Karamu) and Kunzea ericoides (Kanuka) seedlings established on plots in significant numbers: both species were most abundant on topsoil‐removed plots where bare substrate was greatest and plant cover least. Both seed bed treatments and mulching treatments led to measurable differences in overall composition of early successional plant communities. However, absence of plant cover and low soil fertility (both associated with the topsoil‐removed treatment) were the most important factors in seedling success.     

干旱胁迫对鹿角杜鹃种子萌发和幼苗生理特性的影响

为探明鹿角杜鹃种子萌发和幼苗生长期的耐旱性,以鹿角杜鹃干种子和90d苗龄幼苗为材料,采用聚乙二醇(PEG-6000)模拟不同程度的干旱胁迫,研究干旱胁迫对其种子萌发、早期幼苗生长及幼苗的细胞膜透性、MDA含量、有机渗透调节物质和抗氧化酶活性的影响,并对种子萌发率、早期幼苗生长量与PEG胁迫浓度间进行了回归分析。结果表明:(1)5%~25%PEG胁迫范围内,随着干旱胁迫程度的增加,鹿角杜鹃种子的发芽启动时间推迟,发芽持续时间延长,发芽率、发芽势、发芽指数、活力指数和幼苗生长量显著降低;重度干旱胁迫(25%PEG)下,鹿角杜鹃种子完全未萌发。(2)发芽率、发芽势、发芽指数、活力指数以及幼苗生长量的变化均与干旱胁迫程度呈极显著负相关关系,回归分析求得鹿角杜鹃种子萌发的半致死PEG干旱胁迫浓度为15.68%、半致矮PEG干旱胁迫浓度为15.37%。(3)随着PEG胁迫浓度的增加,鹿角杜鹃幼苗叶片SOD活性呈先升后降的趋势,但各胁迫处理仍显著高于CK(0%PEG);细胞膜透性、MDA、脯氨酸、可溶性糖含量、POD和CAT活性则在中度(15%~20%PEG)和重度胁迫下显著升高,与干旱胁迫程度呈极显著正相关关系。研究表明,干旱胁迫显著抑制了鹿角杜鹃种子萌发和早期幼苗生长,使其细胞膜受到损伤,同时鹿角杜鹃可通过体内渗透调节物质和抗氧化酶活性的增加来适应干旱环境,使得自身受抑制、损伤程度降到最低。     

Cover crops influence soil properties and tree performance in an organic apple (Malus domestica Borkh) orchard in northern Patagonia

Little is known about the effects of cover crops on soil properties in organic orchards. To fill this gap, this work aimed at examining the effects of several cover crops on soil fertility, nitrate dynamics, populations of nematodes and tree performance in an organic orchard of apple cv. Royal Gala/EM 26 planted in 1994 at 4 × 2 m. In 1999 the following treatments were randomly applied to the inter-row spaces of 20 tree rows each: permanent cover of alfalfa (Medicago sativa L.) plus fescue (Festuca arundinacea Schribn.), Alfalfa/fescue (AF); permanent cover of strawberry clover (Trifolium fragiferum L.), Strawberry clover (SC); seeding of common vetch (Vicia sativa L.), Vetch (V); and Control (C) (natural vegetation of grasses and legumes with the soil disked twice in late winter, which is the traditional management system used by growers in this region). The cover crops were mowed 3 or 4 times during the growing season and the clippings were left on the ground for decomposition. The trial was conducted in the northern Patagonia region of Argentina in a sandy loam soil with a pH of 7.6 and initial organic matter content of 1.5%. A commercially available certified organic fertilizer containing 5% total N, 2% total P and 4% total K was added annually at a rate of 1.0 kg per tree in equal amounts to each treatment in a radius of 0.4 m around the trunk. After 6 years, soil organic matter in the top soil was 31, 27.9, 23 and 18.6 g kg⁻¹ for SC, AF, V and C treatments, respectively. Total soil N followed the same trend. Apple leaf N declined steadily in all treatments especially in AF and C from year 3 to 5 with values below 1.8% and it was therefore necessary to increase the rate of organic fertilizer. Nitrate concentrations remained under 7.5 mg kg⁻¹ during the winter months in all treatments and increased in spring and summer. A sharp increase in soil nitrate, up to 100 ppm, was observed in late spring in V due to rapid biomass decomposition after maturation of the common vetch. Populations of nematodes especially bacterivores and herbivores increased from September (late winter) to March (fall) in the V treatment. Significant differences were observed when we compared the effects of treatments on the entire nematode population. Tree growth as measured by trunk cross sectional area and canopy volume was significantly lower in the C treatment. Apple yield was also lower in the C treatment in years 5 and 6 while no differences were found among the seeded cover crops. Thus, our soil management treatments did affect soil properties, tree growth and yield. Disking is not a recommended practice because it may decrease the concentration of soil organic matter and leads to poor tree vigor that corresponds to low fruit bearing potential. However, even with the use of permanent cover crops the addition of organic fertilizers is needed in order to sustain yield and tree vigor.     

