西双版纳地区鞘花的繁殖物候特征及影响因素

桑寄生植物作为关键种和关键性食物资源在生态系统中扮演着重要角色,其鞘花的繁殖物候特征不仅会影响自身的繁殖适合度而且还会影响依赖于其获取食物资源的动物。为了解鞘花的繁殖物候特征及其影响因素,探究其与寄主植物和种子散布者之间的相互作用关系。该研究以西双版纳地区分布的鞘花为对象,通过定期观测鞘花和其寄主植物木荷的繁殖物候,测量它们的生物学特性和温湿度等环境因子,并分析鞘花在个体水平和种群水平上的繁殖物候特征以及寄主植物和温湿度对其繁殖物候的影响。结果表明:(1)鞘花的开花物候属于集中大量开花模式,整个种群的花期和果期的持续时间分别约为20 d和72 d,花期和果期的同步性指数都较高,6月中旬鞘花果实被取食的数量和速率最大,之后逐渐降低。(2)鞘花的始花期与木荷的始花期相关性较高,花期和果期与木荷的繁殖物候基本重叠。(3)每月开花和果熟的个体数量与同期和前一个月的平均温度和相对湿度的相关关系均不显著。总之,桑寄生植物的繁殖物候特征可能受到较多因素的影响,若想全面了解半寄生植物的繁殖物候特征,就要综合考虑生物和非生物等多种因子的共同作用。     

荒漠孑遗植物裸果木的开花物候特征

2014年和2015年连续2年对哈密和乌恰裸果木(Gymnocarpos przewalskii)自然种群的开花物候特征进行野外定点观测,并运用相对开花振幅、开花强度和开花同步性等参数对其开花物候特征进行分析。结果表明,2个种群每年均开1次花,4月下旬到5月上旬进入始花期,开花进程基本相似,均呈单峰曲线,具有较高的开花同步性,呈现"集中大量"开花模式;在个体水平上,始花期、终花期在种群和年份间略有差异;单花持续时间在年际间是稳定的,但在种群间有所不同,即乌恰种群的单花持续时间略长于哈密种群;不同裸果木种群的开花数与结籽数间均呈极显著正相关,开花同步性指数与相对开花强度间呈显著负相关;乌恰种群的始花期与开花数间呈负相关。裸果木个体开花振幅、开花同步指数及开花模式在种群和年份间的相似性很大程度上是由其在长期的地理变迁中形成的遗传特性决定的,而始花期和开花持续时间等开花物候特征的差异主要是由环境和气象因子(海拔、气温、降水及光照)差异造成。研究结果对于探讨该属植物的繁殖生物学特性及其保护具有重要意义。     

中国特有植物血水草开花物候与生殖特性

于2008年3-5月对分布在井冈山的血水草(Eomecon chionantha Hance )5个自然种群的开花物候进行了观察,运用开花振幅、相对开花强度和开花同步性等指数研究了其开花物候特征及其对该种生殖成功的影响.结果表明:血水草开花时间为3月下旬-5月上旬,种群花期历时24 ～46 d,个体平均开花持续时间为11～21 d,单花花期一般为3～5d;井冈山血水草种群的开花物候进程呈单峰曲线模式具有一个开花高峰期,表现出一种“集中开花模式”;与大多数亚热带植物一样,血水草具有较低的相对开花强度,分布频率集中在10％ ～30％.开花物候指数与生殖间的相关分析结果表明:始花时间与花期持续时间呈显著负相关,而与开花数和坐果率呈显著正相关;花期持续时间与开花数和坐果率呈显著正相关;同步性指数与始花时间、开花数、花期持续时间呈负相关.血水草“集中开花模式”是其在长期的进化过程中适应周围气候条件及生境的一种生殖保障.     

