Conflicting selection on the timing of germination in a natural population of Arabidopsis thaliana

The timing of germination is a key life‐history trait that may strongly influence plant fitness and that sets the stage for selection on traits expressed later in the life cycle. In seasonal environments, the period favourable for germination and the total length of the growing season are limited. The optimal timing of germination may therefore be governed by conflicting selection through survival and fecundity. We conducted a field experiment to examine the effects of timing of germination on survival, fecundity and overall fitness in a natural population of the annual herb Arabidopsis thaliana in north‐central Sweden. Seedlings were transplanted at three different times in late summer and in autumn covering the period of seed germination in the study population. Early germination was associated with low seedling survival, but also with high survival and fecundity among established plants. The advantages of germinating early more than balanced the disadvantage and selection favoured early germination. The results suggest that low survival among early germinating seeds is the main force opposing the evolution of earlier germination and that the optimal timing of germination should vary in space and time as a function of the direction and strength of selection acting during different life‐history stages.     

Aerial and soil seed banks enable populations of an annual species to cope with an unpredictable dune ecosystem

Background and Aims

Simultaneous formation of aerial and soil seed banks by a species provides a mechanism for population maintenance in unpredictable environments. Eolian activity greatly affects growth and regeneration of plants in a sand dune system, but we know little about the difference in the contributions of these two seed banks to population dynamics in sand dunes.

Methods

Seed release, germination, seedling emergence and survival of a desert annual, Agriophyllum squarrosum (Chenopodiaceae), inhabiting the Ordos Sandland in China, were determined in order to explore the different functions of the aerial and soil seed banks.

Key Results

The size of the aerial seed bank was higher than that of the soil seed bank throughout the growing season. Seed release was positively related to wind velocity. Compared with the soil seed bank, seed germination from the aerial seed bank was lower at low temperature (5/15 °C night/day) but higher in the light. Seedling emergence from the soil seed bank was earlier than that from the aerial seed bank. Early-emerged (15 April–15 May) seedlings died due to frost, but seedlings that emerged during the following months survived to reproduce successfully.

Conclusions

The timing of seed release and different germination behaviour resulted in a temporal heterogeneity of seedling emergence and establishment between the two seed banks. The study suggests that a bet-hedging strategy for the two seed banks enables A. squarrosum populations to cope successfully with the unpredictable desert environment.     

Ecological significance of the aerial seed pool of a desert lignified annual,Blepharis sindica (Acanthaceae)

Reproductive traits of a lignified annual plant, Blepharis sindica were studied in relation to the formation of an 'aerial seed pool' on dead plants in an arid grassland in the Thar Desert of northwestern India. The dead plants remained standing on the soil surface and retained fruits for more than one year. Aerial seed pools developed about 6 cm above the ground. There were no seed pools on or below the ground surface. Only 5.7% of seeds died on dead plants because of insect predation or fungi infection during one year. Seed release was cued by rainfall, and a fraction of seeds on the aerial seed pools was released in each rainfall event. After 13 rainfall events during the monsoon season, 25% of seeds was still retained on the plants. Seed predation on the ground surface was intensive; all cones placed on the soil surface were removed within four days, and 97% of fruits were removed within 10 days. Fifty percent of seeds germinated within 3.5 h, and there were no differences in viability and time required for germination between first year seeds and older seeds. The results indicate that the aerial seed-holding on dead plants is an available way to avoid seed predation in harsh desert environments where seed predation is intense and favorable periods for growth are temporally limited and unpredictable.     

