光温耦合的中国温带地区旱柳花期时空格局模拟

建立基于温度和光周期驱动的旱柳花期物候模型,旨在寻找影响旱柳花期时空变化的主要气象因子,揭示调控植物开花时间的生态机制,还可为改善柳絮造成的环境污染和花粉过敏等人类健康问题提供参考信息和依据。利用中国气象局农业气象观测网提供的中国温带地区1982-2011年49个站点的旱柳开花始期、盛期和末期观测资料及平行的逐日气象数据,分别对6种模型（简单积温模型、温度三基点模型、八时段温度模型、简单积温-日长模型、温度三基点-日长模型和八时段温度-日长模型）进行了参数率定和假设检验,根据外部检验结果,从中选出针对旱柳3个花期的最优物候模型,进而利用连续地理气象数据和最优物候模型重建了1982-2011年旱柳开花始期、盛期、末期和花期长度的时空变化特征。结果表明：光温耦合的物候模型对旱柳花期的模拟效果和外推效果优于仅基于温度的模型。旱柳开花始期和盛期最优模型均为八时段温度-日长模型,末期为温度三基点-日长模型,说明光周期和温度可能是影响旱柳花期开始、繁盛和结束时间的主要气象因子。同时,优选出的物候模型能够较准确地对不同年份和不同地区的旱柳花期进行模拟及预测。重建的1982-2011年旱柳平均开花始期、盛期和末期日期分别为4月24日、4月28日和5月3日,平均花期长度为9 d,始期、盛期和末期出现日期呈现出从海拔低到高、从南向北、从西向东逐渐推迟的空间格局。1982-2011年旱柳开花始期、盛期和末期在大部分地区呈提前趋势,呈显著提前趋势的面积分别占总面积的49.78%、50.01%和53.40%,花期长度变化差异不显著。     

Herbarium records identify sensitivity of flowering phenology of eucalypts to climate: Implications for species response to climate change

Flowering phenology is very sensitive to climate and with increasing global warming the flowering time of plants is shifting to earlier or later dates. Changes in flowering times may affect species reproductive success, associated phenological events, species synchrony, and community composition. Long‐term data on phenological events can provide key insights into the impacts of climate on phenology. For Australia, however, limited data availability restricts our ability to assess the impacts of climate change on plant phenology. To address this limitation other data sources must be explored such as the use of herbarium specimens to conduct studies on flowering phenology. This study uses herbarium specimens for investigating the flowering phenology of five dominant and commercially important Eucalyptus species of south‐eastern Australia and the consequences of climate variability and change on flowering phenology. Relative to precipitation and air humidity, mean temperature of the preceding 3 months was the most influential factor on the flowering time for all species. In response to a temperature increment of 1°C, a shift in the timing of flowering of 14.1–14.9 days was predicted for E. microcarpa and E. tricarpa while delays in flowering of 11.3–15.5 days were found for E. obliqua, E. radiata and E. polyanthemos. Eucalyptus polyanthemos exhibited the greatest sensitivity to climatic variables. The study demonstrates that herbarium data can be used to detect climatic signals on flowering phenology for species with a long flowering duration, such as eucalypts. The robust relationship identified between temperature and flowering phenology indicates that shifts in flowering times will occur under predicted climate change which may affect reproductive success, fitness, plant communities and ecosystems.     

Evidences of olive pollination date variations in relation to spring temperature trends

Airborne pollen concentration patterns reflect flowering phenology of a given species, and it may be a sensitive regional indicator in climate change studies. This paper presents the relationship between a strategic biological event, such as olive flowering, and the air temperature trend, registered over a large scale (1982–2007) in the Umbria region. The aim of the study was to determine relationships between phenological behaviour (flowering) of olive trees and the air temperature trend (1982–2007) in the Umbria region. The phenological data on flowering phase were registered indirectly through an aerobiological monitoring technique. The obtained results showed a strong relationship between phenology and thermal trend. This characteristic was confirmed from results of correlations between temperature (mean temperature from 1st March) and flowering dates, especially that of full flowering (r = −0.9297). Moreover, the results showed an advance trend of 6, 8 and 10 days, respectively of start, full and end of flowering dates. The advance of the recorded flowering time in this period is to ascribe mainly to the increase of mean temperature and above all to that registered in months of May and June.     