巫山淫羊藿开花物候与生殖特征

研究巫山淫羊藿(Epimedium wushanense)3个种群的开花物候进程和生殖特征,分析了不同种群中的花部特征、开花物候指数和结实率差异,以及开花物候指数对巫山淫羊藿生殖成功的影响。结果表明:巫山淫羊藿花期为3月初到4月中下旬,种群开花一般历时22～27d;单株花期(花序)约12～17d;单花花期一般为3～4d。3个种群的个体水平的开花振幅曲线呈单峰型曲线,均具有较高的开花同步指数,表现出一种集中开花式样。相关分析表明:坐果数与开花高峰期、终花期、花期持续时间和花蕾数均呈极显著正相关,但坐果数与始花期无相关关系。开花物候变异系数分析表明:3个种群中花期持续时间、单花持续时间、终花期、开花高峰期和坐果数差异极显著;总花数、开花振幅和结实率差异显著;开花同步指数和始花期无显著差异性。3个种群中除同步性指数外,其余指数的变异系数均有差异,此外,巫山淫羊藿中结实率受到距的长短的影响。表明巫山淫羊藿的开花物候和生殖特征受到微环境的影响,同时生殖特征还受到花部特征距的影响。     

准噶尔无叶豆的开花物候与生殖特征

 为了研究沙漠稀有植物准噶尔无叶豆(Eremosparton songoricum)的开花物候特征及其对生殖成功的影响, 2005和2006年连续两年对其自然种群的开花物候和开花过程中的花部表型变化进行了观察, 并运用相对开花振幅、开花强度和开花同步性等开花物候指数研究了开花物候特征。结果表明: 准噶尔无叶豆在5月下旬至6月中下旬开花, 其种群、个体、花序和单花的花期分别历时26~29 d、8~10 d、5~7 d和2~3 d。单花开花进程依其形态和散粉特征可分为散粉前期、散粉初期、散粉盛期和凋谢期4个时期。其个体水平的开花物候进程(开花振幅曲线)呈渐进式单峰曲线, 具有很高的开花同步指数, 表现出一种集中开花的模式。开花物候指数与座果数之间的相关分析结果表明, 始花日期与花期持续时间存在负相关关系, 而与开花数和座果数存在正相关关系; 花期持续时间与开花数和座果数存在显著正相关关系。准噶尔无叶豆个体开花物候在很大程度上是由其遗传因子决定的, 而开花物候在年度间的变异, 可能是由于荒漠气候的差异(主要是水分和温度的差异)所引起的。作为沙漠窄域分布特有种, 准噶尔无叶豆在环境和人为干扰的双重选择压力下, 为了吸引更多的传粉者访问而达到生殖成功, 形成了大量集中开花的模式。     

科尔沁沙地两种蒿属固沙半灌木的开花物候比较研究

对科尔沁沙地的乌丹蒿和差不嘎蒿的开花物候进行了调查研究,从头状花序、花序枝、个体以及群体水平比较两种植物的开花物候特性。结果表明:(1)乌丹蒿的群体始花期为6月18日,较差不嘎蒿的早9 d,持续时间为19 d,差不嘎蒿开放持续16 d,两种植物群体花期重叠约10 d。在个体水平上,乌丹蒿始花日期比差不嘎蒿早7 d,单株花期长2 d;(2)乌丹蒿二级花序枝上头状花序数比差不嘎蒿的少,开放持续时间更长。两种植物开花频率均为单峰型,二级花序枝开放持续时间频率均以8 d为最高,其次为9 d,头状花序水平上两者开放持续时间频率较高的均为5 d、4 d、6 d,5 d频率最大;(3)差不嘎蒿二级花序枝上近基部的头状花序先开放,自基部向上的第2和第3个位置最先开放的频率相对为高,分别为12.7%和11.9%。而乌丹蒿则是顶端的头状花序先开放,最先开放的频率高达56.3%;(4)两种植物的开花振幅曲线均呈单峰曲线,其中乌丹蒿开花后第9 d开花量达到高峰,差不嘎蒿开花后第6 d开花量达到高峰。两种植物开花物候的相似性反映出它们的亲缘关系及对共同环境的适应性,不同点尤其是最先开放顺序的差异性,表明了两者长期进化的遗传差异。     

