Synchrony and asynchrony: observations and hypotheses for the flowering wave in a long-lived semelparous bamboo

Aim (1) To describe the spatio‐temporal patterns of mass‐flowering and die‐off in a long‐lived, semelparous, clumping bamboo, Bambusa arnhemica, at landscape and local scales. (2) To discuss causal processes in the flowering patterns of semelparous bamboos. Location The entire range of B. arnhemica, in the monsoonal, tropical, north‐west of the Northern Territory of Australia, mostly along watercourses. Methods Landscape‐scale flowering patterns were assessed by a combination of air, boat and ground survey in each year from 2000 to 2002. Areas that flowered prior to 2000, and those in which no flowering occurred, were also recorded, and historic records collated. At local scales, initiation of flowering, rates of die‐off, and subsequent germination densities of seedlings were quantified by ground‐based counts. Results After an estimated 40–50 years of vegetative development, B. arnhemica flowered, seeded prolifically, then died. Flowering occurred synchronously within patches ranging from 0.002 to 3200 km². One or more patches flowered in successive years from 1996 to 2002, forming a temporally‐structured but spatially‐chaotic flowering wave that affected c. 80% of the population. Synchronous flowering took the form of a flowering distribution in which over 95% of clumps within a patch initiated flowering in a central year, most of the remainder flowering the year before or after. Along the Daly River, an exception was observed in which 56% of clumps flowered in the peak year. Seedling densities were three orders of magnitude greater under clumps that flowered in the central rather than the leading year of the flowering distribution. Main conclusions Synchrony is argued to be the primal state in semelparous bamboos, promoted by intense selection acting on a endogenous (genetic or biological) clock whose influence largely overrides that of the environment. A flowering wave may develop within an initially synchronous population when stochastic events interact with the biological clock without permanently altering the clock setting, producing an off‐set patch. Off‐set groups may only survive if sufficient individuals are off‐set by the same amount at the same time and in the same vicinity so as to produce a new synchronously‐flowering patch. This could be driven by two processes. Inter‐year climatic variation may alter the biological clock's perception of time, producing off‐sets at local or regional scales or even affecting entire populations. Severe environmental pressures may also force one‐off changes to flowering schedules, as suggested by a severe flood event prior to flowering on the Daly River. A dynamic hypothesis for a wider range of bamboo flowering patterns is proposed in which synchronous flowering is fragmented and disrupted over time but renewed by allochronic speciation and dispersal.     

Interactive effects of introduced herbivores and post‐flowering die‐off of bamboos in Patagonian Nothofagus forests

Question: In November 2000, Chusquea culeou, a bamboo species dominating Andean forest understories in southern Argentina and Chile, massively flowered and died over a north‐south distance of ca. 120 km. Because bamboo is the major forage for large herbivores in these forests, we examined the interactive influences of the bamboo die‐off and herbivory by introduced cattle on understory and tree regeneration. Location: Lanín National Park, Argentina. Methods: Permanent plots, in and outside livestock exclosures, were installed in a Nothofagus dombeyi forest in patches of flowered and non‐flowered C. culeou. Plots were monitored over four years for changes in understory composition and tree seedling densities and heights. Results: After the C. culeou die‐off, new establishment of N. dombeyi was low, both with and without herbivory. Livestock alone directly increased N. dombeyi seedling mortality through physical damage. However, tree seedling browse ratings and height growth were interactively affected by bamboo flowering and herbivory; unfenced plots in flowered bamboo patches had the shortest seedlings, highest browse ratings, and lowest tree seedling annual growth rates. Understory cover was higher where livestock were excluded, and this effect was intensified in the patches of flowered bamboo. Neither herbivory nor bamboo flowering resulted in major changes in species composition, with the exception of Alstroemeria aurea. Conclusion: Effects of livestock on N. dombeyi regeneration were contingent on flowering of C. culeou. Prior to introduction of livestock, N. dombeyi regeneration was probably successful beneath canopy gaps during windows of opportunity following bamboo die‐off, but now livestock impede tree regeneration. Herbivory during bamboo withering periods also produces more open understories, particularly affecting palatable heliophyllous herb species such as Alstroemeria aurea. The results underscore the importance of assessing herbivore impacts on tree regeneration during relatively short periods of potential tree regeneration immediately following rare bamboo flowering and die‐off.     

