POLLINATION ECOLOGY OF LABIATAE IN A PHRYGANIC (EAST MEDITERRANEAN) ECOSYSTEM

This study was conducted in a phryganic (East Mediterranean) ecosystem at Daphni, near Athens, Greece. The Labiatae, represented by ten species belonging to nine genera, dominate in this ecosystem type. They flower from February to July. Both flowering time and nectar quantity are related to the species ability to tolerate intense water stress. Labiatae are visited by 201 insect species. Of these, 43 are exclusively supported by the family and 37 are monotropous. Solitary bees (mainly Anthophoridae, Megachilidae, Halictidae) constitute 47.3% of pollinators. The family is important in hosting specialized bees (15 species) in phrygana, particularly late in the flowering season. Labiatae species form two equally represented groups in this system; namely, the late winter-early spring (early) flowering, visited by relatively few pollinator species, and the late spring-summer (late) flowering species, visited by numerous pollinators. This temporal distinction is accompanied by different pollination profiles that include duration of anthesis, reward to pollinators, floral attractiveness, and flower character differentiation. All of these attributes are maximized in the early flowering period. This strategy suggests a mechanism for resource partitioning at a time when the pollinator resource is limited and competition for the services of pollinators is expected to be intense. Contrary to the current theory concerning cornucopian species, the copiously rewarding flowers of Labiatae in phrygana are not those abundantly serviced by pollinators.     

Flowering phenological pattern in crowns of four temperate deciduous tree species and its reproductive implications

Temperate deciduous forest trees flower in spring, a period that starts when the trees lack leaves and when weather is unpredictable, including frost events, and ends when the forest becomes green and vertical microclimatic gradients are established. This paper asks whether there are spatio-temporal patterns in the development of flowering in trees, and how they relate to reproductive processes. Using a crane, flowering phenology was studied in the crowns of ca. 200 trees of four species, from early spring (ash) through the period of leaf-unfolding (maples) to early summer (lime). Flowering levels in different crown regions were documented quantitatively and repeatedly during the flowering season and compared among individuals and among species. Early-flowering trees displayed a clear and consistent acropetalous and centrifugal flowering pattern, while this pattern disappeared in species that flowered after leaves unfolded. This pattern was superposed on the basic flowering rhythm of each species, and was influenced by effects of direct sunlight, acting at a small scale in early spring and at a large scale in early summer. As this acropetalous centrifugal pattern contrasts the microclimatic gradients that develop only after leaves unfold, it might indicate physiological processes in the 'awakening' of trees, as well as evolutionary processes that took place in temperate trees during adaptation to a temperate climate.     

Flowering phenology of <Emphasis Type="Italic">Ulex europaeus</Emphasis>: ecological consequences of variation within and among populations

Reproductive phenology of gorse (Ulex europaeus L., Genisteae, Fabaceae) is unusual in that the onset and duration of flowering vary greatly among individuals within populations: some plants initiate flowering in autumn or winter and continue flowering through spring, others initiate flowering in early spring. To understand the origin of this diversity and its ecological consequences, we investigated flowering phenology of randomly sampled individuals from five different natural populations in Brittany (France). Reproductive success was evaluated for individuals with contrasting flowering patterns, from 16 natural populations. Flower production, pod production, seed production and seed predation were estimated. Plants initiating flowering in spring produced larger numbers of flowers and pods over a shorter period than plants flowering from winter to spring, which produced few flowers and pods at a time but over a longer period. Pod production of long-flowering plants did not differ significantly between winter and spring, but their pods were more intensively attacked by seed predators in spring than in winter. We discuss our results in relation to biotic and abiotic parameters. We postulate that long-flowering can be interpreted as a bet-hedging strategy, spreading the risk of pod failure (rotting or freezing) in winter and of seed predation in spring.     