多枝柽柳春夏两季开花物候特征与生殖特性

为了研究多枝柽柳的两季开花物候特征与生殖特性,对分布于古尔班通古特沙漠南缘莫索湾荒漠区的多枝柽柳自然种群各水平的开花物候指数、开花过程中的花部特征变化和结实特性进行了观察和统计。结果表明:(1)多枝柽柳春花期为5月下旬至6月上旬,表现为"集中开花模式",夏花期为6月中旬至9月上旬,表现为"持续开花模式"。(2)多枝柽柳春、夏花期的群体、个体、花序和单花水平的花期持续时间分别为14d、10d、3d、1d和85d、77d、2d、1d。(3)花序水平上夏花期始花日期、开花持续时间、开花数和座果数两两之间均存在显著相关关系。(4)两季花期间单花水平的开花进程与形态结构没有明显差异。(5)春花期的结实率显著高于夏花期,但两季花期结籽率差异不显著。(6)春花期昆虫访花次数显著高于夏花期。(7)两季花期的花粉活力没有明显差异,春花期的柱头可授性持续时间比夏花期长。研究认为,古尔班通古特沙漠南缘莫索湾荒漠区的生境条件恶劣,多枝柽柳特殊的春、夏两季开花结实特性在一定程度上提高了其生殖成功的可能性,是对特殊生境的适应性表现。     

大花百子莲的开花物候与生殖特性

连续2年对引种植物大花百子莲在中国上海市人工种群的开花物候、结实情况进行观测记录,从不同水平分析了大花百子莲的开花物候对其生殖的影响。结果显示:（1）大花百子莲在引种地的开花时间为6月初至7月末;种群花期历时40 d左右,单株花期在14～31 d波动,单花寿命主要为2～3 d;2年的开花振幅平均在6朵·株^-1·d^-1左右,花朵开放百分率值在3.69%~4.87%左右,平均开花式样为11~17朵。（2）大花百子莲种群内2007年比2008年的各项开花物候指标均提前6～9 d;种群开花进程为渐进式的双峰曲线;种群花期重叠时间较长,2年的平均开花同步指数分别为0.663、0.695;植株个体相对开花强度主要分布频度为50%～70%。（3）开花物候指数与座果率的相关分析表明,始花时间与开花数目、座果数、花期长度呈显著负相关关系,开花数目与座果数、花期长度以及花期长度与座果数均呈显著正相关关系,同一花序内单花所发育成果实的数量与单花开放时间呈极显著负相关关系。研究表明,大花百子莲始花时间早的植株比始花时间晚的植株花期更长,开花数目多的植株比开花数目少的植株花期更长,花期长的植株比花期短的植株座果率更高;而且在同一花序上,越早开放的单花其结实的可能性越大。作为引种植物,大花百子莲开花物候年度间具有相似性,说明其在引种地可以适应环境压力,形成稳定的开花时间与开花模式;较长的花期重叠和大量的开花数目可以吸引较多的传粉者访问,为其生殖成功奠定了良好基础。     

濒危植物长柄双花木开花物候与生殖特性

研究了长柄双花木开花过程中花部表型的变化。连续4a对其野生种群、1a对人工种群的开花物候进行了观察,并运用相对开花强度和同步性等开花物候指数分析了开花物候对其生殖的影响。结果如下：长柄双花木开花时间为9月上、中旬至11月中、下旬;单花花期一般为6～7d,单花依其形态和散粉特征可以分为4个时期：散粉前期、散粉初期、散粉盛期和凋谢期。个体开花持续时间49～55d,种群花期历时63～71d。种群内不同年度间开花物候指数没有显著差异,而种群间则存在显著差异,野生种群开花进程为渐进式单峰曲线。人工种群则为“钟”形曲线,二者均属于“集中开花模式”。长柄双花木具有2个相对开花强度的分异趋势,这种分异趋势具有进化意义。开花物候指数与生殖间的相关分析表明,始花时间与开花数量、座果率及花期长度之间均具显著负相关关系,而开花数量与花期长度之间则呈显著正相关,但均为线性相关。长柄双花木开花物候在种群间的差异和种群内年度间的相似性说明,其开花时间可能是由与其相关的复杂的微生境特征和(或)由其遗传因子决定的,同时也反映了种群间的遗传分异和种群内个体间的遗传一致性。作为一种濒危物种,长柄双花木在这种环境的选择压力之下,形成了“大量、集中开放的花”的开花模式,吸引到更多的传粉者的访问,从而达到生殖成功。     