Aerial seed bank dynamics and seedling emergence patterns in two annual Mediterranean Asteraceae

Question: We explored the functional significance of seasonal aerial seed banks in two coexisting, heterocarpic annual Asteraceae with dormant (Chrysanthemum coronarium) and non‐dormant (Anacyclus radiatus) achenes. We hypothesised that the plant achene pool is a significant component of total seed reserves, and that within‐season seedling emergence timing is shaped by achene release patterns. Location: SW Spain. Methods: In an observational study, we established temporal achene release patterns. We also quantified the aerial and soil achene pools throughout the release season, and assessed seedling emergence timing. Sowing experiments were used to explore the influence of achene release dynamics on emergence timing, and to establish achene morph‐specific patterns of between‐year distribution of germination. Results: Achene release extended from late spring to late autumn (Chrysanthemum), or from early autumn to early winter (Anacyclus). Within species, achene morphs differed in release timing. Only in Chrysanthemum, a small achene fraction seemed to persist in the soil, and between‐year germination distribution differed among morphs. In coexisting populations, the Anacyclus plant achene pool was an order of magnitude higher than the soil pool throughout the release season, whereas in Chrysanthemum both pools were of the same magnitude during autumn. Within‐year seedling emergence was significantly staggered beneath parent plants compared with the pattern resulting solely from the germination response in soil, with the exception of Chrysanthemum in one of the two study years. Conclusions: Results suggest that seasonal aerial seed banks are effective within‐season, risk‐reducing traits in ruderal Mediterranean habitats characteristic of the study species.     

Population ecology of halophyte seeds

Some aspects of the population biology of halophytes are considered in this review. Persistent seed banks have been reported for a number of inland- and coastal-salt marsh plant communities. Seeds of perennial grasses are often under-represented, while annuals and some perennial forbs may be over-represented in the seed bank. The persistent seed bank of annual halophytes appears adaptive, and provides multiple seed germination opportunities which may prevent local extinction when environmental stress increases. Somatic seed polymorphism provides a mechanism by which parent plants can respond to changing environments by partitioning their resources into reproductive units which have distinct germination responses. Parental effects may influence either seed morphology and/or physiological requirements of seeds when they are exposed to environmental stress. A prolonged germination period can provide plant populations with numerous opportunities to establish seedling cohorts. Early cohorts will have a selective advantage under moderate conditions because mortality will be low and plants will survive until maturity. However, fluctuations in salinity levels and tidal activity can cause high mortality in early cohorts in salt marsh habitats, providing later cohorts with an opportunity for establishment. Resource allocation to reproductive structures is related to plant size, which itself can be affected by both abiotic and biotic factors. Larger plants were found to produce more seeds than smaller plants in a population, but the mean seed weight was greater in small plants.     

Advantage of early seedling emergence in <Emphasis Type="Italic">Fagus crenata</Emphasis>: importance of cotyledon stage for predator escape and pathogen avoidance

To examine how early-emerging seedlings take advantage of establishment in a deciduous forest, we explored the relationships among the emergence date, growth stage and major causes of mortality (damping-off by fungi and predation by rodents) in seedlings of Fagus crenata. The emergence of current-year seedlings and their survival and damage were followed at short (3–7 days in early spring) intervals for a growing season. The growth stage was divided into two stages, with only cotyledons (CT stage) and with true leaves (TL stage). The survival rate was negatively correlated with the emergence date, indicating the advantage of early seedling emergence. This advantage was largely explained by the lower occurrence of damping-off. In contrast, seedling predation occurred independently of the emergence date, but depended strongly on the developmental stage. Rodents consumed a considerable number of seeds during the early period after emergence, and strongly preferred CT-stage over TL-stage seedlings throughout the growing season. Therefore, seedling predation was inferred to be concentrated in a relatively short period while the remaining seeds were depleted and CT-stage seedlings were abundant. The seedling stage synchronously shifted from the CT to TL stage. This growth-stage transition was independent of the emergence date and appeared to correspond with the timing of seed depletion. Delayed stage transition resulted in a disproportionately high risk of damping-off later in the season. Our results indicate that early seedling emergence is advantageous for F. crenata for resistance to pathogens and that the timing of growth-stage progression of seedlings appears important to escape predation because of the distinct food preference of predators.     