Spiny east Mediterranean plant species flower later and in a drier season than non-spiny species

We compared the flowering phenology of spiny and non-spiny native species belonging to three families (Asteraceae, Fabaceae and Lamiaceae), which include the highest number of spiny species in the flora of Israel. We found that the peak of flowering (when the highest number of species flowered) was 4–8 weeks later for spiny species than for non-spiny species. The flowering peak of non-spiny species was in late March, while that of spiny species was at the beginning of May. The seasonal shift in flowering time from the main season, when most Mediterranean plants bloom, to the end of the flowering season, when fewer species bloom, might be the evolutionary result of a change in phenology reducing the competition for pollinators. Our results clearly indicate that spinescence of plants in the semi-arid east Mediterranean region is associated with a delayed flowering season at the beginning of the dry summer when most of the herbaceous vegetation is already dry. During this season, mammalian grazers consume any edible herbaceous vegetation, selecting for late flowering species that allocate more resources for anti-herbivore defenses than early flowering species. There is a well-known global geographical trend where the occurrence of spiny plants is higher in arid regions than in humid ones. In parallel to the global trend, we show a seasonal one, that non-spiny plants grow and flower in the spring, which is the main flowering season in the Mediterranean basin, while spiny plants flower later, in the hot and dry summer. Under the current trend of global warming, there are prospects of future increase in the dominance of spiny species in the Mediterranean region.     

民勤荒漠植被对气候变化的响应

运用民勤荒漠区1974-2009年物候观测资料和2002-2010年植被样方观测资料以及同期气象资料,分析了荒漠植被对气候变化的响应.结果表明:1961-2010年,民勤荒漠区气温升高,空气湿度增大,年均气温升高速率大于全球水平和中国近百年平均水平;植物对气温变化的响应主要表现在春季物候提前、秋季物候推迟、生长季延长;植被对降水量变化的响应主要表现为植被盖度和纯盖度随降水量减少而降低,植株密度、植物多度随降水量变化而波动;植被盖度和纯盖度与年降水量的相关性较高,然后依次为6-7月和4-5月的降水量;植株密度和植物多度与9月降水量呈正相关;植物春季物候提前的次序是芽初膨大期＞芽开放期＞开花始期＞展叶始期和展叶盛期＞花蕾序出现＞开花盛期＞开花末期＞果实成熟期;秋季物候推迟的次序是叶全变色期＞落叶始期＞叶初变色期＞落叶末期.春季气温升高对民勤荒漠区植物物候的影响大于秋季气温升高对物候的影响.     

Phenology and abundance in relation to climatic variation in a sub-arctic insect herbivore–mountain birch system

The two forest-defoliating geometrid moth species Operophtera brumata and Epirrita autumnata are known to exhibit different altitudinal distribution patterns in northern birch forests. One possible explanation for this is that altitudinal climatic variation differentially affects the performance of two species through mismatching larval and host plant phenology. We explored this hypothesis by investigating the relationship between larval phenology and leaf phenology of Betula pubescens, which is the main host plant of both moth species, along ten replicate altitudinal transects during two springs with contrasting climate in northern Norway. There was a distinct monotonous cline in host plant phenology with increasing altitude in both years of the study, but the development of the leaves were generally 14 days later in the first of the 2 years due to cold spring weather. We found that larval development of both species closely tracked host plant leaf phenology independent of altitude and year. However, at the time of sampling, E. autumnata was approximately one instar ahead of O. brumata at all altitudes, probably reflecting that E. autumnata has faster early instar growth than O. brumata. The abundance of O. brumata was lowest at the altitudinal forest-line, while E. autumnata was lowest near sea level. Our results do not indicate that the altitudinal distribution patterns of the two moth species is due to any phenological mismatch between larval and host plant phenology. We suggest rather that natural enemies at low altitudes limit larval survival and thus abundance of E. autumnata, while an early onset of winter at the forest limit reduces survival of late eclosing adults of O. brumata.     

Estimating the first flowering and full blossom dates of Yoshino cherry (Cerasus × yedoensis ‘Somei-yoshino’) in Japan using machine learning algorithms

Climate change alters the phenology of various plants. For example, increasing temperatures shift the first flowering and full blossom days of Yoshino cherry trees and affect cultural events related to cherry blossoms. We developed models to estimate the first flowering and full blossom dates of Yoshino cherry in Japan based on temperature and phenological data observed at 82 stations in Japan for 68 years (1953–2020). Three machine learning algorithms, namely, the random forest (RF), artificial neural network (ANN), and gradient boosting decision tree (GBDT) algorithms, were utilized, and the hyperparameters were optimized using Optuna. The GBDT models produced the best estimation accuracy, with an overall root mean square error (RMSE) = 1.53 and 1.48 days for the first flowering date and full blossom date, respectively. Furthermore, our analysis using Shapley Additive Explanations (SHAP) revealed that in the RF and GBDT models, the low temperature in winter and high temperature in spring would advance the estimated first flowering and full blossom dates.     