Clarifying springtime temperature reconstructions of the medieval period by gap-filling the cherry blossom phenological data series at Kyoto, Japan

We investigated documents and diaries from the ninth to the fourteenth centuries to supplement the phenological data series of the flowering of Japanese cherry (Prunus jamasakura) in Kyoto, Japan, to improve and fill gaps in temperature estimates based on previously reported phenological data. We then reconstructed a nearly continuous series of March mean temperatures based on 224 years of cherry flowering data, including 51 years of previously unused data, to clarify springtime climate changes. We also attempted to estimate cherry full-flowering dates from phenological records of other deciduous species, adding further data for 6 years in the tenth and eleventh centuries by using the flowering phenology of Japanese wisteria (Wisteria floribunda). The reconstructed tenth century March mean temperatures were around 7°C, indicating warmer conditions than at present. Temperatures then fell until the 1180s, recovered gradually until the 1310s, and then declined again in the mid-fourteenth century.     

Statistical downscaling of monthly mean air temperature to the beginning of flowering of Galanthus nivalis L. in Northern Germany

 We have examined the relationship between phenological data and concurrent large-scale meterological data. As phenological data we have chosen the beginning of the flowering of Galanthus nivalis L. (flowering date) in Northern Germany, and as large-scale meteorological data we use monthly mean near-surface air temperatures for January, February and March. By means of canonical correlation analysis (CCA), a strong linear correlation between both sets of variables is identified. Twenty years of observed data are used to build the statistical model. To validate the derived relationship, the flowering date is downscaled from air temperature observations of an independent period. The statistical model is found to reproduce the observed flowering dates well, both in terms of variability as well as amplitude. Air temperature data from a general circulation model of climate change are used to estimate the flowering date in the case of increasing atmospheric carbon dioxide concentration. We found that at a time of doubled CO₂ concentration (expected by about 2035) G. nivalis L. in Northern Germany will flower ∼2 weeks and at the time of tripled CO₂ concentration (expected by about 2085) ∼4 weeks earlier than presently. Received: 7 August 1996 / Accepted: 27 November 1996     

Effect of climatic change on the duration of the breeding season in three European thrushes

Capsule Evidence for extension of breeding seasons in Song Thrushes and Blackbirds (multiple‐brooded species) and shortening in Fieldfares (a single‐brooded species).

Aims To analyse breeding data from central Europe during 1964–2006 in relation to climatic conditions operating at breeding and wintering grounds, and to compare breeding phenology in urban and rural habitats.

Methods Using chick ringing data we evaluated long‐term changes in breeding phenology of Fieldfares Turdus pilaris, Song Thrushes T. philomelos and Common Blackbirds T. merula. Changes in phenology were analysed for the 5th percentile, median, 95th percentile, inter‐quantile range (IQR) and brood size.

Results All thrushes showed consistent trends towards earlier onset of breeding. However, there was a contrasting pattern in the later phenological characteristics (median, 95th percentile and IQR) that were associated with single‐ versus multiple‐brooded species. The single‐brooded Fieldfares revealed an advancing trend in all phenological phases, which caused shortening of its breeding period. In contrast, the multiple‐brooded Song Thrushes and Blackbirds delayed both medians and the 95th percentiles of breeding dates leading to extension of their breeding seasons. Temperatures at both wintering and breeding grounds were generally negatively associated with the onset of breeding. Urban birds bred significantly earlier than their rural conspecifics, but brood sizes did not differ.

Conclusion Multiple‐brooded species may respond differently to increasing spring temperatures than single‐brooded species.     

Scaling phenology from the local to the regional level: advances from species‐specific phenological models

Plant phenology, the study of seasonal plant activity driven by environmental factors, has found a renewal in the context of global climate change. Phenological events, such as leaf unfolding, exert strong control over seasonal exchanges of matter and energy between the land surface and the atmosphere. Phenological models that simulate the start of the growing season should be efficient tools to predict vegetation responses to climatic changes and related changes in energy balance. Species‐specific phenological models developed in the eighties have not been used for global‐scale predictions because their predictions were inaccurate in external conditions. Recent advances in phenology modelling at the species level suggest that prediction at a large scale may now be possible. In the present study, we tested the performance of species‐specific phenological models in time and space, looking at their ability (i) to predict regional phenology when previously fitted at a local scale, and (ii) to predict phenological trends, linked to climate changes, observed over a long‐term. For that task we used an historical phenological dataset from Ohio from the late ninetieth century and an airborne pollen dataset from Ontario, Québec and Maryland from the late twentieth century. The results show that the species‐specific phenological models used in this study were able to predict regional phenology even though they were fitted locally. The reconstruction of a phenological time series over the twentieth century showed a significant advancement of 0.2 days per year in the date of flowering of Ulmus americana, but very weak trends for Fraxinus americana and Quercus velutina.     

