Seed rain and advanced regeneration in a tropical rain forest

By comparing seed rain, seedling and sapling community structures we assessed the possible role played by vertebrate seed dispersal as a structuring factor in advanced regeneration of closed-canopied sites in the tropical rain forest of Los Tuxtlas, Mexico. Seed weight, initial morphology of seedlings and species abundance were also analyzed to determine if these traits influenced the probability of establishment in the shade.About half of the seed species falling in five closed forest sites (25×25 m) during one year came from fruiting trees growing within the sites (local seeds) and half from fruiting trees found outside the sites (immigrant seeds). Seeds of liana and upper-canopy species were over-represented among immigrant seeds compared with seeds of understory tree species. This probably reflects the activity of frugivorous arboreal mammals, bats, and birds. Species with immigrant seeds had both a lower abundance and a narrower spatial distribution than locally produced seeds. Therefore, immigrant seeds showed higher diversity values than locally produced seeds.Average seed size and the proportion of epigeous seedlings were similar in local and immigrant species. Under closed-canopied sites, factors affecting community organization seem to operate selectively, favoring the establishment of large-seeded, local abundant species in the advanced regeneration. However, the fact that some saplings of immigrant species were found in the plots suggest that a slow species infiltration may be occurring leading to a slow shift in the advanced regeneration species composition. We propose that the influence of seed dispersal on advanced regeneration structure depends on the disturbance history of the patches where seeds land.     

Flowering,fruiting and seed germination in Chilean rain forest myrtaceae: ecological and phylogenetic constraints

Phenological studies in plant communities have generally focused on taxonomically heterogeneous species assemblages, and have only occasionally examined the evolutionary and ecological constraints on the phenological patterns of species within a single family or a genus. Here, we determine relative importance of phylogenetic versus other constraints on the flowering and fruiting periods of 12 species and the germination ecology of 10 species of Myrtaceae sympatric to the temperate rainforest of Isla Grande de Chiloé (42 °Cs 30 S), in southern Chile.We found that, for most species in the family Myrtaceae, flowering was strongly aggregated in January and February. Although this pattern is consistent with the expectation of the 'facilitation' hypothesis (i.e., interspecific overlaps are maximized to attract pollinators), available evidence suggests that pollinators, mainly hymenopterans and dipterans, do not limit fruit production in these species of Myrtaceae in the temperate rainforest. In contrast to flowering, fruiting occurred all-year-round, showing greater segregation in time among the species. According to the their temporal patterns of fruit ripening, two functional groups were defined within the Myrtaceae: those that ripen their fruits immediately after flowering (species in the subtribe Myrtinae) and those in which green fruit develops slowly for several months before ripening (subtribe Myrciinae). Seed germination in the field occurred mainly between August and October. Lab assays showed that the species of Myrtaceae, subtribe Myrtinae, exhibited a long seed dormancy (>40 days), while the seeds of species in the subtribe Myrciinae often germinated within one week after harvesting. The analysis of the phenology of reproductive events in the species of Myrtaceae in this rainforest suggests that: (1) flowering periods patterns are constrained mainly by phylogenetic inertia at the family level, and (2) differences in fruiting patterns and dormancy periods are determined mainly by fruit and seed size, which in turn are associated primarily with phylogenetic closeness within the family, and secondarily with the activity of vertebrate seed dispersers.     

Ecology of seed germination of eight non-pioneer tree species from a tropical seasonal rain forest in southwest China

