Regeneration strategies in a temperate hardwood floodplain forest of the Upper Rhine: sexual versus vegetative reproduction of woody species

The regeneration mechanisms of woody species in the hardwood floodplain forest of the Upper Rhine are not well known, although they are of primary importance for future forest development. To gain a better understanding of the balance between sexual and asexual regeneration strategies and to assess the role of the seed bank in regeneration, the similarities in species composition and abundance of four fundamental compartments involved in regeneration (the seed rain, the seed bank, the recruits and the canopy) were compared in three hardwood forest stands with different flooding conditions. The results show that the floristic composition of the recruits is very similar to that of the canopy, whereas the composition of the seed bank is very dissimilar to the latter and comprises very few hardwood species. However, some species such as Fraxinus excelsior, Carpinus betulus and Acer pseudoplatanus which are very rare in the seed bank release a high number of diaspores, and seedlings of these species germinate abundantly in the field in the year following seed release. Moreover, the germinating seeds originate from the litter layer and not from the soil itself. This shows that most woody species regenerating by seed in the hardwood Rhine forest build transient seed banks and that the role of persistent seed banks for regeneration is very limited. Furthermore, it appears that many woody species have developed strategies favouring vegetative propagation for their regeneration, particularly understorey species, such as Cornus sanguinea and Prunus padus. As vegetatively grown individuals better withstand prolonged inundation in their early life stages than seedlings, species relying on vegetative regeneration strategies might be advantaged by regular and prolonged flooding of the Rhine forest over species regenerating only by seeds.     

The unstable fate of seedlings of the small-seeded pioneer tree species, Betula maximowicziana

The dynamics of the seedling population of the pioneer tree, Betula maximowicziana, were studied for 6 years under various site conditions, with special reference to mortality and mortality factors, in order to better understand their seedling bank formation process. We found that current year seedlings suffered high annual mortality of more than 80%, but that mortality fell with age. Canopy covers and cohorts maintained strong influences on seedling mortality during the first 3 years after emergence. The open site with disturbed soil surface was best suited for 3-year seedling survival. Since the seedlings of B. maximowicziana emerged regardless of canopy covers, high seedling mortality was thought to be the main factor that interrupted their regeneration under a canopy. The occurrence of various hazards together caused high seedling mortality. The major mortality factors, regardless of site conditions, were drought, cold hazard and debilitation, though occurrence of the three factors varied, dependent upon the climate of that particular year. Annual mortality and the occurrence of each mortality factor at a fine scale of 0.2 m² quadrate each showed certain spatio-temporal variations within the same site condition. The extremely high mortality of B. maximowicziana seedlings should be attributed to their inherently small, initial size. The poor root system of the small seedlings may be responsible for their low resistance to various hazards, such as drought and frost heaving.     

Regeneration of commercial tree species in a logged forest in the Selva Maya, Mexico

One of the main threats to the sustainability of community forestry in the Selva Maya is insufficient regeneration of commercial tree species. We evaluated the regeneration status of 22 commercial tree species in a managed semideciduous tropical rain forest in Southern Mexico. The study was carried out in six harvesting areas along a 16-year chronosequence. In each area, 10 transects (1000 m²) were established and all trees >50 cm height and <10 cm diameter were recorded. We evaluated the relationships between seedling and sapling abundance, and canopy cover and disturbance condition (closed forest, canopy gap, log landing, skid road, primary road and secondary road). The area occupied by closed forest canopy increased with age of harvesting area (65–91% of sampled area), while the area occupied by canopy gaps decreased (22–9%). Log landings occupied less than 1% of the sampled area. The predominant canopy cover was 75–80% in all harvesting areas, even in the most recently harvested areas. The highest densities of seedlings and saplings, of both shade tolerant and intolerant species, were found in log landing and skid trails, followed by secondary roads. Even Simarouba glauca, a shade tolerant species, displayed higher densities in sites with ≤65% of canopy cover. Our results support previous findings and indicate that the levels of disturbance caused by existing harvesting procedures may be inadequate to promote sufficient regeneration of not only light demanding desirable species but also for some of the evaluated shade tolerant species of commercial interest. Seedling and sapling densities exhibited by Swietenia macrophylla, for example, are insufficient to support current harvesting rates. The application of a spatial mixed system with patch-cuts of different sizes, a consequence of group felling, could be applied to provide the necessary conditions for the regeneration of the main commercial species.     