二型花柱植株金荞麦繁殖特征

开花物候及繁殖分配是植物适应环境的重要因素。对金荞麦开花物候、繁殖分配及策略进行了研究。结果如下:金荞麦的花果期为每年的8—11月,9月集中开花,其集中开花模式有助于吸引昆虫传粉,提高繁殖成功率;金荞麦单花开花持续时间为1—2 d,种群花期均为85d。L型花序花期为15—26d,S型花序花期为14—27d,两者没有显著差异;L型单花序开花数为26—131朵,S型单花序开花数为36—147朵,两者没有显著差异。L型和S型花序开花动态呈现单峰曲线,在花序开花后第11天L型和S型都达到最大值,分别为7.30%和7.20%,且两种花型具有较高的开花同步性,这有助于其繁殖适应性的提高。同一个花型中,雌蕊长、雄蕊长之间存在极显著负相关,但雌雄总长不存在显著差异,表明雌蕊长、雄蕊长可能存在权衡关系;金荞麦的繁殖器官和营养器官生物量在L型和S型间不存在显著差异,但其花生物量与植株生物量表现出极显著正相关关系。金荞麦L型花生物量分配极显著大于S型,而总生物量不存在显著差异,说明金荞麦植株的营养生长与有性繁殖间存在权衡关系。     

Linking flowering and reproductive allocation in response to nitrogen addition in an alpine meadow

Aims Plants can change in phenology and biomass allocation in response to environmental change. It has been demonstrated that nitrogen is the most limiting resource for plants in many terrestrial ecosystems. Previous studies have usually focused on either flowering phenology or biomass allocation of plants in response to nitrogen addition; however, attempts to link flowering phenology and biomass allocation are still rare. In this study, we tested the effects of nitrogen addition on both flowering phenology and reproductive allocation in 34 common species. We also examined the potential linkage between flowering time and reproductive allocation in response to nitrogen addition.Methods We conducted a 3-year nitrogen addition experiment in Tibetan alpine meadow. We measured first flowering date and the reproductive allocation for 34 common plant species in control, low and high nitrogen added plots, respectively. One-way analysis of variance was used to examine differences of first flowering date and reproductive allocation among treatments. The relationships between the change in species first flowering date and change in reproductive allocation in response to nitrogen addition were examined by calculating Pearson correlation coefficients.Important findings For most species, both first flowering date and reproductive allocation significantly responded to nitrogen addition. Nitrogen addition significantly delayed the first flowering date and reduced the reproductive allocation for all graminoid species, but accelerated flowering and increased reproductive allocation for most forb species. We found that changes in first flowering date significantly negatively correlated with the changes in reproductive allocation over species in response to nitrogen, which indicated a positive relationship between flowering response and plant performance in reproductive allocation. Species that advanced their flowering time with nitrogen addition increased their reproductive allocation, whereas those that delayed flowering time tended to decline in reproductive allocation with nitrogen addition. Our results suggest that species-specific switch from vegetative growth to reproductive growth could influence species performance.     

Morphological changes and resource allocation of Zizania latifolia (Griseb.) Stapf in response to different submergence depth and duration

Plants around ponds, rivers and lakes are subjected to long-term partial or complete submergence. When they are flooded, water level affects the plants simultaneously with duration of submergence. Separate and interactive effects of water level and duration on the growth of the herbaceous perennial Zizania latifolia (Poaceae) were investigated by exposing the plants in greenhouse water tanks to submergence in different water depths and for different time-spans. The plants exhibited great shoot elongation upon submergence and prolonged flood duration, and the basal tiller number of the species decreased with higher water levels. Submergence treatment advanced the flowering date and increased the inflorescence number. Plant total biomass did not differ among all the treatments, while the root:shoot ratio decreased with increased water level, prolonged duration of submergence and their interaction. The high plasticity in morphology and shifts in reproductive strategy and biomass allocation enabled the Zizania plants to survive the compound effect of flooding height and duration. This may explain the occurrence of this species in habitats subjected to long-term flooding. The results obtained in this experiment will contribute to understanding the impact of flooding dynamics on plants and the ways of adaptation responses to prolonged waterlogging.     