We compared various aspects of the seed biology of eight non-pioneer tree species from a tropical seasonal rain forest in Xishuangbanna, SW China, that differ in time of dispersal, size and fresh seed moisture content (MC). Seeds were tested for germination under laboratory conditions after dehydration to different moisture levels and under 3.5, 10 and 30% solar irradiances in neutral-shade houses. For six species, germination was also compared in forest understory (3.5% light) and center of a forest gap (32.5% light). Under continuous dehydration over activated silica gel, 100% of seeds of four species had lost the ability to germinate after 48 h, and those of all species except Castanopsis hystrix (decreased from >90 to 30% germination) had lost the ability to germinate after 120 h. Four species did not differ in final germination percentages at the three irradiances (i.e. uniform germination). However, final germination percentages of Horsfieldia pandurifolia and Litsea pierrei var. szemaois were significantly lower in 30% than in 10 or 3.5% light, and seeds of Antiaris toxicaria and C. hystrix germinated to higher percentages in 30 and 10% than in 3.5% light. Mean time to germination (MTG) of the eight species (forest and shade house data combined) ranged from 5–5 days for Pometia tomentosa to 72–207days for L. pierrei; MTG for four species was ≤21 days. There was no obvious relationship between relative desiccation resistance and either time of dispersal, MTG or uniformity of germination at the three light levels, or between seed size and MC or MTG. However, the relationship between seed MC at maturity (25–60% fresh mass basis) and MC at 50% loss of seed viability (12.4–42.5%) was significant. Seven of the species fit Garwood’s (Ecol Monogr 53:159–181, 1983) rapid-rainy germination syndrome and one, L. pierrei, either her delayed-rainy or intermediate-dry germination syndrome. However, fresh, non-dehydrated seeds of all eight species germinated in ≤30 days at constant 30°C in light.     

重庆四面山常绿阔叶林建群种种子雨、种子库研究

对重庆四面山常绿阔叶林建种种种子雨、种子库的研究表明,建群种早期和晚期的种子雨无活力;种子偏早或偏晚成熟及大籽粒的树种,其种子雨被取食的比例大;种子雨、有活力种子雨、种子库三者的数量变化不一致;有活力种子雨量较大的栲、石栎、小叶青冈、扁刺栲、香桂等,其种子库密度在早期以近几何级数的方式增长,元江栲、银木荷种子库小,存在时间短,翌年无一生年萌发苗;种子库数量动态、消减率动态决定于种子被取食的强度、     

The effect of landscape fragmentation and forest continuity on forest floor species in two regions of Denmark

Abstract. Forest species composition was recorded in 82 forests in the Himmerland and Hornsherred regions in Denmark and analysed with respect to isolation (distance to other forests and areas of forest), forest continuity (older or younger than 200 yr), soil pH, tree species composition and seed dispersal groups. Continuity and isolation measures were correlated with forest species richness in Hornsherred. Myrmecochorous, autochorous, anemoballistic and endozoochorous species were markedly fewer in recent than in ancient forests. In Himmerland, patterns were much weaker and few significant correlations were found between forest species richness or different seed dispersal groups and continuity or isolation of the forests. Differences between the two regions may result from less intensive land use, a more humid climate and a smaller species pool with less species with short distance dispersal in Himmerland. Landscape fragmentation therefore appears to limit forest species’recolonization more in Hornsherred than in Himmerland.     

Vertical distribution of Hymenophyllaceae species among host tree microhabitats in a temperate rain forest in Southern Chile

Question: Are differences in microhabitat preferences of co‐occurring epiphytic Hymenophyllaceae species (filmy ferns) correlated with differences in ecophysiological responses to light availability and humidity in the host tree? Location: The Andean foothills in south‐central Chile. Methods: We evaluated the distribution pattern of nine filmy fern species in microhabitats that differ in light availability and humidity in four host tree species. A DCA was developed to assess Hymenophyllaceae species microhabitat preference in terms of canopy openness (CO) and relative humidity. We assessed whether differences in chlorophyll content, maximum photochemical efficiency (Fv/Fm), photosynthetic capacity (A_max), evapotranspiration (E) and instantaneous water use efficiency (WUE) are consistent with any pattern. Results: CO and relative humidity differed significantly with height in the host trees. While CO increased with height in a host tree, relative humidity decreased. DCA analysis showed that filmy fern species distribution within and among trees was mainly explained by the relative humidity of the microhabitat. Chlorophyll content, chlorophyll a/b ratio, A_max and E differed significantly among filmy fern species. A_max and E were correlated with axis 1 scores from the DCA analysis. Conclusions: The vertical distribution and abundance of filmy fern species in Chilean temperate rain forest seems to be closely related to the different microhabitats offered by host trees. This pattern may reflect interspecific differences in ecophysiological traits related both to light availability and humidity. Our results suggest that humidity is the main environmental factor driving functional responses and habitat preferences of these filmy fern species.     