Afforestation trials with Prosopis chilensis (Mol.) Stuntz and Prosopis flexuosa D.C. in the Dry Chaco, Argentina

In implantation trials carried out with Prosopis chilensis and Prosopis flexuosa, the following variables were considered: implantation time (December and March), implantation technique (direct seeding and seedling transplantation) and age of the seedlings at the time of transplantation (60, 105 and 150 days). Survival was evaluated for more than 9 months after implantation and trials were repeated in 3 consecutive years.

Results from both species were similar and they show that survival after transplantation was higher (60–80%) than from direct seeding (35–45%). Survival from transplantation was independent of the size and age of the seedlings and it was less affected by climatic conditions than direct field sowing. Survival from direct seeding was higher in March than in December. The best time for either implantation alternative would be the end of the summer or beginning of fall, when climatic conditions increase the probability of seedling survival.     

Season and substrate effects on the first-year establishment of current-year seedlings of major conifer species in an old-growth subalpine forest in central Japan

Seasonal and substrate effects on first-year seedling establishment of major conifer species (Abies mariesii, Abies veitchii, Picea jezoensis var. hondoensis, and Tsuga diversifolia) were assessed in an old-growth subalpine forest in central Japan. Older seedlings (≥1 year) were more abundant on woody debris (WD), such as fallen logs, than on soil. It is known that this preferential seedling establishment on WD is determined by the first-year seedling establishment. The present results indicate that first-summer seedling survivorship clearly contributes to the creation of such seedling–substrate associations, because mortality during the first-summer was greater on soil than WD but that during the first-autumn/winter was not different. Although the standardized mortality rate of current-year seedlings on WD was not so different among the three seasons, that on soil significantly decreased in the winter; indicating that soil is not an unsuitable substrate for seedling survivorship during snow-covered winter. Thus, it can be concluded that biotic and abiotic factors enhancing seedling mortality on soil are most active during summer, and seedling–substrate associations seem to be determined mainly during the first-summer soon after seedling emergence. Furthermore, the first-year survivorship of P. jezoensis var. hondoensis, which has smaller-sized seeds/seedlings that are disadvantageous for early survival, decreased with progressing WD decay toward soil. This suggests that the properties of WD also affect current-year seedling survivorship, especially for species sensitive to substrate properties. In conclusion, initial survivorship affects seedling bank dynamics, and such early dynamics are greatly regulated by the highly heterogeneous substrate availability on forest floors.     

Tree Regeneration and Sapling Damage of Pedunculate Oak Quercus robur in a Grazed Forest in Galicia, NW Spain: A Comparison of Continuous and Rotational Grazing Systems

Forest grazing has been recognised as being a useful tool in fire-risk reduction, in addition to having the potential to preserve or enhance forest biodiversity if managed correctly. Concern for natural regeneration of forest trees in Europe has also prompted interest in the effects of large herbivores on seedling and sapling growth and mortality. An investigation was carried out into sapling damage and density of natural regeneration of oak (Quercus robur) in a mature, pony-grazed, Pinus radiata forest in Galicia, NW Spain under two different grazing regimes (continuous and rotational). In all treatments significantly more oak seedlings and saplings were recorded in areas of grass sward than areas dominated by taller ground flora species. Damage to oak saplings was assessed from the form (height and canopy) relative to stem diameter. The height and average canopy diameter of similar-aged saplings were found to be significantly greater in ungrazed (control) than both continuous and rotationally grazed treatments. Height and canopy diameters of similar-aged oak were not significantly different between the two grazing treatments. Significant differences were observed in tree form, with unbrowsed saplings having the greatest height to canopy width ratio and those in the continuously browsed plots having the smallest. An obvious decrease in the goodness of fit (R²) of regression analyses were found in continuously grazed areas compared to rotational and control plots for both height and canopy data. The differences in damage observed were not significantly different enough to suggest one method of grazing over the other as being better for minimising sapling damage. Management requirements are more likely to dictate grazing regime. Overall, stock density is likely to have a more significant effect on damage than stocking system.     