Factors affecting seed germination and seedling establishment of a long-lived desert shrub (<Emphasis Type="Italic">Coleogyne ramosissima</Emphasis>: Rosaceae)

Long-lived desert shrubs exhibit infrequent, episodic recruitment from seed. In spite of this long time scale, selection on life history attributes that affect seedling recruitment should be strong. We studied factors affecting germination phenology and seedling establishment for Coleogyne ramosissima, a dominant shrub species in the ecotone between warm and cold deserts in western North America. We also examined ecotypic differentiation in establishment strategy in response to selection regimes in two contrasting habitats. We followed patterns of dormancy loss, germination, emergence, and survival in reciprocal field experiments at warm winter Mojave Desert and cold winter Colorado Plateau study sites. Seed germination took place in late winter, under winter rain conditions at the warm desert site and under snow at the cold desert site. Distinctive germination phenologies for the two seed populations at contrasting field sites followed predictions based on laboratory germination experiments. There was no seed bank carryover across years. Seedling survival at the end of three growing seasons was remarkably high (mean survival 54%). Most seedling mortality was due to sprout predation by rodents early the first spring in unprotected caches. Emergence and establishment at each site were significantly higher for seeds from the local population, supporting the idea of ecotypic differentiation in establishment strategy. Establishment success was an order of magnitude greater overall at the Colorado Plateau site, which represents the leading edge of an upward elevational shift in distribution for this species under the current climatic regime. The Mojave Desert site is on the trailing edge of this shift, and recruitment there is apparently a much less frequent occurrence.     

入侵植物小花山桃草种群构件生物量结构及种子萌发特征

通过野外设置样方调查和室内萌发试验,研究小花山桃草种群各构件生物量的结构特征和它们之间的关系模型、繁殖分配以及种子萌发特点。结果表明:(1)小花山桃草根、茎、叶、花(果)序生物量与植株高度之间以及各构件生物量之间均呈正相关关系,可用幂函数模型或线性函数模型较好地表达;(2)各构件生物量在个体生物量中所占的比率表现为茎>花序>叶>根;(3)小花山桃草的繁殖投入和繁殖分配都随植株个体的增大而增加;(4)小花山桃草个体大小和繁殖投入之间为线性关系,而个体大小和繁殖分配之间为幂函数关系;(5)小花山桃草存在一个较小的繁殖阈值(0.6043g);(6)小花山桃草种子在有光照(12h)和黑暗条件下发芽率均可达到85%以上;未经贮藏的种子不萌发,低温沙藏(1～2℃)和室温干藏(14～32℃)一个半月种子萌发率分别可达92.5%和79%;低温沙藏时种子即可发芽,且发芽率可达61%。在研究地区,小花山桃草几乎整个生长季都可萌发,甚至初冬还有幼苗产生。小花山桃草构件生物量结构和繁殖分配特征、种子萌发特点等都有助于其入侵能力的提高,是其成功入侵我国的重要原因。     

盐生植物种子萌发对环境的适应对策

盐生环境是一种严峻的胁迫环境,对植物的生长、发育、繁殖等生活史的各阶段都产生着重要的影响。盐生植物是生长在盐渍土壤上的一类天然植物区系,它们在长期的进化过程中形成了一系列适应盐生生境的特殊生存策略。一般情况下,盐生植物种子对环境的适应能力,是植物对盐生环境适应性的重要体现;而植物发育早期对盐度的适应能力又是决定物种分布和群落组成的关键因素。在对国内外相关文献进行分析归纳的基础上,从盐分对种子萌发的影响机理及植物种子萌发对盐生环境的适应对策两个方面综述了植物种子休眠萌发与盐生环境的关系。     

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.     

Climate change shifts population structure and demographics of an alpine herb,Anemone narcissiflora ssp. sachalinensis (Ranunculaceae), along a snowmelt gradient

Alpine ecosystems, characterized by cold climates and short growing seasons, are thought to be most vulnerable to climate change. Warmer temperatures and earlier snowmelt extend the growing season length and increase drought stress for alpine plants, resulting in changes to their distribution. Anemone narcissiflora ssp. sachalinensis is a perennial herb that grows in the alpine snow-meadows of northern Japan. In the last few decades, its distribution has shifted toward later snowmelt habitat in the Taisetsu Mountains of Hokkaido. We recorded demographic data for this species at early, middle and late snowmelt habitats over four years (2009–2012), and constructed transition matrix models to evaluate how demographic parameters and population growth rate vary between local habitats along a snowmelt gradient. The proportion of reproductive plants was low and seed production was limited in the early snowmelt habitat, with drier soil conditions, in comparison to the middle and late snowmelt habitats, with moist soil conditions. Evidence of the transition from small plants to those in the reproductive stage was limited in the early snowmelt habitat, suggesting that growth was inhibited; the local population in this habitat was estimated to be sustained by seed migration from later snowmelt habitats. These results indicate that advancing snowmelt under climate change may decrease the reproductive activity and population growth rate of snow-meadow plants if seed migration from later snowmelt populations is limited, resulting in the extinction of local populations.     