神农箭竹开花前后植物体内N、P、K元素的动态

对神农箭竹(Fargesia murielae)N、P、K的含量进行分析,结果表明:N、P、K在竹子植物体内呈非均匀分布。N和P在各器官中的分布规律为:叶>鞭、根>竿;K在未开花竹和正开花竹中分布规律为:鞭>根>叶>竿,在已开花竹中分布规律为:叶>鞭>竿>根。随着开花过程的进行,N的含量在叶、竿、鞭和根等器官中逐渐减少;P的含量在竿、鞭和根等器官中逐渐降低,在叶片中先升高后降低;K的含量在鞭和根中呈下降趋势,在竿和叶中先降低后升高。     

带状间伐对巴山木竹林复壮更新及大熊猫取食的影响

间伐是竹林复壮更新的有效手段之一.以陕西佛坪国家自然保护区为研究区域,对大熊猫(Ailuropoda melanoleuca)冬季栖息地非主要采食区域竹林老龄化严重且密度过大的巴山木竹(Bashania fargesii)林进行为期3年(2017至2019年)的带状间伐样方监测,研究间伐对竹林复壮更新以及大熊猫觅食选择...     

Bamboo-Dominated Forests of the Southwest Amazon: Detection,Spatial Extent,Life Cycle Length and Flowering Waves

We map the extent, infer the life-cycle length and describe spatial and temporal patterns of flowering of sarmentose bamboos (Guadua spp) in upland forests of the southwest Amazon. We first examine the spectra and the spectral separation of forests with different bamboo life stages. False-color composites from orbital sensors going back to 1975 are capable of distinguishing life stages. These woody bamboos flower produce massive quantities of seeds and then die. Life stage is synchronized, forming a single cohort within each population. Bamboo dominates at least 161,500 km² of forest, coincident with an area of recent or ongoing tectonic uplift, rapid mechanical erosion and poorly drained soils rich in exchangeable cations. Each bamboo population is confined to a single spatially continuous patch or to a core patch with small outliers. Using spatial congruence between pairs of mature-stage maps from different years, we estimate an average life cycle of 27–28 y. It is now possible to predict exactly where and approximately when new bamboo mortality events will occur. We also map 74 bamboo populations that flowered between 2001 and 2008 over the entire domain of bamboo-dominated forest. Population size averaged 330 km². Flowering events of these populations are temporally and/or spatially separated, restricting or preventing gene exchange. Nonetheless, adjacent populations flower closer in time than expected by chance, forming flowering waves. This may be a consequence of allochronic divergence from fewer ancestral populations and suggests a long history of widespread bamboo in the southwest Amazon.     

四川引种大花序桉花期与蒴果发育时空分布特征研究

为了探索大花序桉（Eucalyptus cloeziana）引种后花期和蒴果发育规律,对四川宜宾引种的5个种源7年生大花序桉花期和蒴果发育进行了研究。结果表明：（1）大花序桉花芽主要分布在树冠下层内膛,部分花芽直接从主干上萌发,同一个花枝的花蕾随机开放;（2）同一种源的大花序桉始花期差距不超过5 d,末花期差距不超过10 d,花期持续41~62 d,5个种源大花序桉花期从当年12月中下旬至翌年5月上旬,因此种源间存在花期不遇;（3）不同种源大花序桉开花率存在显著性差异,其中72号种源开花率（53.94%）最高,74号种源尚未生殖生长;（4）根据蒴果颜色和果爿状态将蒴果发育分为：发育早期、果爿形成期、果爿张开期和散种期。     

圈养大熊猫食谱组成与营养成分分析

为了探讨圈养大熊猫主食竹食谱组成及主要影响因素,并为圈养单位提供科学合理的食物营养配比和饲养管理建议。本研究通过对成都大熊猫繁育研究基地１０ 只成年大熊猫４ 年（２００９. ０３ －２０１３.０２）的饲养数据,３只成年大熊猫１ 年（２０１２. ０７ －２０１３. ０６）的觅食行为观察数据,以及投饲的３ 种竹笋和５ 种主食竹常规养分和总黄酮含量的统计分析。结果发现,基地圈养大熊猫偏好竹笋,喜食巴山木竹叶、白夹竹茎和苦竹茎,随机采食箬竹叶和刺竹茎;基地大熊猫根据投饲竹种类和季节的变化,形成了较为稳定的食谱组成;对投饲竹各部位成分比较分析发现,投饲竹养分含量的变化呈现出一定的规律性：其中粗蛋白、粗脂肪、蛋白质和能量比都以竹叶最高,竹枝次之,竹茎最低;干物质和粗纤维的含量则以竹茎最高,竹枝次之,竹叶最少;竹茎和竹枝呈现钙少磷多的现象,而竹叶则是钙多磷少;各投饲竹不同部位总黄酮含量以叶中最高,枝中其次,茎中最低。研究结果表明,圈养大熊猫主食竹食谱的选择遵循能量和营养学的规律,选择适口性强、营养价值高的食物假说。     