Recruitment limitation in forests: Lessons from an unprecedented mountain pine beetle epidemic

Since the mid-1990s the forests of central British Columbia have undergone an unprecedented Mountain Pine Beetle (Dendroctonus ponderosae Hopkins) (MPB) epidemic that has resulted in extensive mortality of canopy lodgepole pine (Pinus contorta var. latifolia Engelm.). This study investigated how seed-source availability, seedbed substrate, overstory structure, and time since MPB attack interact to affect post-MPB seedling recruitment of the dominant tree species of these forests. In addition to post-MPB recruitment, these forests may be regenerated by trees established in the understory prior to MPB disturbance. Accordingly, we examined abundance and patterns of all regeneration less than 130 cm tall. We found post-MPB recruitment was sparse. Subalpine fir (Abies laciocarpa (Hook.) Nutt.) comprised the majority of the post-MPB recruitment. It increased with local parent tree basal area and increased strongly with proximity to a major seed source. This resulted in a patchy distribution for subalpine fir post-MPB regeneration. Lodgepole pine post-MPB recruitment was limited by overstory shading. Recruitment of pine decreased as the total overstory basal area increased. Interior spruce (Picea glauca × engelmannii) post-MPB recruitment was similarly limited by total overstory basal area. Seedbed substrates were uniform and dominated by moss. Substrate type distribution did not change as time since MPB disturbance increased. The overall low post-MPB recruitment observed was likely due to a lack of disturbance to the moss-dominated forest floor. Moss is known to be a poor substrate in northern forests. The distribution of all regeneration less than 130 cm tall showed the same trends as the post-MPB regeneration. We believe the post-MPB seedling recruitment dynamics of these forests was not substantially changed from conditions prior to MPB disturbance. There was no pulse of regeneration up to 10 years post-MPB disturbance. Unless this changes, future stand structure will be dominated by the seedling bank established prior to the MPB epidemic. Subalpine fir dominated the seedling bank (68%) and post-MPB recruitment (94%). This suggests that MPB-disturbed forests are undergoing a substantial shift in landscape-level species composition.     

Effects of seed mass and emergence time on seedling performance in Castanopsis chinensis

Seed mass and emergence time may influence fitness of plants. The experiments were conducted to investigate the effects of seed mass and emergence time on the performance of seedlings of Castanopsis chinensis, a large evergreen tree species with limited tolerance of shade. The treatment consisted of three light regimes: 100%, 15% and 2% of full sunlight. Germination rate and germination time showed a significant correlation with seed mass. Germination rate differed among the three light regimes. Early-emerging large-seeded seedlings in 100% light produced the heaviest seedlings, while those resulting from smaller late-emerging seeds and grown under 2% light regime produced the lightest seedlings. Relative growth rate (RGR) decreased with seed mass in all three light regimes, and seedlings of C. chinensis have a higher RGR in high and intermediate light levels compared to low light. In contrast, the effect of emergence time on RGR was not significant. The growth of C. chinensis seedlings were significantly influenced by seed size, emergence time, and light conditions. For each month, the mean height of seedlings was significantly greatest for large-seeded seedlings and significantly lowest for small-seeded ones, irrespective of emergence time and light conditions. Regeneration success of C. chinensis appears to be regulated by the interactive effect of seed mass, emergence time and light regime.     

Scots pine (Pinus sylvestris L.) plantations contribute to the regeneration of holly (Ilex aquifolium L.) in mediterranean central Spain