Male-biased sex allocation in late-blooming flowers driven by resource limitation in the clonal perennial Aconitum kusnezoffii (Ranunculaceae)

Flowering phenology and clonal growth are known to affect resource and pollen availability, and therefore select for adaptive or constrained sex allocation strategies to some degree. However, the consequences of temporal sex allocation patterns for reproductive fitness across the flower, inflorescence, and genet levels have rarely been examined. Moreover, experimental tests of the underlying regulatory mechanisms are scarce. We examined the association of flowering phenology and inflorescence position with temporal sex allocation and reproductive success in the protandrous perennial clonal herb, Aconitum kusnezoffii, over four consecutive growing seasons by examining more than 39 000 flowers. We also conducted controlled experiments to test the effects of resource and pollen limitation on the female reproductive success of lateral inflorescences. We found that some male functions were positively correlated with flowering phenology, whereas female reproductive success was negatively correlated with flowering phenology and inflorescence position. Lateral inflorescences invested more in male function than terminal inflorescences and therefore yielded fewer and smaller seeds. Resource limitation may serve as the key mechanism underlying this differentiated pattern. Decreased female reproductive success was consistently observed at the flower and inflorescence levels as flowering occurred later in the growth season. Late-blooming lateral inflorescences specialized in the male function, and their female reproductive success was constrained by early-blooming terminal inflorescences. This might be the first attempt to systematically demonstrate sex allocation strategy differentiation in a protandrous plant species at the inflorescence level. In addition, our study provides empirical evidence of dichogamy selecting for specialized sex allocation strategies among inflorescences.     

常绿阔叶树种栲树开花物候动态及花的空间配置

 基于定株观测和随机枝取样法,对浙江天童常绿阔叶林内栲树（Castanopsis fargesii）的开花物候动态及其雌花、雄花的空间配置进行了研究。结果表明：在栲树的生殖枝上,并非所有的芽都分化、萌发生成花序,栲树花芽的分化和发育集中在一级生殖枝上。生殖枝上花芽的分化与该枝的空间位置密切相关。栲树花期明显晚于春季的展叶期,与叶片生长时间重叠。盛花期集中于5月下旬,约持续8 d左右,属于同步发生的花期。栲树雄花序的数量明显高于雌花序,雄花序约占花序总数的77.88%,雌花序仅占22.12%。大量雄花和花粉的存在是保证雌花接受花粉和完成受精的基础。花序在植冠层中的空间配置明显不同：在同一植冠内,向阳面和背阴面生殖枝上芽萌发成花序的比率存在明显差异(p<0.01),阳面生殖枝上顶芽萌发成花序的比率高于阴面生殖枝的比率,并且,阳面的每个生殖枝上平均花序数和雄花数量均高于阴面生殖枝,花序的分化和发育与枝系的生长发育状况有密切关系。     

雌雄异熟植物露蕊乌头开花时间对雌雄功能期及表型性别的影响

开花时间决定了植物雌雄功能的交配机会, 最终影响繁殖成功。交配环境假说认为雌雄异熟植物开花时间的差异能引起植物表型性别的变异, 改变种群内的交配环境, 影响植物对雌雄功能的最佳性分配。为了研究开花时间对雌雄异熟植物的雌雄性别时期及表型性别的影响, 本文以毛茛科雄性先熟植物露蕊乌头(Aconitum gymnandrum)为实验材料, 记录了雄性和雌性功能期, 分析了植株开花时间、花的雌雄功能期和表型性别的关系。结果表明: 在植物同一花序内, 较晚开放的花有更长的雄性期和更短的雌性期, 性分配在时间上偏雄。雌雄功能期在时间上的相对分配随植物开花时间的变化表现出相似的趋势： 较晚开的花或较晚开花的个体, 花的雄性功能期相对于雌性功能期更长, 在时间上更偏向雄性功能。而且, 开花时间的差异影响种群内花的性比和植物个体的表型性别动态。随着开花时间由早到晚的变化, 种群内早期以雄花为主,末期以雌花为主, 种群内性别环境由偏雄向偏雌变化, 因此植株个体的平均表型性别则从偏雌转向偏雄。本文结果支持交配环境假说, 雄性先熟的露蕊乌头开花早期, 种群内花的性别比偏雄, 种群表型性别环境偏雄, 因而植物个体平均表型性别偏雌, 性别分配(即时间分配)偏向雌性功能, 而晚开花个体的平均性别偏雄, 更偏向雄性功能的分配。     