Female investment in offspring size and number shifts seasonally in a lizard with single-egg clutches

The timing of reproduction strongly influences reproductive success in many organisms. For species with extended reproductive seasons, the quality of the environment may change throughout the season in ways that impact offspring survival, and, accordingly, aspects of reproductive strategies may shift to maximize fitness. Life-history theory predicts that if offspring environments deteriorate through the season, females should shift from producing more, smaller offspring early in the season to fewer, higher quality offspring later in the season. We leverage multiple iterations of anole breeding colonies, which control for temperature, moisture, and food availability, to identify seasonal changes in reproduction. These breeding colonies varied only by the capture date of the adult animals from the field. We show that seasonal cohorts exhibit variation in key reproductive traits such as inter-clutch interval, egg size and hatchling size consistent with seasonal shifts in reproductive effort. Overall, reproductive effort was highest early in the season due to a relatively high rate of egg production. Later season cohorts produced fewer, but larger offspring. We infer that these results indicate a strategy for differential allocation of resources through the season. Females maximize offspring quantity when environments are favorable, and maximize offspring quality when environments are poor for those offspring. Our study also highlights that subtle differences in methodology (such as capture date of study animals) may influence the interpretation of results. Researchers interested in reproduction must be conscious of how their organism’s reproductive patterns may shift through the season when designing experiments or comparing results across studies.     

Seedling establishment in a masting desert shrub parallels the pattern for forest trees

The masting phenomenon along with its accompanying suite of seedling adaptive traits has been well studied in forest trees but has rarely been examined in desert shrubs. Blackbrush (Coleogyne ramosissima) is a regionally dominant North American desert shrub whose seeds are produced in mast events and scatter-hoarded by rodents. We followed the fate of seedlings in intact stands vs. small-scale disturbances at four contrasting sites for nine growing seasons following emergence after a mast year. The primary cause of first-year mortality was post-emergence cache excavation and seedling predation, with contrasting impacts at sites with different heteromyid rodent seed predators. Long-term establishment patterns were strongly affected by rodent activity in the weeks following emergence. Survivorship curves generally showed decreased mortality risk with age but differed among sites even after the first year. There were no detectable effects of inter-annual precipitation variability or site climatic differences on survival. Intraspecific competition from conspecific adults had strong impacts on survival and growth, both of which were higher on small-scale disturbances, but similar in openings and under shrub crowns in intact stands. This suggests that adult plants preempted soil resources in the interspaces. Aside from effects on seedling predation, there was little evidence for facilitation or interference beneath adult plant crowns. Plants in intact stands were still small and clearly juvenile after nine years, showing that blackbrush forms cohorts of suppressed plants similar to the seedling banks of closed forests. Seedling banks function in the absence of a persistent seed bank in replacement after adult plant death (gap formation), which is temporally uncoupled from masting and associated recruitment events. This study demonstrates that the seedling establishment syndrome associated with masting has evolved in desert shrublands as well as in forests.     