The aim of this study is to evaluate the recruitment possibilities of holly (Ilex aquifolium L.) populations under Pinus sylvestris plantations in the mountains of Central Spain. Population recruitment of holly under P. sylvestris L. plantations was studied at two localities. We sampled holly seed rain, seed bank, and seedling emergence and survival (of both emerged and planted seedlings) in both localities. Seed germination was analysed by randomly sowing Ilex seeds and their subsequent recollection and analysis over the following 3 years (only in one locality). Seed rain, seed bank and emerged seedling densities did not differ between localities. Seed bank density was between 10 and 20 seeds m⁻², considerably higher than the observed seed rain (0.4–1.2 seeds m⁻²), and also higher than the emerged seedling density (0–4 seedlings m⁻²). But Ilex is not very efficient in germinating under pine forests (29.1% germination after 3 years). This may indicate a seed accumulation process of at least 5 years. Seedling survival for the first year differed among the studied sites (50–60 vs. 17%). Nevertheless these survival rates are higher than those found for the same seedling populations in different forest habitats (not Scots pine forests) from near locations. Differences in summer water stress between the study sites are proposed as the main cause of the observed differences in seedling survival. It is concluded that P. sylvestris forests may provide one of the best regeneration habitats for holly in the Mediterranean area, where this species is close to the Southern limit of its distribution.     

不同种子催芽方法对杜松育苗的影响

采用3种不同的方法对杜松种子进行催芽处理实验,结果表明:相比于混砂催芽方法和温、冷水交替浸种催芽法,采用热水浸种、变温催芽方法处理的种子,播种后出苗时间短,苗齐,其1年生幼苗高生长、地径以及壮苗数量都较高。在育苗上,热水浸种、变温催芽方法要好于其它两种方法。     

Density-dependent selfing and its effects on seed performance in a tropical canopy tree species, Shorea acuminata (Dipterocarpaceae)

In the conservation and management practices of natural forests, sound reproduction and regeneration form the basis of the maintenance and viability of the tree populations. To obtain and serve biological information for sustainable forest management, we investigated reproductive biology and inbreeding depression in seeds of an important dipterocarp tree species, Shorea acuminata (Dipterocarpaceae), by both field and laboratory experiments. Results of parental analysis of immature and mature seeds showed that selfing rates varied greatly, from 7.6 to 88.4% among eight mother trees, and the mean overall selfing rate was 38.3%. Observed outcrossing events within a 40-ha study plot were predominantly (76.5%) short-distance events with a mating distance (md) ≤ 100 m. Since the selfing rate sharply decreased with increase in the number of flowering conspecifics (i.e., individuals of the same species) within a 100-m radius from the mother trees, the local density of flowering conspecifics appears to be the key factor determining the outcrossing rate in S. acuminata. However, the extremely high selfing rate (88.4%) observed for one tree could not be simply explained by the low local density of flowering conspecifics. Instead, differences in its flowering phenology (its flowering peaked ca. a week earlier than most of the other examined individuals) may have severely limited its receipt of pollen from other conspecifics, and thus promoted selfing. Since there were no significant differences in the proportion of selfed progeny between immature and mature seed stages, there was no evidence of selective abortion of selfed seeds during seed development. However, the seed mass of outcrossed progeny was heavier than that of selfed progeny, and heavier seeds showed higher success rates at germination and seedling establishment. These results suggest that inbreeding depression resulted in reductions in seed mass and may reduce the fitness of selfed seeds in S. acuminata. In addition, the outcrossing rate of S. acuminata was more sensitive to low local conspecific flowering-tree densities than that of a sympatric bee-pollinated dipterocarp species with greater pollination distances. These results suggest that the management of local adult-tree densities is important for avoiding selfing and inbreeding depression in future generations, especially in a species like S. acuminata with predominantly short-distance pollination.     

我国樟子松人工林天然更新研究进展

樟子松人工林天然更新问题的研究一直在进行,但人工林内幼苗数量过少及难以发育为幼树的问题一直难以彻底解决。文中综述了影响我国沙地樟子松人工林天然更新能力的因素,如光照、温度、水分、土壤环境、生物因子及人工营林技术等的研究概况。在研究过程中发现,樟子松人工林的遗传多样性与土壤微生物对樟子松人工林天然更新影响方面的研究尚少有报道,或可以为解决樟子松人工林天然更新问题提供更深入的思路。     

Seed mass effects on performance of Pinus halepensis Mill. seedlings sown after fire