Effects of fern thickets on woodland development on landslides in Puerto Rico

Abstract. Thicket-forming ferns are common colonizers of disturbed habitats in the tropics, but little is known about their ecology. The effects of thickets formed by the fern Dicranopteris pectinata on tree seedlings on five landslides in the Luquillo Experimental Forest in northeastern Puerto Rico were both positive and negative. Soil moisture and total soil N were higher under fern thickets than in adjacent open areas and soil bulk density and soil surface temperatures were lower. Germination of seeds of the tree Cecropia schreberiana was higher for seeds sown under fern thickets than for those sown into adjacent open areas. Tree seedlings of Tabebuia hetero-phylla exhibited a threefold reduction in photosynthesis under ferns, probably resulting from a twelvefold reduction of photosynthetic photon flux density. Growth of Tabebuia seedlings was reduced under ferns but the distribution of seedlings of naturally occurring woody plants was not strongly correlated with the presence of fern thickets. Although fern thickets on low-nutrient landslide soils appear to facilitate germination, they inhibit growth of tree seedlings and may, therefore, delay forest development on landslides in Puerto Rico.     

Effects of high- and low-intensity fires on soil properties and plant growth in a Bolivian dry forest

We compared soil nutrient availability and soil physical properties among four treatments (high-intensity fire, low-intensity fire, plant removal, and harvesting gap) and a control (intact forest understory) over a period of 18 months in a tropical dry forest in Bolivia. The effect of treatments on plant growth was tested using a shade intolerant tree species (Anadenanthera colubrina Vell. Conc.) as a bioassay. Surface soils in high-intensity fire treatments had significantly greater pH values, concentrations of extractable calcium (Ca), potassium (K), magnesium (Mg), and phosphorus (P), and amounts of resin-available P and nitrogen (N) than other treatments; however, a loss of soil organic matter during high-intensity fires likely resulted in increased bulk density and strength, and decreased water infiltration rates. Low intensity fires also significantly increased soil pH, concentrations of extractable Ca, K, Mg, and P, and amounts of resin-available P and N, although to a lesser degree than high-intensity fires. Low-intensity fires did not lower soil organic matter contents or alter soil physical properties. Plant removal and harvesting gap treatments had little effect on soil chemical and physical properties. Despite the potentially negative effects of degraded soil structure on plant growth, growth of A. colubrina seedlings were greater following high-intensity fires. Evidently, the increase in nutrient availability caused by high-intensity fires was not offset by degraded soil structure in its effects on seedling growth. Long-term effects of high intensity fires require further research.     

Leaf Litter Decomposition and Substrate Chemistry of Early Successional Species on Landslides in Puerto Rico1

Decomposition is a critical process for nutrient release and accumulation of soil organic matter in disturbed soils, such as those found on landslides. I conducted a decomposition experiment on five landslides in the Luquillo Mountains of Puerto Rico as part of an investigation of the successional roles of two of the most common plant colonists to landslides, Cecropia schreberiana Miq. (Cecropiaceae) a pioneer tree species, and Cyathea arborea (L.) Sm. (Cyatheaceae) a pioneer tree fern. I compared leaf litter decomposition over one year and the initial and 1‐yr chemistry for both species. Initial litter chemistry differed between the two species, as Cecropia had slightly higher nitrogen (9.2 mg/g) than Cyathea (8.2 mg/g) and higher lignin (28.6%) than Cyathea (26.0%), but water‐soluble carbon and nonpolar extractable carbon (fats and oils, waxes, chlorophylls) were higher in Cyathea than Cecropia. Total carbon, acid‐soluble carbon, total phosphorus, and pH did not differ significantly between leaf litter species. Across all five landslides, Cyathea (k= 0.93 ± 0.06) leaves decomposed significantly faster than Cecropia (k= 0.68 ± 0.06). The differences in these species leaf litter decomposition rates and chemical composition could potentially influence organic matter dynamics and nutrient cycling rates in these early successional systems.     

Seed rain and soil seed bank of Picea asperata Mast. in subalpine coniferous forest of western Sichuan,China

Abstract

Dragon spruce (Picea asperata Mast.) is widely planted on clear-cuts from natural subalpine coniferous forests in western Sichuan. To assess the natural regeneration potential of this species in spruce plantations of different ages, field studies on the seed rain, seed bank, and seedling recruitment were conducted in 20-year, 30-year, and 60-year plantations, and in a retained natural forest ca. 150 years old for comparison. Moreover, a series of temperature and light regimes were also designed in March 2003 to test germination/dormancy responses of the P. asperata seeds to different conditions. In the plantations considered, both the densities of seed rain and soil seed bank increased with increasing stand age, whereas they were both low beneath the natural forest partially due to low adult density of P. asperata adult trees. P. asperata has a transient seed bank, and ca. 60% of seeds were found in the litter layer. Seed decay and seed predation were the two most important factors affecting soil seed bank dynamics, which caused a number of seed losses from the soil. Only a small fraction germinated and produced seedlings, and the indices of the losses from the seed bank via germination in the 30-year plantation, 60-year plantation and natural forest were 4.75, 5.10, and 2.80%, respectively. High seedling mortality was observed after seedling emergence, and most of the germinated seedlings died out within one growing season. The P. asperata seeds showed a high percent germination and no significant differences under most light and temperature regimes except for full sunlight or temperature below 5°C. In conclusion, despite a substantial number of seed produced, the high depletion of soil seed, the low seedling output, and high seedling mortality may obscure the natural regeneration potential of this tree species.     