Soil seed bank composition and diversity in a managed temperate deciduous forest

Little is known about the influence of forest management on the interaction between seed bank and aboveground vegetation. We surveyed seed banks and vegetation in 10 forest stands under similar abiotic conditions but submitted either to a coppice-with-standards treatment (n=5) or to a selective-cutting system (n=5). We analyzed species composition and diversity, community ecological profile, and distribution of taxa among different life forms, strategy, morphology and functional type categories. A total of 2085 seedlings (8296 seedsm^–2) germinated-corresponding to 28 species, among which Juncus effusus was the most abundant. Fifty-seven percent of the species were also recorded in the aboveground vegetation, the dominant species being Rubus fruticosus agg., but only 28% of the aboveground species were present in the seed bank. Our results suggest that (1) vernal geophytes and shade-tolerant perennials, which group most true forest species, are not incorporated in the seed bank, (2) parent plants of most seeds were present either in the stand in an earlier dynamic stage or apart from the stand and long-distance dispersed, (3) as expected, early-successional species are well represented in the seed bank, (4) forestry vehicles seem to be a major means of dispersion for stress-tolerant species normally found in forest lanes and wheel tracks. We conclude that seed banks contain species that have a potentially negative impact on the true forest flora and, thus, forest management should minimize soil disturbance and retain remnants of old-coppice woods to conserve disturbance-sensitive true forest species.     

Distribution and abundance of vines along the light gradient in a southern temperate rain forest

Question: Are vines light‐demanding species? Location: Temperate evergreen rain forest of southern Chile (40°39′S, 72°11′W). Methods: In 45 plots of 25 m² distributed in treefall canopy gaps, secondary forest stands and old‐growth forest (15 plots per light environment), all climbing and non‐supported vines were counted and identified to species level, and canopy openness was quantified using hemispherical photographs. Vine abundance and diversity (species richness and Simpson's index) were compared in the three light environments and similarity between vine communities was estimated using Jaccard's similarity coefficient. We also determined the relationship between light niche breadth and local dominance at the species level. Results: In total there were 2510 vine individuals of 14 species. Canopy openness was significantly different in the three light environments. Species richness, diversity, community composition and density of vines were similar in treefall gaps, secondary and old‐growth forest. Of the seven more common vine species, which accounted for 91% of all vines, three had even distribution, two were more abundant in the shaded understorey, and two had higher density in well‐lit sites. Local dominance of vine species and niche breadth were not significantly associated. Conclusions: Our study in a temperate rain forest questions the widespread notion of vines as pioneer‐like species, which may be a consequence of the abundance of some lianas in disturbed sites of tropical forests. Functional arguments are needed to justify a general hypothesis on light requirements of vines, which constitute a vast group of species.     

Seasonal rhythms of seed rain and seedling emergence in two tropical rain forests in southern Brazil