Pinus halepensis is a widely distributed species in the Mediterranean basin. It is generally well adapted to regenerate after wildfire, except when fire intervals are too short (≤15 years). In these latter situations, direct seeding could be a good alternative for restoring pine woodland after fire. Under dry climate, low seedling growth and survival can greatly limit the use of seeding. Early studies have shown that seedlings grown from large seeds have higher seedling establishment, growth and survival. Seed size grading may however reduce the genetic diversity of a seed lot by eliminating part or all of the families with relatively small seeds. An alternative to improve seed lot quality without losing genetic variability could be collecting and sieving seeds from each family separately. In order to explore the influence of seed mass on P. halepensis seedling performance, seeds from five half-sib families differing in mean seed mass were sown under greenhouse and field conditions. Final seedling emergence was unrelated to seed mass and half-sib family. The time of emergence was also unrelated to seed mass but it varied among families. Seed mass showed a positive effect on seedling height and diameter, both at population level and within family throughout the study period (9 months under greenhouse conditions and 20 months under field conditions). We also observed a negative relationship between seed mass and relative growth rate for seedling diameter, but it was not high enough to fully compensate the initial differences due to seed mass at the end of the 20-month study period. Seedling predation had a considerable impact on seedling survival, and it was not related to seed mass. When seedlings killed by predation were excluded from the survival analysis, larger seedlings, coming from larger seeds, showed slightly better survival, but only during the first growing period. The small advantages obtained from large seed mass in seedling development do not seem to justify the increased operational costs derived from seed mass selection for each family.     

Enrichment of logging gaps with moabi (Baillonella toxisperma Pierre) in a Central African rain forest

Studies of regeneration in African rain forests suggest that without silvicultural treatments, natural succession in logging gaps may not result in the establishment of timber species. In this paper we present the results of an experimental enrichment planting with moabi (Baillonella toxisperma Pierre), a valuable and important timber species harvested in Central Africa. Although forest gaps are generally considered as favourable for the regeneration of this species, a survey conducted in a forest concession in south-eastern Cameroon provided an estimate of only 12.7 seedlings ha⁻¹, suggesting that the species was, in fact, poorly represented in logging gaps within the study area. To further investigate the dynamics of the moabi in logging gaps, 795 seeds were sown in 15 logging gaps and 410 nursery-raised seedlings were planted in 15 other gaps. A biannual monitoring program over a 30-month period showed a lower survival rate for seedlings from sowing (75.9%) compared to that of nursery-raised seedlings (95.3%). Planted seedlings reached an average of 229.3 cm tall whereas seedlings from sowing were 167.5 cm tall, with the observed difference roughly corresponding to the average height of the nursery-raised seedlings at the time they were introduced to the logging gaps. After 30 months, the diameters of planted seedlings (16.8 mm) were also greater than those of the directly sown individuals (12.5 mm). Forest gap characteristics significantly influenced the growth of the plants. Factors accounting for the differences were total solar radiation, the soil content of coarse sand, the topographic position of the gap, the vegetation cover and the density of Macaranga spp. Whilst total solar radiation had a positive influence on growth, the remaining factors had impacted growth negatively. A streamlined technique was tested by planting 7 seedlings in 250 gaps. Without additional site maintenance, 29.3% of the moabi seedlings emerged naturally from the competing vegetation after 24 months. With biannual maintenance some 89.4% of seedlings became successfully established. Clearance operations had no significant influence on the height of plants whilst plant diameter was greater in cleared gaps. The total cost of the enrichment technique was 5.5 EUR per gap without maintenance and 7.5 EUR per gap with a single maintenance measure. Whilst long-term monitoring is needed, this study suggests a high survival rate of moabi introduced in logging gaps, and a growth rate 10 times higher than previously reported under canopy cover. These findings, combined with the low costs of the enrichment technique, support the use of silvicultural measures in logging gaps to restore the forest.     

Effect of natural disturbances on the abundance of Norway spruce (Picea abies (L.) Karst.) regeneration in nemoral forests of the southern boreal zone

The impact of natural disturbances on the canopy (trees ≥14 m high) and sapling stratum (>0.3 and ≤14 m high) composition was studied in nemoral old-growth forests located within the southern boreal zone in Central Russia (Central Forest Reserve, 32°29′–33°01′E, 56°26′–56°31′N). I hypothesized that the current disturbance regime does not allow the maintenance of current spruce abundance in the canopy, and, as a result, there is a continuous shift in the canopy composition towards a greater abundance of deciduous species. Three 300×20 m² transects were established to estimate the proportions of stand under non-closed unexpanded canopy gaps. Data on sapling composition of 49 canopy gaps were used to analyze pattern of gap refuting in these forests. Additionally, data from three forest inventories showed changes in canopy composition over a period from 1972 to 1990.