Integration of flowering dates in phenology and pollen counts in aerobiology: analysis of their spatial and temporal coherence in Germany (1992–1999)

We studied the possibility of integrating flowering dates in phenology and pollen counts in aerobiology in Germany. Data were analyzed for three pollen types (Betula, Poaceae, Artemisia) at 51 stations with pollen traps, and corresponding phenological flowering dates for 400 adjacent stations (< 25 km) for the years 1992–1993 and 1997–1999. The spatial and temporal coherence of these data sets was investigated by comparing start and peak of the pollen season with local minima and means of plant flowering. Our study revealed that start of birch pollen season occurred on average 5.7 days earlier than local birch flowering. For mugwort and grass, the pollen season started on average after local flowering was observed; mugwort pollen was found 4.8 days later and grass pollen season started almost on the same day (0.6 days later) as local flowering. Whereas the peak of the birch pollen season coincided with the mean flowering dates (0.4 days later), the pollen peaks of the other two species took place much later. On average, the peak of mugwort pollen occurred 15.4 days later than mean local flowering, the peak of grass pollen catches followed 22.6 days after local flowering. The study revealed a great temporal divergence between pollen and flowering dates with an irregular spatial pattern across Germany. Not all pollen catches could be explained by local vegetation flowering. Possible reasons include long-distance transport, pollen contributions of other than phenologically observed species and methodological constraints. The results suggest that further research is needed before using flowering dates in phenology to extrapolate pollen counts.     

Phenological behaviour of Parthenium hysterophorus in response to climatic variations according to the extended BBCH scale

Considering the importance of ecological and biological traits in imparting invasive success to the alien species, the phenological behaviour of an alien invasive weed Parthenium hysterophorus was documented according to the extended BBCH scale in four different seasons. A phenological calendar was prepared using both two‐ and three‐ digit coding system, precisely describing the developmental stages of the weed. The phenological documentation is further supplemented with the dates corresponding to a particular growth stage, pictures of the representative growth stages and meteorological data of all the four seasons. Results revealed that the phenology of the weed altered in response to the changing temperature and humidity conditions but no apparent climatic condition could inhibit its germination or flowering. However, the emergence of inflorescence was highly sensitive to the temperature/photoperiodic conditions. Variations in the phenological traits of P. hysterophorus with changing environmental conditions explain the acclimatisation potential of the weed permitting its vast spread in the non‐native regions. Since the given phenological illustrations are accurate, unambiguous and coded as per an internationally recognised scale, they could be exploited for agronomic practices, weed management programmes, and research purposes.     

Non-linear regression models for time to flowering in wild chickpea combine genetic and climatic factors

Background

Accurate prediction of crop flowering time is required for reaching maximal farm efficiency. Several models developed to accomplish this goal are based on deep knowledge of plant phenology, requiring large investment for every individual crop or new variety. Mathematical modeling can be used to make better use of more shallow data and to extract information from it with higher efficiency. Cultivars of chickpea, Cicer arietanum, are currently being improved by introgressing wild C. reticulatum biodiversity with very different flowering time requirements. More understanding is required for how flowering time will depend on environmental conditions in these cultivars developed by introgression of wild alleles.

Results

We built a novel model for flowering time of wild chickpeas collected at 21 different sites in Turkey and grown in 4 distinct environmental conditions over several different years and seasons. We propose a general approach, in which the analytic forms of dependence of flowering time on climatic parameters, their regression coefficients, and a set of predictors are inferred automatically by stochastic minimization of the deviation of the model output from data. By using a combination of Grammatical Evolution and Differential Evolution Entirely Parallel method, we have identified a model that reflects the influence of effects of day length, temperature, humidity and precipitation and has a coefficient of determination of R²=0.97.

