Maritime climate influence on chaparral composition and diversity in the coast range of central California

We investigated the hypothesis that maritime climatic factors associated with summer fog and low cloud stratus (summer marine layer) help explain the compositional diversity of chaparral in the coast range of central California. We randomly sampled chaparral species composition in 0.1‐hectare plots along a coast‐to‐interior gradient. For each plot, climatic variables were estimated and soil samples were analyzed. We used Cluster Analysis and Principle Components Analysis to objectively categorize plots into climate zone groups. Climate variables, vegetation composition and various diversity measures were compared across climate zone groups using ANOVA and nonmetric multidimensional scaling. Differences in climatic variables that relate to summer moisture availability and winter freeze events explained the majority of variance in measured conditions and coincided with three chaparral assemblages: maritime (lowland coast where the summer marine layer was strongest), transition (upland coast with mild summer marine layer influence and greater winter precipitation), and interior sites that generally lacked late summer water availability from either source. Species turnover (β‐diversity) was higher among maritime and transition sites than interior sites. Coastal chaparral differs from interior chaparral in having a higher obligate seeder to facultative seeder (resprouter) ratio and by being dominated by various Arctostaphylos species as opposed to the interior dominant, Adenostoma fasciculatum. The maritime climate influence along the California central coast is associated with patterns of woody plant composition and β‐diversity among sites. Summer fog in coastal lowlands and higher winter precipitation in coastal uplands combine to lower late dry season water deficit in coastal chaparral and contribute to longer fire return intervals that are associated with obligate seeders and more local endemism. Soil nutrients are comparatively less important in explaining plant community composition, but heterogeneous azonal soils contribute to local endemism and promote isolated chaparral patches within the dominant forest vegetation along the coast.     

Phenological patterns in a natural population of a tropical timber tree species, Milicia excelsa (Moraceae): Evidence of isolation by time and its interaction with feeding strategies of dispersers

? Premise of the study: Population genetic structuring over limited timescales is commonly viewed as a consequence of spatial constraints. Indirect approaches have recently revealed reproductive isolation resulting from flowering time (so-called isolation by time, IBT). Since phenological processes can be subject to selection, the persistence of flowering asynchrony may be due to opposing selective pressures during mating, dispersal, and regeneration phases. Our study aimed to investigate phenology, fruit handling by animals, and their interaction in a timber tree species, Milicia excelsa. ? Methods: We analyzed phenological data collected over 6 years on 69 genotyped trees in a Cameroonian natural rainforest complemented by data from germination trials and field observations of dispersers. ? Key results: Initiation of flowering was correlated with variation in temperature and relative humidity, but was also affected by genetic factors: pairwise differences in flowering time between nearby individuals correlated with kinship coefficient, and earliness of flowering remained stable over time. A decrease in mean seed production per fruit with increasing flowering time suggests selection against late bloomers. However, germination rate was not affected by seed collection date, and the main seed disperser, the bat Eidolon helvum, seemed to increase in abundance at the end of the reproductive season and preferred trees in open habitats where early and late bloomers are expected. ? Conclusions: The pairwise approach performs well in detecting IBT. The persistence of different mating pools in such a case may result from a trade off between selective forces during the mating and seed dispersal processes.     

Significance of summer fog and overcast for drought stress and ecological functioning of coastal California endemic plant species