准噶尔盆地东南缘梭梭种子雨特征

为了研究梭梭种子散布规律,通过布设种子雨收集器结合室内实验分析,对准噶尔盆地东南缘梭梭种子雨特征进行了研究。研究显示:(1)梭梭种子雨的累积密度达到平均189粒/m2,其中有活力种子占约80%;(2)种子散布的高峰集中在11月初到11月15日时间段,其落种量占整个种子雨的65%。其后种子雨密度随时间逐渐减小;(3)整个种子雨过程,不同时期散落的种子雨质量存在差异,表现为不同时期散落种子的萌发率呈现出先增大后减小的趋势;(4)变异函数分析表明,梭梭种子雨在8.12 m的有效变程内,种子雨具有明显的空间格局,其由空间自相关和随机因素引起的空间异质性各占50.0%。准噶尔盆地东南缘梭梭种子雨密度大且质量较高,同时其时空分布异质性较高,这些特征均将影响梭梭种群的分布格局和种群更新。     

Wind-dispersed seed deposition patterns and seedling recruitment of Artemisia halodendron in a moving sandy land

* BACKGROUND AND AIMS: Artemisia halodendron is a native sub-shrub that occurs mainly in moving and semi-fixed sandy lands in Inner Mongolia, China. Information on the spatial patterns of wind-dispersed seed deposition and seedling recruitment of A. halodendron inhabiting moving sandy lands is very limited. The aim of this study was to examine wind-dispersed seed deposition patterns and post-dispersal recruitment of A. halodendron seedlings. * METHODS: The spatial patterns of wind-dispersed seed deposition and seedling recruitment of A. halodendron were examined by investigating the numbers of deposited seeds, emerged and surviving seedlings using sampling points at a range of distances from the parent plant in eight compass directions for two consecutive growing seasons. * KEY RESULTS: Wind-dispersed seed deposition showed considerable variation between directions and years. Wind transported A. halodendron seeds only a few meters away from the parent plant in all eight directions. Seedling emergence and establishment also showed between-direction and between-year variability, but the spatial pattern of seedling distribution differed from that of seed deposition. Only a very small fraction (<1 %) of the deposited seeds emerged in the field and survived for long enough to be included in our seedling censuses at the end of the growing season. * CONCLUSIONS: The spatial variation in wind speed and frequency strongly affects the pattern of seed deposition, although the variation in seed deposition does not determine the spatial pattern of seedling recruitment. Seeds of A. halodendron are not dispersed very well by wind. The low probability of recruitment success for A. halodendron seedlings suggests that this species does not rely on seedling recruitment for its persistence and maintenance of population.     

Effects of herbivory and its timing across populations of Trillium grandiflorum (Liliaceae)

The goal of this study was to identify the degree to which the frequency and timing of herbivory by white-tailed deer (Odocoileus virginianus) and subsequent plant response varied across 12 populations of the perennial herb Trillium grandiflorum. Effects of natural and experimental herbivory on the stage and size of reproductive plants were measured. Both the frequency and timing of herbivory varied across T. grandiflorum populations. Reproductive plants were more likely to regress to nonreproductive stages in the next growing season when (1) reproductive plants were consumed by deer (vs. intact reproductive plants); (2) reproductive plants were consumed early in the growing season (vs. reproductive plants consumed late in the growing season); (3) reproductive plants were smaller in size. Clipped plants that remained reproductive were smaller in the following season than unclipped controls. Plant size was positively correlated with the number of ovules, suggesting that reductions in the growth rate of reproductive plants diminish their future reproductive success. Populations with high levels of natural herbivory had a greater proportion of reproductive plants that regressed to nonreproductive stages, probably because reproductive plants in these populations were smaller in size. However, the plant response to herbivory was similar across populations.     

Controls over reproductive phenology among ungulates: allometry and tropical‐temperate contrasts

Ungulates inhabiting high latitudes schedule the timing of conceptions so that offspring are born during the most favourable nutritional conditions for reproductive success. The optimal period for births is less reliably predictable in tropical and subtropical savanna environments where plant growth is governed by rainfall, suggesting that reproductive phenology could be influenced more proximately by resources affecting the body condition of females around the time of conceptions. To assess how these controls operate, we compared the timing of births and conceptions among tropical and subtropical savanna ungulates with the patterns shown by ungulates in northern temperate or subarctic latitudes. The association between the timing of births and the onset of plant growth early in the growing season is less consistent among tropical savanna ungulates than among ungulates inhabiting northern temperate environments, and apparently subject to other influences affecting vegetation phenology. Nevertheless, birth peaks seem to coincide with the time of the year when forage quality is expected to be best for offspring survival and growth for most tropical or subtropical ungulates with gestation periods shorter than a year. When gestation time exceeds one year, proximal effects of nutritional conditions around the time of conceptions apparently become overriding and birth synchrony with early season plant growth is no longer effective. Proximate nutritional influences on conceptions may also govern the somewhat diffuse spread of births shown by ungulate populations in equatorial latitudes where photoperiod cues controlling oestrus and mating cannot be used to schedule the later timing of births.     