The current status of nemoral forests is characterized by the high proportion of stand area under treefall gaps (71%). The loss of spruce from the canopy caused by treefalls (53% of the total basal area of gap-makers) was slightly greater than its canopy abundance (45%). Canopy gaps of all sizes encouraged spruce regeneration which might be due to a decrease in sapling mortality and/or more active recruitment of spruce seedlings. After a gap was formed, the presence of spruce in sapling strata increased. However, within both small (<200 m² in size) and large (>200 m²) gaps, tall (>6 m) spruce saplings did not reach the level of its abundance in the tree canopy. In gaps, tall (>6 m) saplings of lime (Tilia cordata) and elm (Ulmus glabra) grew more quickly than those of spruce and maple. These data suggested a decrease in canopy spruce and an increase in deciduous species in the near future which supported the original hypothesis. Analysis of forest inventory records revealed similar changes in the canopy structure over the past two decades. However, the observed high proportion of stand area under gaps implies that for the next few decades large areas of nemoral communities will be occupied by relatively young stands. This may, in turn, decrease the frequency of large-scale treefalls revegetated mainly by deciduous saplings.     

伯乐树种子育苗试验

本文通过对伯乐树种子采用5种方式,作对比育苗试验,结果表明,C方式出土时间最早、出苗率最高、出苗最整齐,最高平均发芽率为86%,一年生苗平均高18cm,最高28cm,平均地径0.4cm,最大地径0.6cm。     

Impact of bark stripping by sika deer, Cervus nippon, on subalpine coniferous forests in central Japan

We investigated the effect of bark stripping by sika deer, Cervus nippon, on forest regeneration in subalpine coniferous forests on Mt. Ohdaigahara and in the Ohmine Mountains of central Japan. Bark stripping by sika deer occurred in five major tree species: Abies homolepis; Abies Veitchii; Tsuga diversifolia; Picea jezoensis var. hondoensis; and Chamaecyparis obtusa. The percentage of damaged trees on Mt. Ohdaigahara was higher than in the Ohmine Mountains, probably because of the higher deer density. On Mt. Ohdaigahara, the DBH distributions of stems for P. jezoensis var. hondoensis, A. homolepis, T. diversifolia andC. obtusa were bell-shaped with fewer smaller and larger trees. On the other hand, in the Ohmine Mountains the distributions for P. jezoensis var. hondoensis and A. Veitchii showed a reverse-J shaped with more smaller trees. Larger overstory conifers on Mt. Ohdaigahara were killed by bark stripping when 100% barked, although in the Ohmine Mountains ca. 50% of the trees survived even when 100% barked. After the disappearance of the overstory conifers on Mt. Ohdaigahara, the dwarf bamboo, Sasa nipponica, expanded into the forest floor due to changes in light reaching the forest floor. Since S. nipponica is the main forage of deer in this area, this increase caused a corresponding increase in the deer population, which in turn, could cause a further decline in the coniferous forests.     

海南霸王岭陆均松天然更新土壤种子库研究

[目的]了解自然条件下陆均松(Dacrydium pierrei Hickel)土壤种子库的组成和时空动态,揭示其天然林更新机制。[方法]从2014年4月开始,对海南霸王岭陆均松土壤种子库种子数量、成分类型、时空动态进行为期3年的观测,用地统计学等方法分析其与环境因子的关系。[结果](1)陆均松土壤种子库中完好种子、霉烂种子、空心种子、缺损种子占种子库种子总数的比例分别为0.48%、45.55%、29.68%、24.30%;(2)距母树不同距离所含种子数量与距离呈反"J"形曲线递减关系;(3)土壤种子库种子具有明显的空间格局;(4)种子数量垂直分布特征为:枯落物层(87.78%)0~5 cm层(11.01%)6~10 cm层(1.20%),0~5 cm土壤层种子分布急剧减少,至6~10 cm层几乎没有种子分布;(5)土壤种子库无活力种子的数量和比例都较高,最高的100%,有活力种子数量少且萌发率低(14%),种子寿命不到1 a。[结论]土壤种子库的上述特征表明:陆均松具有极大的天然更新障碍,陆均松土壤种子库完好种子数量少,种子质量差、萌发能力弱、寿命较短,种子大量集中在枯落物层,无法与土壤有足够的接触从而降低萌发率。     