Local snow melt and temperature—but not regional sea ice—explain variation in spring phenology in coastal Arctic tundra

The Arctic is undergoing dramatic environmental change with rapidly rising surface temperatures, accelerating sea ice decline and changing snow regimes, all of which influence tundra plant phenology. Despite these changes, no globally consistent direction of trends in spring phenology has been reported across the Arctic. While spring has advanced at some sites, spring has delayed or not changed at other sites, highlighting substantial unexplained variation. Here, we test the relative importance of local temperatures, local snow melt date and regional spring drop in sea ice extent as controls of variation in spring phenology across different sites and species. Trends in long‐term time series of spring leaf‐out and flowering (average span: 18 years) were highly variable for the 14 tundra species monitored at our four study sites on the Arctic coasts of Alaska, Canada and Greenland, ranging from advances of 10.06 days per decade to delays of 1.67 days per decade. Spring temperatures and the day of spring drop in sea ice extent advanced at all sites (average 1°C per decade and 21 days per decade, respectively), but only those sites with advances in snow melt (average 5 days advance per decade) also had advancing phenology. Variation in spring plant phenology was best explained by snow melt date (mean effect: 0.45 days advance in phenology per day advance snow melt) and, to a lesser extent, by mean spring temperature (mean effect: 2.39 days advance in phenology per °C). In contrast to previous studies examining sea ice and phenology at different spatial scales, regional spring drop in sea ice extent did not predict spring phenology for any species or site in our analysis. Our findings highlight that tundra vegetation responses to global change are more complex than a direct response to warming and emphasize the importance of snow melt as a local driver of tundra spring phenology.     

澳门九澳山海滨群落10种植物盛花期物候对极端气候事件的响应

为探究极端气候事件对植物的影响,对澳门九澳山海滨群落10种植物2012-2017年盛花期物候进行了观察。结果表明,植物的盛花期一般在3-9月,其中有4种植物为5月。2013和2016年早春澳门的极端强降水使植物的盛花期出现了明显的提前或者推迟。9种植物的盛花期与盛花期前0～2个月和上一年的秋冬季的月均温度或月降水存在显著相关性。温度和降水对植物盛花期的影响差异不大,但晚花植物对降水更加敏感。这为澳门和邻近岛屿的生态恢复和园林树种选材提供参考。     

Mexical plant phenology: is it similar to Mediterranean communities?

The sclerophyllous, evergreen vegetation found in Mexico under tropical climate is named 'Mexical' (MEX) and presents many traits that have been thought to converge under a Mediterranean climate. Flowering phenology is strongly similar across Mediterranean-type ecosystems (MTEs) and this paper investigates MEX plant phenology in this context. The common history of the vegetation and the differences in the climatic conditions experienced by MEX and MTE taxa provide an ideal scenario to infer the relative importance of natural selection and historical constraints in the phenological response of plants to climatic conditions. This study has involved collecting field and bibliographic data on flowering phenology of MEX communities to detect (1) similarities at the community level between MTEs and MEX, (2) similarities between Tertiary and Quaternary taxa in MTEs and MEX, and (3) similarities between congeneric taxa from MEX and MTEs (taxa sharing a common ancestor but having evolved under different climates). Flowering in MEX does not occur mainly in spring, as in MTEs, but in summer, suggesting a response that maximizes water use in the rainy season. Flowering phenology of MEX species differed from their MTE congeneric species, suggesting that even though a common ancestor is shared, environmental pressures have led to different phenological responses in MEX and MTE plants. The flowering season for species that originated in the Tertiary and Quaternary did not differ in MEX, as expected, because of climatic uniformity along the whole time line. In MTEs, flowering differences between Tertiary and Quaternary species were not congruent, suggesting that the balance between the historical constraints and the selective force of the Mediterranean climate is different among the three MTEs, and a particular explanation is needed for each. © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 138 , 297–303.     