Conclusions

We used our model to test two important hypotheses. We propose that chickpea phenology may be strongly predicted by accession geographic origin, as well as local environmental conditions at the site of growth. Indeed, the site of origin-by-growth environment interaction accounts for about 14.7% of variation in time period from sowing to flowering. Secondly, as the adaptation to specific environments is blueprinted in genomes, the effects of genes on flowering time may be conditioned on environmental factors. Genotype-by-environment interaction accounts for about 17.2% of overall variation in flowering time. We also identified several genomic markers associated with different reactions to climatic factor changes. Our methodology is general and can be further applied to extend existing crop models, especially when phenological information is limited.

     

Regional unified model‐based leaf unfolding prediction from 1960 to 2009 across northern China

Using first leaf unfolding data of Salix matsudana, Populus simonii, Ulmus pumila, and Prunus armeniaca, and daily mean temperature data during the 1981–2005 period at 136 stations in northern China, we fitted unified forcing and chilling phenology models and selected optimum models for each species at each station. Then, we examined performances of each optimum local species‐specific model in predicting leaf unfolding dates at all external stations within the corresponding climate region and selected 16 local species‐specific models with maximum effective predictions as the regional unified models in different climate regions. Furthermore, we validated the regional unified models using leaf unfolding and daily mean temperature data beyond the time period of model fitting. Finally, we substituted gridded daily mean temperature data into the regional unified models, and reconstructed spatial patterns of leaf unfolding dates of the four tree species across northern China during 1960–2009. At local scales, the unified forcing model shows higher simulation efficiency at 83% of data sets, whereas the unified chilling model indicates higher simulation efficiency at 17% of data sets. Thus, winter temperature increase so far has not yet significantly influenced dormancy and consequent leaf development of deciduous trees in most parts of northern China. Spatial and temporal validation confirmed capability and reliability of regional unified species‐specific models in predicting leaf unfolding dates in northern China. Reconstructed leaf unfolding dates of the four tree species show significant advancements by 1.4–1.6 days per decade during 1960–2009 across northern China, which are stronger for the earlier than the later leaf unfolding species. Our findings suggest that the principal characteristics of plant phenology and phenological responses to climate change at regional scales can be captured by phenological and climatic data sets at a few representative locations.     

The strength of assortative mating for flowering date and its basis in individual variation in flowering schedule

Although it has been widely asserted that plants mate assortatively by flowering time, there is virtually no published information on the strength or causes of phenological assortment in natural populations. When strong, assortative mating can accelerate the evolution of plant reproductive phenology through its inflationary effect on genetic variance. We estimated potential assortative mating for flowering date in 31 old‐field species in Ontario, Canada. For each species, we constructed a matrix of pairwise mating probabilities from the individual flowering schedules, that is the number of flower deployed on successive dates. The matrix was used to estimate the phenotypic correlation between mates, ρ, for flowering date. We also developed a measure of flowering synchrony within species, S, based upon the eigenstructure of the mating matrix. The mean correlation between pollen recipients and potential donors for flowering date was  = 0.31 (range: 0.05–0.63). A strong potential for assortative mating was found among species with high variance in flowering date, flowering schedules of short duration and skew towards early flower deployment. Flowering synchrony, S, was negatively correlated with potential assortment (r = ?0.49), but we go on to show that although low synchrony is a necessary condition for phenological assortative mating, it may not be sufficient to induce assortment for a given phenological trait. The potential correlation between mates showed no seasonal trend; thus, as climate change imposes selection on phenology through longer growing seasons, spring‐flowering species are no more likely to experience an accelerated evolutionary response than summer species.     

Small‐scale spatial autocorrelation and the interpretation of relationships between phenological parameters

Question: Are flower production and associated phenological variables (onset, end, duration, and three measures of flowering synchrony) randomly distributed in space or, alternatively, is there a neighbourhood structure (spatial autocorrelation) in the values of these variables? To which extent does spatial autocorrelation affect the correlation tests between phenological traits? Location: A tree savanna reserve in Southeastern Brazil (22°15′S,47°08′W). Methods: The flowering season of Chromolaena odorata was followed for all (96) individuals in a completely mapped area of 3000 m². The phenological traits were estimated by counting flower heads in anthesis on individual plants every seven days for 14 weeks. Results: Flowering time was unimodally distributed, but with different peak dates depending on whether individual flower heads or plants were counted. Three phenological traits and canopy closure above the plants showed some degree of spatial autocorrelation, which caused loss of up to 35% of degrees of freedom in nine of 36 correlation tests. Such a decrease in the degrees of freedom resulted in loss of significance for correlations in three pairs of variables. Conclusions: We hypothesize that the spatial autocorrelation in phenological traits between C. odorata neighbours may be driven by genetic similarity among neighbouring plants and/or spatial structuring of environmental factors. Because location and distance between samples may affect their statistical independence, we suggest that spatial autocorrelation should be taken into account in future studies of plant phenology, e.g. by using effective sample size in statistical tests.     