三峡库区两种耐水淹植物的存活率和碳水化合物储备关系

野古草和秋花柳是三峡库区消落带两种强水淹耐受能力的植物物种。以往研究显示植物的水淹耐受性和体内碳水化合物储备有关。为了探明野古草和秋花柳水淹下的高存活率是否和碳水化合物储备有关, 研究了在室外6个月的模拟水淹条件下两个物种在不同水淹时间(40、90、120和180d)和不同水淹深度下(不水淹、根部水淹和完全淹没)的生物量积累、存活率和碳水化合物含量和分布。结果表明: (1)野古草和秋花柳对长期水淹具有很高的耐受性, 根部水淹植物6个月处理后完全存活; 而完全淹没条件下, 野古草仅在4个月, 秋花柳仅在6个月处理后才开始死亡; (2)碳水化合物主要储备在野古草的茎和秋花柳的茎与主根中, 野古草的根和秋华柳的细根中碳水化合物含量很低; (3)水淹深度和水淹时间对植物生物量积累和碳水化合物含量影响显著(P 0.05):与未水淹植株相比, 根部水淹仅略微降低了生物量积累以及可溶性糖和淀粉含量 (P 0.05), 且保持基本稳定或增加的趋势, 而完全淹没的植株生物量随水淹时间逐渐降低, 碳水化合物含量在前90天快速下降 (P 0.05), 之后缓慢下降或保持不变。研究结果表明, 野古草和秋花柳强的水淹耐受性是和它们高的碳水化合物储备以及水淹条件下对碳水化合物的动用能力有关, 后期的死亡率增加与碳水化合物储备消耗殆尽有关, 野古草和秋花柳对碳水化合物储备对水淹的响应的差异可能和它们的碳水化合物储备在不同组织中的分配模式有关。       

Effects of enhanced UV-B radiation on fitness of an alpine species Cerastium glomeratum Thuill

Aims Information about how species respond to extreme environments, such as high UV-B radiation, is very useful in estimating natural ecosystem structure and functions in alpine areas. Our aim is to examine the effect of enhanced UV-B radiation on the fitness of an alpine meadow annual species on Qinghai-Tibet Plateau.Methods Plants of Cerastium glomeratum Thuill. were exposed to ambient (control) or ambient plus supplemental UV-B radiation (enhanced), simulating a 9% ozone depletion over Gannan, China (102°53′E, 34°55′N, 2900 m in altitude), up to leaf senescence and fruit maturation. Plant height, flower phenology, biomass allocation and reproductive parameters of the species were measured.Important findings Plant height in C. glomeratum was reduced by enhanced UV-B radiation at early growth stages and compensated with ongoing development. Fruit biomass, aboveground biomass, total biomass and reproductive effort (fruit dry mass/aboveground biomass) were not affected by enhanced UV-B radiation, but a significant increase in root/shoot ratio was found. Enhanced UV-B radiation delayed onset of flowering by 1 day and shortened duration of flowering by 5 days in C. glomeratum. But because of the long period of flowering time (83–88 days), this did not make any significant effect on flower number, seed number, pollination success (number of seeds per fruit) or reproductive success (fruit to flower ratio) in C. glomeratum. Enhanced UV-B radiation had no effect on seed germination and seed mass either. And the high production and low germination rate of the seed might be the strategy of C. glomeratum to survive the extreme environments on alpine meadow. All these results showed that C. glomeratum was tolerant to enhanced UV-B radiation.     

Interactive Effects of Arbuscular Mycorrhizal Fungi and Copper Stress on Flowering Phenology and Reproduction of Elsholtzia splendens

Plant responses to heavy metal contamination may depend on the presence of arbuscular mycorrhizal fungi (AMF). Elsholtzia splendens is an indicator species for the presence of copper (Cu) mines because both its flowering phenology and reproduction are tolerant to heavy metals. To test whether effects of Cu on the flowering phenology and reproduction of E. splendens depend on the presence of AMF, we conducted a factorial experiment with two Cu treatments (with or without Cu addition) crossed with two AMF treatments (with or without AMF inoculation). Without AMF, Cu addition significantly delayed the onset dates, ending dates and peak dates of flowering and decreased flowering duration. However, AMF inoculation reversed the effects of Cu stress, with recovered flowering onset and ending dates and increased the flowering duration. Cu addition significantly decreased inflorescence width and number, inflorescence biomass, vegetative biomass and total seed number, but significantly increased 1000-seed weight. AMF inoculation significantly increased vegetative biomass. Two-way ANOVA results showed that the interactive effects between Cu addition and AMF inoculation were significant on the inflorescence number, vegetative biomass and total seed number. These results indicate that AMF can alleviate the Cu stress on the flowering phenology and reproduction of E. splendens.