杧果花期延迟技术研究进展

杧果Mangifera indica花期大多集中在2～3月,易受低温阴雨天气影响,导致其授粉受精不良、落花落果,并加重白粉病和炭疽病的危害。本文简要介绍低温阴雨天气对杧果花期的影响,并详细阐述生产上推迟花期的常用措施,如回缩、重修剪、摘花穗、抹花、应用植物生长调节剂、采后灌水、培养二次秋梢或利用冬梢结果等,为杧果开花调控及花期管理提供参考。     

The roles of shifting and filtering in generating community‐level flowering phenology

Plant phenologies are key components of community assembly and ecosystem function, yet we know little about how phenological patterns differ among ecosystems. Community‐level phenological patterns may be driven by the filtering of species into communities based on their phenology or by intraspecific responses to local conditions that shift when species flower. To understand the relative roles of filtering and shifting on community‐level phenological patterns we compared patterns of first flowering dates (FFD) for herbaceous species at Konza Prairie, KS, USA with those from the colder Fargo, ND, USA area and from Chinnor, England, which has a less continental climate. Comparing patterns of FFD supports that Konza's flowering patterns are potentially influenced both by filtering species that flower early in the growing season and by phenological shifting. Konza species flowering dates were earlier in the spring and later in the fall compared to Fargo, but were not shifted compared to Chinnor, which had a unique suite of early‐flowering species. In all, comparing flowering phenology among three sites reveals that intraspecific responses to climate can generate phenological shifts that compress or stretch community‐level phenological patterns, while novel niches in phenological space can also alter community‐level patterns. Community flowering patterns related to climate suggest that climatic warming has the potential to further distribute flowering of the Konza flora over a longer period, but also could further open it to introductions of non‐native species that have evolved to flower early in the season.     

荚蒾属植物花期物候对春季异常气象事件的响应

在全球气候变化背景下异常气象事件频繁发生。华北地区在经历了2010年春季的异常冷湿后,2016年春季又出现了异常暖旱的气象事件。对荚蒾属(Viburnum)植物2009–2016年春季的2次气象事件及正常年份的多年花期物候进行观测,发现春季异常暖旱时荚蒾属植物的花期整体提前,但花期时长均未缩短;海拔跨度小的琼花(V.macrocephalum)和欧洲绣球(V.opulus),其始花期、花前积温和花期长短年际变化幅度较大,而海拔跨度较大的李叶荚蒾(V.prunifolium)、黑果荚蒾(V.lantana)和红蕾荚蒾(V.carlesii)无论春季冷湿或暖旱,其始花期、花前积温和花期长短都表现出较小的可塑性,并且这3个物种的始花期也相对固定。春季气候暖旱时荚蒾枝条的同化产物分配增加,枝条增粗明显,叶片生长速率加快,叶面积更大但叶片更薄,远果叶的同化产物分配显著增加,可能意味着果实发育的同化产物供给受到影响。针对异常气象事件中专属植物的花期物候格局的研究,不仅能更精准地追踪物种繁育特征的环境可塑性及系统认识专属植物的气候变化响应,而且可为物候学大尺度数据分析提供基本信息。     

Marine influence controls plant phenological dynamics in Mediterranean Mexico

Aims To assess the role of moisture in phenological timing in the mediterranean coastal flora of Baja California, and specifically to assess the role of coastal fog and ocean-derived moisture in plant phenology. Moisture seems to be the primary driver of flowering times and durations at the arid end of the mediterranean-climate region, where rainfall is often sporadic (temperature and day length can be expected to play a much lesser role as they are not growth limiting). We aimed to understand: What factors drive climatic variation between sites? Are there general flowering patterns allowing us to identify phenological categories? Do flowering patterns vary in relation to site-specific weather? and most importantly, does maritime influence on weather affect flowering dynamics in coastal mediterranean ecosystems?Methods The southernmost extent of the California Floristic Province (in Baja California, Mexico) is a biological diversity hotspot of high endemism and conservation value, with two steep moisture gradients: rainfall (N–S) and coastal fogs (W–E), providing an ideal study system. We installed five weather stations across the moisture gradients, recording data hourly. We monitored flowering phenology in the square kilometer surrounding each weather station from 2010 to 2013. About 86 plant taxa were monitored across the five sites, every 6–8 weeks. Averaged climatic data is presented with general trends in flowering, and specific flowering syndromes were observed. Data for flowering intensity across the sites was analyzed using a principal components analysis.Important findings Data analysis demonstrates a general seasonal pattern in flowering times, but distinct differences in local weather and phenology between the five study sites. Three flowering syndromes are revealed in the flora: (i) water responders or spring bloomers, (ii) day-length responders or fall-blooming taxa and (iii) aseasonal bloomers with no seasonal affinity. The two moisture gradients are the strongest drivers of flowering times. Inland sites showed higher phenological variation than coastal sites where seasonality is dampened by ocean-derived moisture, which extends and buffers perennial plant phenology and is a probable driver of local endemism. Phenological controls vary globally with climate and geography; moisture is the primary driver of phenology in mediterranean climates and fog is an important climatic variable in coastal Mexico.     