Seed Set and Natural Regeneration of Dendrocalamus membranaceus Munro after Mass and Sporadic Flowering in Yunnan,China

The flowering periods of woody bamboos, seed set, natural regeneration and death after flowering have been rarely observed and evaluated in the field. Dendrocalamus membranaceus Munro, a tropical woody bamboo mainly distributed in the Yunnan, displayed both sporadic as well as gregarious (mass) flowering and fruited from 2006 to 2013 following severe droughts. The aim of this study is to examine potential differences in seed set and natural regeneration between the two flowering patterns in natural D. membranaceus forests. We investigated and analyzed seed set, seed germination, seedling growth and mortality in both mass and sporadic flowering populations. Observations were made over a period of three years to record changes in bamboo seedling density, height and culm diameter. We observed a low natural seed set ranging from 1.76% to 7.49%, and a relatively high seed germination rate in the nursery from 59.6% to 71.0% for both types of flowering populations. Seeds germinated in 5–7 days after sowing and the germination period lasted 10–15 days. Seed set and germination rates in mass-flowering populations were significantly higher than those of sporadically flowering stands. The seedlings within sporadically flowering populations died within two years. In comparison, seedling mortality in the mass flowering population increased over two periods of observation from 64.92% to 98.89%, yet there was good seedling establishment left over, which showed mean height and mean culm diameter increasing by 1053.25% and 410.71%, respectively, in the second year of observations, and 137.10%, and 217.48%, respectively, in the third year. There are significant differences in seed set, natural regeneration ability and sustainability of bamboo populations between the mass flowering and sporadically flowering populations of D. membranaceus. Sporadic flowering populations failed to produce effective regeneration, while mass flowering populations were able to regenerate successfully. This study provides useful insights for conservation and natural forest management of D. membranaceus. We consider the merits of introducing other genetic provenances towards long-term maintenance of the stand features at sporadically flowering sites; meanwhile, the most economic option for mass flowering stands is to allow natural regeneration to take place through protecting such sites from further disturbance.     

Ecological consequences of a massive flowering event of bamboo (Chusquea culeou) in a temperate forest of Patagonia,Argentina

Question: What changes occur as a consequence of the massive flowering and senescence of the dominant understory species of bamboo, Chusquea culeou (E. Desvaux)? In this study, we documented some of the ecological consequences of this rare event that occurred in 2001, the previous flowering having occurred more than 60 years ago. Location: Nothofagus temperate forest, Patagonia, Argentina. Methods: We assessed changes in environmental variables and bamboo biomass post‐flowering in an old‐growth southern beech forest. In addition, we monitored the demography of emergent Ch. culeou seedling and Nothofagus nervosa saplings, comparing non‐flowered (live understory) and flowered (senescent understory) patches within the forest matrix. Results: Bamboo flowering dramatically increased light availability in the forest understory but, surprisingly, other environmental changes were not observed. Bamboo seedlings emerged in both patch types, and experienced gradual but modest mortality through time. Bamboo dieback promoted higher survivorship and an increment in biomass, height, number of leaves and buds in the saplings of Nothofagus nervosa. Conclusion: The high density of bamboo seedlings 5 years after the flowering event and the independence of emergence from environmental variables suggest that understory regeneration is a gradual process that is not strongly regulated by initial seedling density or resource limitation. In contrast, microenvironmental conditions created after the flowering event significantly increased Nothofagus sapling growth and survival. These results suggest that overstory forest regeneration could be enhanced in this temperate forest in the first years after this infrequent bamboo flowering event.     