The effect of timing of forest fire on phenology and seed production in the fire-dependent herbs Geranium bohemicum and G. lanuginosum in Sweden

The seasonal distribution of fires is one fire regime variable which has received little attention with regard to its effects on plants. For species with a short life-span that recruits after fire, the seasonal timing of a fire can be expected to be important due to effects on potential growth period and reproduction. We observed phenology and reproductive output in two annual and fire-dependent Geranium spp. in the southern part of the European boreal forest. In a garden experiment with the two species under two levels of nutrition, we established cohorts of seedlings at several dates over three summers. Time from germination to flowering and first mature seed differed little between the two species and levels of nutrition; i.e. plant size or level of nutrition had almost no effect on phenology. However, emergence time controlled the timing of reproduction. Most plants emerging before the second week of July in the garden experiment bolted the same year. Plants emerging later behaved as winter-annuals and started to flower in June the following year. A similar dichotomy was observed for populations of Geranium spp. at a number of burnt forest sites that differed in date of fire. This response is likely controlled by photoperiod. Nevertheless, at sites that burnt early some plants did not bolt in the same season; probably an effect of variable seedling emergence dates in the populations. In both the field and garden experiment, there were plants entering reproduction too late to produce mature seeds. Our results indicate that management fires should be conducted either very early, or during July and August to achieve a high seed production in these rare forest plants.     

Improving seed germination and seedling growth of Omphalea oleifera (Euphorbiaceae) for restoration projects in tropical rain forests

To improve the restoration of tropical rain forests, we tested the germination of seeds of Omphalea oleifera collected from soil (S) and from trees (T) in the 2001 dry season (Spring), at the beginning of a dry season (2005a, winter) and in the rainy season (2005b, winter). All seeds had high water content (WC, 31–33%), and the lipid content varied from 14 to 46%. Seedlings from seeds collected in 2001 were subjected to moderate water stress as a preconditioning treatment for severe stress. T-seeds collected in the dry season had high WC, rapid and high germination percentage; S and T-seeds collected in winter (2005) had also high WC but were dormant. GA₃ (250 ppm) broke this dormancy. S-seeds collected in the dry season or at the beginning of it had relatively low WC and low and delayed germination. Some 2001 S-seeds produced albino seedlings. The critical water content for maintaining ecological longevity in these seeds was ∼15%. Seeds collected in 2005b that were dehydrated for 20 days in a moist and fresh atmosphere lost their viability, showing recalcitrant behavior. T-seeds with the lowest lipid content (2005a) after dehydration maintained low germination (15 ± 18%). In all samples the seed size varied widely and was not predictive of seed WC. Embryos taken from dehydrated seeds had two to four times higher WC than the seeds, but germination did not take place. Laboratory and field germination of dormant seeds showed that viability may be maintained for at least 2–3 months on a moist substrate (soil or agar). Moderate water stress at the seedling stage reduced the efficiency of biomass production. Response to this water stress was expressed more in physiological traits than in morphological characters, consequently biomass allocation was maintained and plants retained most of their morphological characteristics (root:shoot ratio, leaf area ratio, specific leaf area, leaf weight ratio). Moderate water stress did not increase the tolerance of seedlings to severe stress, causing leaf shedding and plant death. For restoration purposes we recommend that T-seeds be germinated immediately avoiding dehydration. The use of S-seeds could result in unhealthy seedlings. Seed recalcitrance and the response to moderate water stress restrict germination and establishment to small gaps, where this species naturally grows. We suggest that before introducing O. oleifera in restoration programs, a plant cover should be built to reduce soil water deficit. It is necessary to improve methods to increase potential seed longevity in storage.