Seasonal tropical forests show rhythms in reproductive activities due to water stress during dry seasons. If both seed dispersal and seed germination occur in the best environmental conditions, mortality will be minimised and forest regeneration will occur. To evaluate whether non-seasonal forests also show rhythms, for 2 years we studied the seed rain and seedling emergence in two sandy coastal forests (flooded and unflooded) in southern Brazil. In each forest, one 100 x 30-m grid was marked and inside it 30 stations comprising two seed traps (0.5 x 0.5 m each) and one plot (2 x 2 m) were established for monthly monitoring of seed rain and a seedling emergence study, respectively. Despite differences in soil moisture and incident light on the understorey, flooded and unflooded forests had similar dispersal and germination patterns. Seed rain was seasonal and bimodal (peaks at the end of the wetter season and in the less wet season) and seedling emergence was seasonal and unimodal (peaking in the wetter season). Approximately 57% of the total species number had seedling emergence 4 or more months after dispersal. Therefore, both seed dormancy and the timing of seed dispersal drive the rhythm of seedling emergence in these forests. The peak in germination occurs in the wetter season, when soil fertility is higher and other phenological events also occur. The strong seasonality in these plant communities, even in this weakly seasonal climate, suggests that factors such as daylength, plant sensitivity to small changes in the environment (e.g. water and nutrient availability) or phylogenetic constraints cause seasonal rhythms in the plants.     

Seed viability of five wild Saudi Arabian species by germination and X-ray tests

Our objective was to evaluate the usefulness of the germination vs. the X-ray test in determining the initial viability of seeds of five wild species (Moringa peregrina, Abrus precatorius, Arthrocnemum macrostachyum, Acacia ehrenbergiana and Acacia tortilis) from Saudi Arabia. Usually several days were required to determine the viability of all five species via germination tests. However, X-ray test will give immediate results on filled/viable seeds. Seeds of all species, except Acacia ehrenbergiana and Acacia tortilis showed high viability in both germination (96–72% at 25/15 °C, 94–70% at 35/25 °C) and X-ray (100–80%) test. Furthermore, there was a general agreement between the germination (19%, 14% at 25/15 °C and 17% and 12% at 35/25 °C) and X-ray (8%, 4%) tests in which seed viability of Acacia ehrenbergiana and Acacia tortilis was very low due to insect damaged embryo as shown in X-ray analysis. Seeds of Abruspreca torius have physical dormancy, which was broken by scarification in concentrated sulfuric acid (10 min), and they exhibited high viability in both the germination (83% at 25/15 °C and 81% at 35/25 °C) and X-ray (96%) tests. Most of the nongerminated seeds of the five species except those of Acacia ehrenbergiana and Acacia tortilis, were alive as judged by the tetrazolium test (TZ). Thus, for the five species examined, the X-ray test was proved to be a good and rapid predictor of seed viability.     

重庆四面山常绿阔叶林建群种种子雨、种子库研究

对重庆四面山常绿阔叶林建群种种子雨、种子库的研究表明,建群种早期和晚期的种子雨无活力;种子偏早或偏晚成熟及大籽粒的树种,其种子雨被取食的比例大;种子雨、有活力种子雨、种子库三者的数量变化不一致;有活力种子雨量较大的栲、石栎、小叶青冈、扁刺栲、香桂等,其种子库密度在早期以近几何级数的方式增长,元江栲、银木荷种子库小,存在时间短,翌年无一年生萌发苗;种子库数量动态、消减率动态决定于种子被取食的强度、种子衰老的速度以及种子对病菌、逆境的抗性和种子萌发的整齐性.     

紫茎泽兰种子种群动态及萌发特性

在紫茎泽兰的入侵地四川省德昌县, 选择撂荒地、农田和马尾松林3个不同的生境, 从种子雨、种子库角度研究了紫茎泽兰种子种群动态规律,及其种子在不同光照和土壤基质条件下的萌发特性. 结果表明,3个生境下紫茎泽兰种子种群仅存在于当年3～6月,于4月底达到年高峰值. 其年降种子雨量在撂荒地中高达2.4×10⁵粒·m^-2,并均分布于2 cm土层以上, 且在撂荒地生境下紫茎泽兰种子的千粒重和活力均显著高于另外两个生境. 到7月上旬, 3个生境下土壤中均未发现有活力的种子. 在实验的光照条件下, 全光照对紫茎泽兰的种子萌发有抑制作用,而在28%光照条件下萌发率最高;砂壤土较适宜紫茎泽兰种子的萌发,生红土中其萌发速度及萌发率均最低.     