西安和宝鸡木本植物花期物候变化及温度敏感度对比

植物物候是指示生态系统对气候变化响应的重要证据。已有研究多基于代表性站点的物候观测数据研究物候特征及其对气候变化的响应规律。同一气候区内,不同站点的物候变化及对温度变化响应的敏感度是否一致仍需深入探讨。本文选择同属于暖温带湿润区汾渭平原气候区的西安和宝鸡为研究区,利用"中国物候观测网"在两个站点21个共有物种的开花始期和开花末期数据,比较了1987—2016年两站点各植物花期物候变化特征及其对温度变化响应的敏感度差异。结果表明,西安和宝鸡各物种的开花始期和开花末期均以提前趋势为主。大部分物种开花始期在西安的提前趋势(平均趋势-0.57 d/a)明显强于在宝鸡的提前趋势(平均趋势-0.29 d/a),但开花末期趋势差异不显著。除紫薇和迎春的敏感度差异较大外,其他物种开花始期和开花末期的温度敏感度在两站点间非常接近,无显著差异。由此可见,在同一气候区的不同站点,因增温幅度不同,植物的始花期变化存在较大差异,不能用单站点的物候变化反映整个气候区的物候变化。但同一植物在单站点的温度敏感度可以较好的反映同一气候区其他站点的植物物候-气候关系。本文研究结果可为利用有限站点的物候观测数据分析区域物候变化及对气候变化的响应提供科学依据。     

Changes in the phenology of the ground beetle Pterostichus madidus (Fabricius, 1775)

Abstract A growing body of evidence shows that climate change can alter the phenology of plants and animals. In this study long‐term data from the UK Environmental Change Network (ECN) were analyzed to investigate whether there has been a change in the phenology of the ground beetle Pterostichus madidus (Fabricius, 1775). Pitfall trap data were available from 12 ECN sites across the United Kingdom, most of which have been in operation for more than 15 years. Weather and vegetation datasets were also utilized. Pitfall trap lines were categorized to eight vegetation types. Trend analysis over time was carried out first using all the available dates of capture events, then the datasets grouped by vegetation type and site. Shifts in high‐activity periods were also analyzed. P. madidus appearance dates advanced significantly at seven sites and in five vegetation types. Peak activity advanced at two sites. At one site the timing of activity became significantly later. The last day of activity did not change significantly, supporting the theory that the cessation of the activity period is more likely to be controlled by photoperiod than temperature. The relationships between phenological variables and climatic factors were also investigated. However, no significant correlations were detected. These results demonstrate that between 1992 and 2008, phenology of P. madidus at seven sites from the eight analyzed has changed. Global warming may be driving these changes and future work will investigate underlying processes.     

Comparing fruiting phenology across two historical datasets: Thoreau’s observations and herbarium specimens

Background and AimsFruiting remains under-represented in long-term phenology records, relative to leaf and flower phenology. Herbarium specimens and historical field notes can fill this gap, but selecting and synthesizing these records for modern-day comparison requires an understanding of whether different historical data sources contain similar information, and whether similar, but not equivalent, fruiting metrics are comparable with one another.MethodsFor 67 fleshy-fruited plant species, we compared observations of fruiting phenology made by Henry David Thoreau in Concord, Massachusetts (1850s), with phenology data gathered from herbarium specimens collected across New England (mid-1800s to 2000s). To identify whether fruiting times and the order of fruiting among species are similar between datasets, we compared dates of first, peak and last observed fruiting (recorded by Thoreau), and earliest, mean and latest specimen (collected from herbarium records), as well as fruiting durations.Key ResultsOn average, earliest herbarium specimen dates were earlier than first fruiting dates observed by Thoreau; mean specimen dates were similar to Thoreau’s peak fruiting dates; latest specimen dates were later than Thoreau’s last fruiting dates; and durations of fruiting captured by herbarium specimens were longer than durations of fruiting observed by Thoreau. All metrics of fruiting phenology except duration were significantly, positively correlated within (r: 0.69–0.88) and between (r: 0.59–0.85) datasets.ConclusionsStrong correlations in fruiting phenology between Thoreau’s observations and data from herbaria suggest that field and herbarium methods capture similar broad-scale phenological information, including relative fruiting times among plant species in New England. Differences in the timing of first, last and duration of fruiting suggest that historical datasets collected with different methods, scales and metrics may not be comparable when exact timing is important. Researchers should strongly consider matching methodology when selecting historical records of fruiting phenology for present-day comparisons.     