Localization of lily symptomless virus and cucumber mosaic virus in anther- and filament-derived calluses and effect of callus culture duration on virus-free bulblet production in Lilium × ‘Enchantment’

Bamboos represent one of the world’s great natural and renewable resources. The study reports precocious rhizome formation in multiple shoots of elite, rare, woody bamboo Bambusa bambos var. gigantea. Multiple shoots were initiated from embryonic axes of caryopses inoculated on MS-basal medium supplemented with 5.0 μM BAP and 2% sucrose. Transfer of shoots to MS basal medium supplemented with BAP (2.5 μM, 5.0 μM), GA₃ (0.1 μM) and NAA (50.0 μM) and 5% sucrose led to 58% to 100% rhizome induction within four weeks of culture. Subsequently, these rhizomes developed roots on auxin media and formed culm shoots showing regeneration of plantlets after eight weeks. Incorporation of TIBA inhibited rhizome formation. The plantlets with rhizomes were transferred to soil. Precocious rhizome formation will lead to early establishment besides providing propagules on demand and mass multiplication of bamboos through rhizome banks.     

黑龙江省植物花期对气温变化的响应

为了解黑龙江省植物花期对气温变化的响应趋势。利用1951~2007年采集于黑龙江省的188种植物标本信息,分析黑龙江省植物花期变化与年均温、上年均温和春季、夏季、秋季均温变化的关系。结果表明:黑龙江省42.0%以上植物的花期与气温间存在显著的相关性(P<0.05)。这些植物中,25%以上物种的相对采集时间指数与年均温、上年均温及各季节均温存在显著正相关(P<0.05)关系,且多为草本植物。其余草本植物及木本植物的相对采集时间指数与年均温、上年均温及各季节均温多呈显著负相关(P<0.05),如春季均温每升高1℃,木本植物花期平均提前12.65%。年均温、上年均温和春季、夏季、秋季均温每升高1℃,黑龙江省植物花期分别整体提前4.47%、7.08%、3.87%、5.84%和6.73%。花期对气温变化的响应,在短花期植物中相对明显。相较于春季、秋季及年均温,黑龙江省植物花期对夏季均温变化的响应更为敏感。     

Clonal Structure,Seed Set,and Self-Pollination Rate in Mass-Flowering Bamboo Species during Off-Year Flowering Events

Bamboos are typical examples of highly synchronized semelparous species. Their mass-flowering events occur at supra-annual intervals but they sometimes flower on a small scale in off-years. If some bamboo ramets (culms) of a genet flower and die in off-years, whereas other culms of the same genet do not flower synchronously, the genet can still survive blooming in an off-year and could participate in the next mass-flowering event. At genet level, the effect might be similar to that achieved by synchronously reproducing iteroparous plants. In addition, if multiple genets flower simultaneously in off-years, cross-pollination will be promoted. However, it is not known whether all the culms in a genet flower synchronously and whether multiple genets flower in off-years. We determined the clonal structure of three temperate dwarf bamboo species, i.e., Sasa senanensis, S. kurilensis, and S. palmata, at 24 off-year flowering sites and the surrounding areas in northern Japan using seven microsatellite markers. We also estimated seed set at seven of the sites and self-pollination rates at five sites to determine off-year reproductive success. Next, we investigated whether seed sets at the culm level were related to flowering area and/or number of flowering genets, using generalized linear mixed-effect models (GLMMs). Multiple genets flowered at 9/24 flowering sites. We found that 40/96 of the genets identified had some flowering culms. Non-flowering culms were present in 24/40 flowering genets. Seed set was in the range 2.2%–12.5% and the self-pollination rate was 96.3%. In the best GLMM, seed set increased with flowering area. Seeds were produced in off-years, but cross-pollination was rare in off-years. We suggest that some dwarf bamboos may exhibit iteroparity or imperfectly synchronized semelparity at the genet level, a characteristic similar to that of other reproductively synchronous plants. We also found synchronous flowering of a few genets even in off-years.     

Seed germination and seedling growth in the arrow bamboo <Emphasis Type="Italic">Fargesia qinlingensis</Emphasis>

Improving natural regeneration of bamboos after they die following mass flowering is critical for conservation of giant pandas. However, little is known about factors that affect seed germination and seedling growth of bamboos. We studied seed germination and seedling growth in Fargesia qinlingensis, which mass flowered in a giant panda habitat in the Qinling Mountains of China in early 2000, in laboratory and greenhouse conditions. Seed germination rate was tested under light and dark conditions 5 and 12 months after seed collection. Germination rate displayed no significant difference under light or dark conditions 5 months after seed collection, but was significantly greater in the dark than under light 12 months after seed collection, suggesting light inhibition of seed germination. A 2×2 factorial design was conducted to test the effects of nitrogen (N fertilization and non-N fertilization) and light [full sun and shade (i.e., 14% full sun)] on seedling growth and biomass allocation. N fertilization significantly increased seedling growth, resulting in greater seedling height, more branches, more leaves, greater stem biomass, and greater leaf biomass. Seedlings under 14% full sun conditions had a significantly lower percentage of biomass allocated to the stem. The root/shoot ratio was significantly greater in non-N/shade than non-N/full sun, while there was no significant difference in this ratio between N/shade and N/full sun, suggesting that nitrogen fertilization compensated for the effect of shade on biomass allocation. Our results suggest that N fertilization could be employed in restoration of F. qinlingensis stands after die-off following mass flowering.     