Conservation of temperate forest birds in Chile: implications from the study of an isolated forest relict

We characterize the bird assemblage of an isolated relict temperaterainforest (Santa Inés) in terms of its structure, composition anddynamics. The relict character and long-term isolation of this forest remnant,immersed in a matrix of semiarid scrub, provide a natural experiment to assessthe consequences of long-term isolation and forest area reduction for temperatebird species. Bird surveys were conducted in a forest fragment and thesurrounding scrub matrix between April 1999 (austral autumn) and October 2000(austral spring) on a seasonal basis. Within the forest fragment wedistinguished two microhabitat types: creeks and areas far from creeks (i.e.slopes). A total of 36 species were recorded in the study site, of which 21were inside the relict forest. Highest bird abundance and richness wererecorded during winter and spring, and these were always higher in creek plotsrather than slope plots. Comparisons between this assemblage and thosedistributed in the continuous temperate forest (located more than 900km southwards) indicate that this bird assemblage shares moresimilarities, with regard to bird species composition, to southern temperateforest localities and to other relict forests, than to nearer scrub habitats.However, there are eight species, endemic to temperate forests in southern SouthAmerica, missing from our system. In this regard, our analyses indicate thatthese eight endemic species should be of great conservation concern and willlikely be the ones that will go extinct first if the fragmentation of thetemperate forest continues.     

Seed rain of fleshy and dry propagules in different habitats in the temperate rainforests of Chiloé Island,Chile

In temperate rainforests on Chiloé Island in southern Chile (42°S), most canopy trees bear fleshy, avian‐dispersed propagules, whereas emergent tree species have dry, wind‐borne propagules. In the present study, the following hypothesis was tested: regardless of species, fleshy propagules are deposited in greater numbers in canopy gaps and in forest margins and hence have a more heterogeneous seed shadow than wind‐dispersed propagules. To test this hypothesis, the seed rains of these two types of propagules were compared in the following forest habitats: (i) tree‐fall gaps (edges and centre); (ii) forest margins with adjacent pastures; and (iii) under closed canopy (forest interior). Seed collectors (30‐cm diameter) were placed in two (15 and 100 ha) remnant forest patches (n = 60–100 seed collectors per patch) distributed in the four habitats. Seeds were retrieved monthly from each collector during two reproductive seasons (1996, 1997). In both years, the seed rain was numerically dominated by two species with dry propagules (Laureliopsis philippiana and Nothofagus nitida) and three species with fleshy fruits (Drimys winteri, Amomyrtus luma, and Amomyrtus meli). The seed shadows of the two species with dry, wind‐dispersed seeds differed markedly. Seeds of L. philippiana were deposited predominantly in canopy openings, whereas N. nitida seeds fell almost entirely in the forest interior. The fleshy‐fruited species, Drimys and Amomyrtus spp., had similar seed deposition patterns in the various habitats studied, but the between‐year differences in seed rain were greater in Drimys winteri than in Amomyrtus spp. Although no more than 10% of fleshy‐fruited propagules reached the margins of the patch, approximately 7% of these were carried there by birds. Every year, canopy gaps (pooling data from edges and centres) concentrated approximately 60% of the total seed rain of both propagule types in both forest patches. Forest margins received less than 20% of the total seed rain, which was largely dominated by fleshy‐fruited species. Seed shadows were a species‐specific attribute rather than a trait associated with propagule type and dispersal mode.     