Flowering phenology and reproduction of a forest understorey plant species in response to the local environment

Temperate forest understorey plants are subjected to a strong seasonality in their optimal growing conditions. In winter and early spring, low temperatures are suboptimal for plant growth while light becomes limited later in spring season. We can thus expect that differences in plant phenology in relation to spatiotemporal environmental variation will lead to differences in reproductive output, and hence selection. We specifically studied whether early flowering, a paradoxical pattern that is observed in many plant species, is an adaptive strategy, and whether selection for early flowering was confounded with selection for flower duration or was attributable to environmental variables. We used Geum urbanum as a study species to investigate the effect of relevant environmental factors on the species’ flowering phenology and the consequences for plant reproductive output. We monitored the phenology of four to six plants in each of ten locations in a temperate deciduous forest (Belgium). We first quantified variation in flowering time within individuals and related this temporal variation to individual flower reproductive output. Then, we studied inter-individual variation here-in and linked this to reproduction at the plant level, hence studying the selection differential. We found that flowering within individual plants of Geum urbanum was spread over a long period from June to October. Reproductive output of individual flowers, measured as total seed mass per flower, declined during the season. We found no indication for selection for early flowering but rather for longer flower duration. Larger plants had an earlier flowering onset and a higher seed mass, which suggests that these factors covary and are condition dependent. None of the studied environmental variables could explain plant size, although soil pH and to a lesser extent light availability had a positive direct effect on seed mass per plant. Finally, we suggest that the high intra-individual variation in flowering time, which might be a risk spreading strategy of the plant in the presence of seed predation, limits the potential for selection on flowering phenology.

     

Significance of intra-inflorescence variation on flowering time of a spring ephemeral, <Emphasis Type="Italic">Gagea lutea</Emphasis> (Liliaceae), under seasonal fluctuations of pollinator and light availabilities

I studied the relationship between seed-set patterns within inflorescences and temporal variations in light and pollinator availabilities for 2 years in the spring ephemeral species Gagea lutea in a deciduous forest. Timing of canopy closure and seasonal trend of pollinator frequency did not synchronize with the annual fluctuation in flowering phenology. In the early snowmelt year, seed-set success reflected the seasonal pollinator abundance from early to middle flowering periods. In the late snowmelt year, however, seed-set rates were independent of pollinator activity and decreased with canopy closing even after hand pollination. The restricted seed production by defoliation and the increase in seed-set rates at the forest edge suggested that seed production was supported by current photosynthetic carbon gain. Thus, annual fluctuations of reproductive success can explain the variation in flowering phenology within a population although seasonal light deterioration would serve as a selective force for flowering in the early season.     

Exploring the causes of variation in phenology and morphology in Mediterranean geophytes: a genus-wide study of Cyclamen