Establishment and Control of Hay-scented Fern: A Native Invasive Species

Hay-scented fern (Dennstaedtia punctilobula (Michx.) Moore) is a native forest understory species that behaves as an invasive plant under certain conditions. Previous work has shown that both increased understory light intensity following overstory thinning and removal of competing plants by herbivores can lead to accelerated growth of hay-scented fern, allowing it to develop dense, nearly monospecific understories that inhibit tree seedling regeneration. To study the relationship between these two factors, we sampled 28 forest stands thinned at different times and subjected to different levels of browsing by white-tailed deer (Odocoileus virginianus), and concluded that more than 15 years of intensive browsing following thinning was necessary for ferns to form closed understory canopies with densities of >90 fronds/m² and canopy heights of 60–80 cm; thinning alone or intensive browsing alone was not sufficient to cause this level of fern invasion. We applied three treatments to dense fern understories to determine the relative importance of the fern canopy and the dense mat of roots, rhizomes, and dead fronds in the inhibition of tree seedling establishment. Results after two years were: (1) complete removal of the organic mat produced a large germination response of woody and herbaceous species; (2) mixing the organic mat into the mineral soil produced an initial germination response but poor seedling survivorship, as the fern canopy regrew to near pretreatment density; (3) repeatedly clipping the ferns for two years without disturbing the organic mat resulted in a lower germination response than the removal treatment, but rapid growth of seedlings.     

Distance‐dependent effects of soil‐derived biota on seedling survival of the tropical tree legume Ormosia semicastrata

Question: How do seed germination and subsequent seedling survival of O. semicastrata (Hance forma litchiifolia How) vary with respect to distance from parent trees and conspecific density in different types of tropical forest? Are there effects of soil biota on O. semicastrata that systematically depend on distance from parent trees and conspecific density? Do soil pathogens differently affect survival of O. semicastrata in different types of tropical forest? Location: Tropical lowland rain forest and tropical montane rain forest in Jianfengling National Nature Reserve, Hainan Island, China. Methods: Individual adult O. semicastrata trees were selected in lowland rain forest and montane rain forest. Soil was collected at a distance of 0‐5 m or 15‐20 m from the parent tree. Soil samples from each distance were combined into a bulk sample. Half of the soil sample was sterilized by autoclaving. Surface‐sterilized seeds were then added to the soil material in shade‐houses at both forests. Results: Germination of O. semicastrata seeds at low‐ or high‐seed density was barely affected by the sterilization procedure. In both forests, seedlings grown in non‐sterilized soil collected close to parent trees had significantly higher mortality compared to those in sterilized soil. In contrast, seedling survival with soil collected far from parent trees was not affected by the soil sterilization procedure. Conclusions: Host‐specific pathogens concentrated in the soil around parent trees may regulate community structure of tropical trees at the stage of seedling development.     

The effect of environmental heterogeneity on clonal behaviour of Prunella vulgaris L.

Melinis minutiflora Beauv. (Poaceae) is an African grass that is invading mid-elevation Trachypogon savannas in Venezuela. The objective of this study was to investigate the influence of soil fertility, competition and soil disturbance in facilitating Melinis' invasion and growth in these savanna sites. We manipulated soil fertility by adding nitrogen (+N), phosphorus and potassium (+PK), or nitrogen, phosphorus, and potassium (+NPK). We simultaneously manipulated the competitive environment by clipping background vegetation. In a separate experiment, we mechanically disrupted the soil to simulate disturbance. We hypothesized that germination and growth were bottlenecks to early establishment in undisturbed savanna, but that disturbance would alleviate those bottlenecks. We measured Melinis seed germination and subsequent establishment by adding seeds to all plots. We examined Melinis growth by measuring biomass of Melinis seedling transplants, 11 months after they were placed into treatment plots. Germination and establishment of Melinis from seed was extremely low. Of the 80,000 seeds applied in the experiment, only 28 plants survived the first growing season. Mortality of Melinis seedling transplants was lowest in PK fertilized plots, but in the absence of PK mortality increased with N additions and clipping. By contrast, fertilization of the savanna with NPK greatly increased Melinis seedling biomass and this effect was greatly enhanced when competition was reduced (e.g. clipping). Melinis transplant growth responded strongly to soil disturbance- a response not fully explained by removal of competitors (clipping) or changes in soil nutrients and moisture. We suspect that disruption of the soil structure allowed for greater root proliferation and subsequent plant growth. We believe that native savanna is relatively resistant to Melinis invasion, since Melinis seedlings persisted in intact savanna but exhibited little or no growth during the first year. The significant enhancement of Melinis seedling growth with clipping and nutrient additions suggests that low soil nutrients and the presence of native savanna species are important factors in the ability of native savanna to resist Melinis establishment. However, the potential for Melinis growth increases enormously with soil disturbance.