Impacts of forest fragmentation on species composition and forest structure in the temperate landscape of southern Chile

Aim Few studies have explicitly examined the influence of spatial attributes of forest fragments when examining the impacts of fragmentation on woody species. The aim of this study was to assess the diverse impacts of fragmentation on forest habitats by integrating landscape‐level and species‐level approaches. Location The investigation was undertaken in temperate rain forests located in southern Chile. This ecosystem is characterized by high endemism and by intensive recent changes in land use. Method Measures of diversity, richness, species composition, forest structure and anthropogenic disturbances were related to spatial attributes of the landscape (size, shape, connectivity, isolation and interior forest area) of forest fragments using generalized linear models. A total of 63 sampling plots distributed in 51 forest fragments with different spatial attributes were sampled. Results Patch size was the most important attribute influencing different measures of species composition, stand structure and anthropogenic disturbances. The abundance of tree and shrub species associated with interior and edge habitats was significantly related to variation in patch size. Basal area, a measure of forest structure, significantly declined with decreasing patch size, suggesting that fragmentation is affecting successional processes in the remaining forests. Small patches also displayed a greater number of stumps, animal trails and cow pats, and lower values of canopy cover as a result of selective logging and livestock grazing in relatively accessible fragments. However, tree richness and β‐diversity of tree species were not significantly related to fragmentation. Main conclusions This study demonstrates that progressive fragmentation by logging and clearance is associated with dramatic changes in the structure and composition of the temperate forests in southern Chile. If this fragmentation process continues, the ability of the remnant forests to maintain their original biodiversity and ecological processes will be significantly reduced.     

Gap-phase dynamics and coexistence of a long-lived pioneer and shade-tolerant tree species in the canopy of an old-growth coastal temperate rain forest of Chiloé Island, Chile

Aim A major question with regard to the ecology of temperate rain forests in south‐central Chile is how pioneer and shade‐tolerant tree species coexist in old‐growth forests. We explored the correspondence between tree regeneration dynamics and life‐history traits to explain the coexistence of these two functional types in stands apparently representing a non‐equilibrium mixture. Location This study was conducted in northern Chiloé Island, Chile (41.6° S, 73.9° W) in a temperate coastal rain forest with no evidence of stand disruption by human impact. Methods We assessed stand structure by sampling all stems within two 50 × 20 m and four 5 × 100 m plots. A 600‐m long transect, with 20 uniformly spaced sampling points, was used to quantify seedling and sapling densities, obtain increment cores, and randomly select 10 tree‐fall gaps. We used tree‐ring analysis to assess establishment periods and to relate the influences of disturbances to the regeneration dynamics of the main canopy species. Results Canopy emergent tree species were the long‐lived pioneer Eucryphia cordifolia and the shade‐tolerant Aextoxicon punctatum. Shade‐tolerant species such as Laureliopsis philippiana and several species of Myrtaceae occupied the main canopy. The stem diameter distribution for E. cordifolia was distinctly unimodal, while for A. punctatum it was multi‐modal, with all age classes represented. Myrtaceae accounted for most of the small trees. Most tree seedlings and saplings occurred beneath canopy gaps. Based on tree‐ring counts, the largest individuals of A. punctatum and E. cordifolia had minimum ages estimated to be > 350 years and > 286 years, respectively. Shade‐tolerant Myrtaceae species and L. philippiana had shorter life spans (< 200 years). Most growth releases, regardless of tree species, were moderate and have occurred continuously since 1750. Main conclusions We suggest that this coastal forest has remained largely free of stand‐disrupting disturbances for at least 450 years, without substantial changes in canopy composition. Release patterns are consistent with this hypothesis and suggest that the disturbance regime is dominated by individual tree‐fall gaps, with sporadic multiple tree falls. Long life spans, maximum height and differences in shade tolerance provide a basis for understanding the long‐term coexistence of pioneer and shade‐tolerant tree species in this coastal, old‐growth rain forest, despite the rarity of major disturbances.     