The phenology and morphology of Mediterranean plants are constrained by drought in summer and cold temperatures in winter. In this study we examine how climatic factors and phylogenetic constraints have shaped variation in the phenology and morphology of 17 species of the genus Cyclamen cultivated in uniform garden conditions. We quantify the extent to which traits differ among subgenera and thus represent conserved traits within evolutionary lineages. We also explore whether leaf, flowering and seed-release phenology are correlated among species, and thus whether variation in flowering phenology results from selection on dispersal phenology. Our results show a significant influence of subgenus membership on leaf and flowering phenology but not on morphological traits or the timing of seed release. Among-species variation in foliage height, leaf size and seed mass (but not in floral traits) is correlated with chromosome number. Leaf traits show that species with a shorter vegetative period have a higher capacity for resource acquisition. Major phenological shifts, i.e. spring vs. autumn flowering and a decoupling of leaf and flower phenology in autumnal flowering species, thus occurred prior to the diversification of species in each subgenus and not as a response to selection on dispersal timing. Leaf and flowering phenology illustrate a gradient of strategies from autumn flowering in the absence of leaves (hysteranthous species) to spring flowering with fully developed foliage (synanthous species). In the former, flowering is uncoupled from resource acquisition by simultaneous photosynthesis, indicative that hysteranthy is a response to temporal unpredictability in the onset of rain after the summer drought. Our results support the idea that whereas leaf development is controlled primarily by moisture availability and secondarily by temperature, flowering is temperature dependent, above a minimum moisture threshold. © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society, 2004, 145 , 469–484.     

青藏高原东缘高寒草甸植物群落的开花物候

利用2008年整个生长季的地面花期物候观测数据,初步研究了青藏高原东缘高寒草甸植物群落开花物候的动态、格局、参数间的联系及其与物种共存的关系.结果表明:植物群落的花期从5月初开始,8月上旬进入开花数目的高峰期,到10月初结束,遍及整个生长季节;不同物种进入花期的时间、峰值期及持续时间不同,各物种花期物候生态位相分离,但群落中大部分物种开花峰值期主要集中在7-8月;不同物种的开花峰值期时间与花期持续时间有负相关关系:开花越早的植物其花期持续时间越长;物种间的开花物候差异主要是由植物本身的特性和环境条件共同决定的,而群落水平上的开花物候格局主要受非生物因素的控制.

Abstract：

Based on the observation on the flowering phenology in the whole growth season of 2008, the dynamics, patterns, and relationships among various parameters of flowering phenology of alpine meadow plant community in eastern Qinghai-Tibetan Plateau were studied, and the potential effects of above-mentioned factors on the species coexistence of the plant community were approached. The flowering period of the plant community started from early May, came into peak period in the first ten days of August, and ended in early October, extending all over whole growth period. Different species had their different first flowering date, peak flowering period, and flowering duration, and their flowering phenologieal niches differentiated obviously. Most species had their peak flowering period centralized in July-August. There was a negative rela-tionship between peak flowering period and flowering duration: the earlier the flowering, the lon-ger the flowering duration. The differences in flowering phonology among the species were mainly determined by species per se characteristics and environmental conditions, but the flowering phe-nological pattern on community level was primarily controlled by abiotic factors.     

Phenological change in a spring ephemeral: implications for pollination and plant reproduction

Climate change has had numerous ecological effects, including species range shifts and altered phenology. Altering flowering phenology often affects plant reproduction, but the mechanisms behind these changes are not well‐understood. To investigate why altering flowering phenology affects plant reproduction, we manipulated flowering phenology of the spring herb Claytonia lanceolata (Portulacaceae) using two methods: in 2011–2013 by altering snow pack (snow‐removal vs. control treatments), and in 2013 by inducing flowering in a greenhouse before placing plants in experimental outdoor arrays (early, control, and late treatments). We measured flowering phenology, pollinator visitation, plant reproduction (fruit and seed set), and pollen limitation. Flowering occurred approx. 10 days earlier in snow‐removal than control plots during all years of snow manipulation. Pollinator visitation patterns and strength of pollen limitation varied with snow treatments, and among years. Plants in the snow removal treatment were more likely to experience frost damage, and frost‐damaged plants suffered low reproduction despite lack of pollen limitation. Plants in the snow removal treatment that escaped frost damage had higher pollinator visitation rates and reproduction than controls. The results of the array experiment supported the results of the snow manipulations. Plants in the early and late treatments suffered very low reproduction due either to severe frost damage (early treatment) or low pollinator visitation (late treatment) relative to control plants. Thus, plants face tradeoffs with advanced flowering time. While early‐flowering plants can reap the benefits of enhanced pollination services, they do so at the cost of increased susceptibility to frost damage that can overwhelm any benefit of flowering early. In contrast, delayed flowering results in dramatic reductions in plant reproduction through reduced pollination. Our results suggest that climate change may constrain the success of early‐flowering plants not through plant‐pollinator mismatch but through the direct impacts of extreme environmental conditions.