带状采伐对毛竹地下构件养分含量的影响

选择立地条件基本一致的毛竹(Phyllostachys edulis)纯林,分别设置宽度为6 m、8 m、10 m,垂直长度为30 m的样地进行条带式采伐,在采伐带相邻的位置设置相同宽度的保留带,另随机选取3块不进行采伐的常规经营毛竹林作为对照,测定不同宽度带状采伐毛竹竹鞭、鞭根、竹篼和篼根的养分情况,分析毛竹地下构件在不同宽度采伐后的恢复潜力,探寻合理的带状采伐宽度,为毛竹林的科学管理提供理论依据。结果表明:(1)采伐带毛竹地下竹鞭、鞭根和篼根有机碳含量均随采伐带宽的增加先增后降,竹篼有机碳含量则持续上升;3种宽度采伐带篼根、8 m采伐带鞭根和10 m采伐带竹篼有机碳含量均显著高于保留带(P<0.05),8 m采伐带鞭根、3种带宽采伐带竹篼和篼根有机碳含量均与对照差异显著;6 m和10 m采伐带篼根、8 m采伐带鞭根有机碳含量所占地下构件有机碳比例较保留带有大幅提升。(2)3种采伐带宽中竹鞭、鞭根和竹篼全氮含量均表现为8 m>6 m>10 m,篼根全氮含量则随着带宽的增加持续上升;8 m采伐带鞭根和10 m采伐带竹篼与保留带有显著差异,3种带宽鞭根、竹篼和篼根全氮含量均与对照有差异;各构件总氮含量所占比例均基本表现为竹鞭>竹篼>鞭根>篼根,8 m采伐带鞭根和10 m采伐带竹鞭、鞭根、篼根总氮所占比例显著增加。(3)采伐带毛竹竹鞭、竹篼和篼根全磷含量均随带宽的增加呈先增后降的趋势,而鞭根全磷含量持续上升;8 m采伐带竹鞭全磷含量显著高于对照,其竹篼全磷含量显著高于相应保留带和对照;8 m和10 m采伐带均以竹篼全磷含量所占比例最高,6 m采伐带则是竹鞭全磷占比最高。(4)采伐带毛竹地下构件全钾含量均以8 m带宽最高,8 m采伐带的竹篼和篼根全钾含量分别显著高于6 m采伐带17.69%和106.90%,显著高于10 m采伐带102.87%和121.12%;8 m采伐带竹鞭、鞭根和篼根全钾含量均显著高于保留带和对照;随着带宽的增加,全钾含量占比在采伐带竹鞭和竹篼中呈下降趋势,在鞭根中则呈上升趋势,在篼根中则先上升后降低。研究认为,带状采伐处理能促进毛竹地下构件的发育,且8 m采伐带毛竹地下构件对养分的吸收效率显著增加,养分元素含量相对较高,可以为竹林恢复生长提供大量能量和生长潜力,有利于采伐后毛竹林的快速恢复,是较为合理的采伐带宽度。     

海拔对大熊猫主食竹结构、营养及大熊猫季节性分布的影响

大熊猫主要采食竹子,因此主食竹对大熊猫生存具有重要的作用。在秦岭的佛坪地区的大熊猫主要取食巴山木竹以及秦岭箭竹。本文研究了海拔对这两种竹林的结构与营养含量的影响以及海拔与大熊猫季节性分布的关系。结果表明：（1）海拔对竹林基径、株高有显著影响（p0.05）;整齐度、均匀度、基径和株高分布的偏度值和峰度值均随海拔变化而变化。（2）海拔对主食竹营养含量的显著影响随季节而变化（春季粗蛋白和总糖p=0.02;夏季粗纤维p=0.01;秋季粗蛋白p=0.04、粗纤维p=0.04和总糖p0.01）。每个竹种在叶、枝、杆三个部位间的营养均有显著性差异（p0.05）。（3）海拔对竹林结构及营养的显著影响决定了大熊猫对主食竹的取食策略,夏季在高海拔活动,其余季节在低海拔活动。本文的研究结果对理解海拔、主食竹结构、营养以及大熊猫迁移活动之间的关系有重要作用,为圈养大熊猫的饲料配比提供理论依据,也为野外大熊猫的保护和规范保护区内部的人类活动提供科学支撑。     