Density effects of flowering phenology and mating potential in Nicotiana alata

Summary We investigated effects of plant density on floral phenology and potential mating in artificial populations of the outcrossing ornamental Nicotiana alata planted at three densities. Path analysis revealed that increasing plant density yielded significantly earlier peak flowering dates, significantly earlier last flowering dates, and significantly lower plant biomass. Direct effects of density on final flower number were not significant. Variation among replicate plots for first date of flowering was larger than variation among densities, indicating that factors other than density influence floral initiation.We did not record actual mating, but determined from phenological data the number and identity of potential mates. Increased density had several effects on potential mating patterns and on potential N_e, effective population number. At high density, fewer focal plants flowered for shorter durations. This led to less overlap in flowering time among plants, decreasing the number of potential parental combinations possible among the progeny. Two outcomes of high density, the lower total number of plants flowering and the lower number of plants flowering at most census dates, tended to reduce potential N_e. In contrast, it was low density, where variance in flower number was greatest, that was most likely to yield the greatest reduction in N_e due to variance in progeny number.At high density the potential for assortative mating among tall plants was much greater and occurred later than among large plants at low density. Much of the potential high density assortative mating occurred late in the phenology of individual plants, when there was likely to be lower fruit set.We discuss how ecological agents that alter flowering phenology can potentially alter the genetics of populations, the level and timing of assortative mating and, if genetic variation for response to such ecological agents exists, the potential selection regime.     

气候变化背景下青藏高原东部猪毛蒿物候的变化规律和影响因素

近几十年来青藏高原正经历着广泛而深远的气候变化,这种变化对当地物种的物候及分布格局产生了显著影响。猪毛蒿作为干旱半干旱地区的优势物种和影响群落稳定性的关键物种,青藏高原东缘是其重要的分布区域之一,然而其物候将如何应对气候变化目前我们尚不清楚。为此,基于青海省东部连续20年的气象数据和原位物候观测实验,探讨猪毛蒿物候的变化规律及量化不同气候因子的相对贡献率。结果表明：1)过去20年间年均温和年均每日日照时长分别呈现出显著的上升和下降趋势,而年降水和年均每日最大风速没有显著的变化;2)20年间猪毛蒿返青-开花的时间间隔和开花-结果的时间间隔并未表现出显著的变化趋势,结果-枯黄的时间间隔显著缩短;3)所有气候因子均对猪毛蒿不同物候的时间间隔有显著影响,其中最大风速是影响猪毛蒿物候时间间隔最重要的气候因子。这一研究发现可以为气候变化情景下青藏高原高寒植物的保护和利用提供理论依据。     

浙江古田山亚热带常绿阔叶林开花物候:气候因素、系统发育关系和功能性状的影响

植物的开花物候受气候因素、植物系统发育关系和功能性状的影响。然而当前植物开花物候研究中未见同时考虑这3个因素的报道。为了解它们相互之间的影响, 本研究利用中国东部地区浙江省古田山国家级自然保护区亚热带常绿阔叶林24 ha大样地(GTS; 118°03′50′′-118°11′12.2′′ E, 29°10′19′′-29°17′41′′ N)设置的130个种子雨收集器5年的开花数据检验这3个因素对开花的影响。结果表明, 古田山植物的开花高峰期集中在5月, 群落开花格局明显受温度和降雨的影响。利用植物DNA条形码数据研究发现, 植物间系统发育关系对古田山植物开花时间有显著影响, 亲缘关系近的物种开花时间更相近。植物的平均开花时间受最大树高的影响, 但不受传粉方式、花色、种子质量和扩散方式的影响。该研究结果说明气候因素、植物系统发育关系和功能性状都可能影响植物开花物候格局, 同时考虑这3个因素能够帮助我们更好地理解开花物候格局。