The role of seed dispersal in the natural regeneration of rain forest after strip-cutting in the Peruvian Amazon

Seed dispersal and forest regeneration were studied on a 30×150 m strip cleared by strip-cutting, a system of forest management designed for sustained yield (Hartshorn 1989), in high terrace rain forest in the Department of Loreto, Peru. After one year the strip was dominated by seedlings of a few bat- and bird-dispersed pioneer tree species (Cecropia spp., Melastomataceae, and Alchornea triplinervia); stump sprouts from cut trees and saplings that survived the clearing were less numerous. The density of saplings (>2 m in height) surviving the clearing was 903 per hectare; 94% of these survived the subsequent 18 months. About 30% of 417 stumps (>7.5 cm diameter at breast height) resprouted within 3 months, with an additional 10% sprouting in the subsequent 10 months. Sprouting frequency was greater for small stumps than large and varied greatly among plant families. Seed deposition over this year was much lower in the interior of the strip, both in species richness and numbers of seeds, than within the forest; strip edges were intermediate in richness and number. The decline in seed input from forest to edge to strip, both in species and in numbers of seeds, was most pronounced for bird-dispersed taxa (primarily Melastomataceae); bat- and wind- dispersed taxa were more evenly distributed. The similarity in bat species composition between the strip and nearby primary forest was higher than the similarity in bird species composition between these habitats, reflecting a failure of many forest bird species to venture into the strip. The predominance of Cecropia spp. and other pioneers of minimal commercial value in the regeneration question the sustainability of strip-cutting. Subsequent succession and future tree species composition on the cleared strip will depend not only on the survivorship and growth of sprouts, survivors, and seedlings, but also on responses of different seed-dispersing animal taxa to changes in the species composition and structure of the vegetation in the strip.     

Light-dependent changes in the relationship between seed mass and seedling traits: a meta-analysis for rain forest tree species

Seed mass is considered to be an important attribute for the establishment success of plant species being linked with their seed production, establishment, and survival. This meta-analysis shows that seed mass is also closely correlated to growth-related species attributes of the established phase of rain forest tree species, and that the strength of this relationship varies with light conditions. Seed mass is an especially good predictor of species traits under high-light conditions, when the species attain their full growth potential. At high irradiance (>20% of full light) seed mass is negatively correlated with RGR, NAR, LAR, SLA and LMF. At low irradiance (<5% of full light), seed mass is only negatively correlated with LAR and SLA. Correlations between seed mass and morphological traits are therefore strongest at low irradiance where light interception is important. Conversely, correlations between seed mass and a physiological trait are strongest at high irradiance, where maximisation of photosynthetic rates is important. The strength of the correlation between growth parameters and seed mass declines over time, and disappears after 1–4 years. Seed mass appears to be a good proxy for the shade tolerance of tropical tree species, especially at the younger stages of the life cycle.     

温带半干旱地区一年生植物种子的萌发特性

在实验室条件下研究了中国温带半干旱地区科尔沁沙地的23种1年生植物的种子萌发特性（新采集种子、冷藏和干藏种子）.大籽蒿、虎尾草、冠芒草、沙蓬和地锦的新种子萌发率达90%左右,11种植物新种子萌发率均低于70%,说明这些植物的新种子具有或多或少的休眠属性.经过150d的冷干藏后,大籽蒿、虎尾草萌发率保持在90%以上,说明这两种植物完全没有休眠机制;冠芒草、沙蓬和地锦的种子萌发率下降较多,可能是储藏的环境条件导致的2次休眠现象;冷藏和干藏处理均能使绿珠藜、毛马唐、细叶益母草、雾冰藜、金狗尾草、苋菜、马齿苋、碱地肤和水稗草的种子在生长季开始时完成生理后熟,萌发率达到80%以上;干藏有利于促进毛马唐、细叶益母草、马齿苋和鹤虱的种子成熟,冷藏有利于促进绿珠藜和金狗尾草的种子成熟;黄蒿、灰绿藜、画眉草和烛台虫实在不同处理下的萌发率都比较低,说明种子内在生理休眠作用较强,具有减少种子一次性萌发数量的风险分摊策略.大多数1年生植物均能在较短时间内达到最终萌发率的90%,表现出迅速萌发的特性;黄蒿、灰绿藜、碱地肤和沙蓬种子则在不同处理中表现出延长萌发时间的策略来适应半干旱地区不确定的环境条件.最后,探讨了几种主要1年生植物的种子萌发对策与其对环境适应机制之间的关系.