汶川地震对大熊猫主食竹——拐棍竹竹笋生长发育的影响

为了了解汶川地震对大熊猫主食竹生长发育的影响,2009年10月—2011年4月,在地震重灾区四川龙溪-虹口国家级自然保护区内设置150个样方,就地震强度、中度、轻度干扰对大熊猫主食竹拐棍竹(Fargesia robusta)竹笋生长发育的影响进行了研究。研究结果显示:(1)发笋物候上,地震强度干扰的发笋时间(Julian时间)较中度和轻度干扰显著推迟,而发笋期不受影响。(2)发笋量和当年生幼竹数是拐棍竹更新生长的数量指标。地震强度和中度干扰样方的发笋量都显著高于轻度干扰,而发笋量在强度和中度干扰间差异不显著;地震强度干扰样方的当年生幼竹数显著高于中度和轻度干扰,中度和轻度干扰样方间差异不显著。这说明地震的强度和中度干扰都有利于拐棍竹竹笋和当年生幼竹数量的更新。(3)基径和株高是拐棍竹更新生长的质量指标。强度干扰下拐棍竹的基径和株高都显著小于中度和轻度干扰,中度干扰样方的基径与轻度干扰差异不显著,但株高显著矮于轻度干扰样方。表明强度干扰下拐棍竹的质量最差,中度干扰的基径已基本得到恢复,但株高仍受到抑制。(4)地震不同干扰等级拐棍竹退笋原因划分为:外力机械损伤、动物取食、生长停止、其他4种。对地震不同干扰等级拐棍竹退笋原因比率的分析发现,强度干扰样方因外力机械损伤和生长停止退笋的比率显著高于中度和轻度干扰样方,而因动物取食退笋的比率显著低于中度和轻度干扰样方。证明强度干扰样方的竹笋更容易受到外力机械损伤和生长停止退笋,而中度和轻度干扰的竹笋更倾向于动物取食致退,揭示了拐棍竹的生长是与震后不同程度的干扰相适应的。     

竹子分子生物学研究进展(英文)

对2003年以来的竹子分子生物学研究进展进行了综述,包括现代分子手段在竹子分类学研究中的开发与应用,鞭芽发育、快速生长、开花、抗逆等相关的重要功能基因研究,基因组测序和转录组测序,遗传转化体系的建立等。这些为今后竹子生物学的研究提供了依据。     

Population ecology in natural<Emphasis Type="Italic">Syneilesis palmata</Emphasis> stands: I. relationship between flowering and plant size

In this study, the relationship was explored between flowering potential and plant size in a population ofSyneilesis palmata (THUNB.) MAX., Usannamul. For this perennial shade-tolerant herb, the flowering rate, plant size, and changes that occurred in the next year after flowering were monitored in two different types of natural stands: moderately shaded (MS; for 11 years) and severely shaded (SS; 7 years). The correlation coefficients (CC) between inflorescence number and other factors, i.e., widths of the first leaf, second leaf, or total leaf breadth, were significant at the 1% level in both stands. In particular, the CC was >0.9 between the number of inflorescences and total leaf breadth. Although the mean flowering rates averaged 6.8% in MS and 8.6% in SS, those values varied widely, ranging from 0 to 53% among years. These rates were not related to petiole length but did increase in proportion to the width of the leaf blade. Flowering began when plants moved beyond the medium size class; the rate in the MS environment was 100% for the largest-sized plants. For plants in the SS, however, the highest rate was 35%. Over the 11 years, 62.8% of all plants in the MS never flowered, while another 30.8% bloomed only once. In comparison, plants observed for 7 years in the SS showed a non-flowering rate of 78.1% and a one-time-only rate of 19.2%. In the next year after flowering, 57.6% of the MS plants exhibited reduced growth while 25.8% of those individuals each split into two plants. In the SS, 37.8% had decreased development the following year, and a mortality rate of 37.8%.