冻融过程对荒漠短命植物种子萌发的影响

荒漠区冻融交替显著改变土壤温度和水分条件,并进一步影响荒漠植物种子萌发.为解析荒漠土壤冻融过程对植物种子萌发的影响,本研究以古尔班通古特沙漠4种典型短命植物[东方旱麦草(Eremopyrum orientale)、卵果鹤虱(Lappula patula)、尖喙拢牛儿苗(Erodiumoxyrrhychum)和条叶庭荠(...     

Incorporating seed fate into plant–frugivore networks increases interaction diversity across plant regeneration stages

Plant–animal mutualistic interactions, such as pollination and seed dispersal, affect ecosystem functioning by driving plant population dynamics. However, little is known of how the diversity of interactions in these mutualistic networks determines plant regeneration dynamics. To fill this gap, interaction networks should not only account for the number of seeds dispersed by animals, but also for seed fate after dispersal. Here, we compare plant–animal networks at both the seed dispersal and seedling recruitment stage to evaluate how interaction diversity, represented by different network metrics, changes throughout the process of plant regeneration. We focused on a system with six species of frugivorous birds and three species of fleshy‐fruited trees in the temperate secondary forest of the Cantabrian Range (northern Iberian Peninsula). We considered two plant cohorts corresponding to two fruiting years showing strong differences in fruit and frugivore abundance. Seed dispersal interactions were estimated from a spatially‐explicit, field‐validated model predicting tree and bird species‐specific seed deposition in different microhabitats. These interactions were further transformed into interactions at the seedling recruitment stage by accounting for plant‐ and microhabitat‐specific seed fates estimated from field sampling. We found that network interaction diversity varied across plant regeneration stages and cohorts, both in terms of the evenness and the number of paired interactions. Tree–bird interactions were more evenly distributed across species pairs at the recruitment stage than at the seed deposition stage, although some interactions disappeared in the seed‐to‐seedling transition for one plant cohort. The variations in interaction diversity were explained by between‐plant differences in post‐dispersal seed fate and in inter‐annual fruit production, rather than by differences between frugivores in seed deposition patterns. These results highlight the need for integrating plant traits and disperser quality to predict the functional outcome of plant–animal mutualistic networks.     

VARIATION IN POPULATION GROWTH RATE IN THE WOODLAND ANNUAL IMPATIENS PALLIDA (BALSAMINACEAE)

Variation in population growth rate over environmental gradients was determined for the annual Impatiens pallida (Balsaminaceae), by monitoring age- and size-specific survivorship and fecundity in five populations of this summer-flowering and primarily woodland species. All I. pallida seedlings emerged within a few days of each other and a size hierarchy was established within a month, and remained unchanged thereafter. Light and initial seedling density together explained 67% of the variance in mean adult plant size. As a result of differences among populations in the amount of disturbance, survivorship to the time of floral bud production ranged from 39% to 93% of the initial seedling total. Survivorship during the reproductive period was most affected by plant size and soil moisture. For plants surviving to flowering time, the probability of setting seed, the type of seed and the number of seeds produced per plant were significantly positively correlated with plant size. More than 82% of all seeds were produced by plants greater than or 1 m tall. Cleistogamy was the major form of reproduction in 4 out of 5 populations monitored. The net reproductive rate (R₀) differed greatly among populations, ranging from 1.23 to 16.27. Population growth rate increased with increasing resource availability and decreasing disturbance during the growing season. The magnitude of R₀ depended primarily on 1) timing, intensity and frequency of disturbance, 2) length of the reproductive period, and 3) population size structure.