Aim Fog drip is a crucial water source for plants in many ecosystems, including a number of global biodiversity hotspots. In California, dozens of rare, drought‐sensitive plant species are endemic to coastal areas where the dominant summer moisture source is fog. Low clouds that provide water to these semi‐arid ecosystems through fog drip can also sharply reduce evaporative water losses by providing shade. We quantified the relative hydrological importance of cloud shading vs. fog drip. We then examined how both factors influence the range dynamics of an apparently fog‐dependent plant species spanning a small‐scale cloud gradient. Location The study area is on Santa Cruz Island off the coast of southern California. It is near the southern range limit of bishop pine (Pinus muricata D. Don), a tree endemic to the coasts of California and Baja, Mexico. Methods We measured climate across a pine stand along a 7 km, coastal–inland elevation transect. Short‐term (1–5 years) monitoring and remote sensing data revealed strong climatic gradients driven primarily by cloud cover. Long‐term (102 years) effects of these gradients were estimated using a water balance model. Results We found that shade from persistent low clouds near the coast reduced annual drought stress by 22–40% compared with clearer conditions further inland. Fog drip at higher elevations provided sufficient extra water to reduce annual drought stress by 20–36%. Sites located at both high elevation and nearer the coast were subject to both effects. Together, these effects reduced average annual drought stress by 56% and dramatically reduced the frequency of severe drought over the last century. At lower elevation (without appreciable fog drip) and also near the inland edge of the stand (with less cloud shading) severe droughts episodically kill most pine recruits, thereby limiting the local range of this species. Main conclusions Persistent cloud shading can influence hydrology as much as fog drip in cloud‐affected ecosystems. Understanding the patterns of both cloud shading and fog drip and their respective impacts on ecosystem water budgets is necessary to fully understand past species range shifts and to anticipate future climate change‐induced range shifts in fog‐dependent ecosystems.     

Greater temperature sensitivity of plant phenology at colder sites: implications for convergence across northern latitudes

Warmer temperatures are accelerating the phenology of organisms around the world. Temperature sensitivity of phenology might be greater in colder, higher latitude sites than in warmer regions, in part because small changes in temperature constitute greater relative changes in thermal balance at colder sites. To test this hypothesis, we examined up to 20 years of phenology data for 47 tundra plant species at 18 high‐latitude sites along a climatic gradient. Across all species, the timing of leaf emergence and flowering was more sensitive to a given increase in summer temperature at colder than warmer high‐latitude locations. A similar pattern was seen over time for the flowering phenology of a widespread species, Cassiope tetragona. These are among the first results highlighting differential phenological responses of plants across a climatic gradient and suggest the possibility of convergence in flowering times and therefore an increase in gene flow across latitudes as the climate warms.     

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...     

Phenological response of tundra plants to background climate variation tested using the International Tundra Experiment

The rapidly warming temperatures in high-latitude and alpine regions have the potential to alter the phenology of Arctic and alpine plants, affecting processes ranging from food webs to ecosystem trace gas fluxes. The International Tundra Experiment (ITEX) was initiated in 1990 to evaluate the effects of expected rapid changes in temperature on tundra plant phenology, growth and community changes using experimental warming. Here, we used the ITEX control data to test the phenological responses to background temperature variation across sites spanning latitudinal and moisture gradients. The dataset overall did not show an advance in phenology; instead, temperature variability during the years sampled and an absence of warming at some sites resulted in mixed responses. Phenological transitions of high Arctic plants clearly occurred at lower heat sum thresholds than those of low Arctic and alpine plants. However, sensitivity to temperature change was similar among plants from the different climate zones. Plants of different communities and growth forms differed for some phenological responses. Heat sums associated with flowering and greening appear to have increased over time. These results point to a complex suite of changes in plant communities and ecosystem function in high latitudes and elevations as the climate warms.     

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.     

Links between plant species’ spatial and temporal responses to a warming climate

To generate realistic projections of species’ responses to climate change, we need to understand the factors that limit their ability to respond. Although climatic niche conservatism, the maintenance of a species’s climatic niche over time, is a critical assumption in niche-based species distribution models, little is known about how universal it is and how it operates. In particular, few studies have tested the role of climatic niche conservatism via phenological changes in explaining the reported wide variance in the extent of range shifts among species. Using historical records of the phenology and spatial distribution of British plants under a warming climate, we revealed that: (i) perennial species, as well as those with weaker or lagged phenological responses to temperature, experienced a greater increase in temperature during flowering (i.e. failed to maintain climatic niche via phenological changes); (ii) species that failed to maintain climatic niche via phenological changes showed greater northward range shifts; and (iii) there was a complementary relationship between the levels of climatic niche conservatism via phenological changes and range shifts. These results indicate that even species with high climatic niche conservatism might not show range shifts as instead they track warming temperatures during flowering by advancing their phenology.     

利用控制实验研究植物物候对气候变化的响应综述

物候是植物在长期适应环境过程中形成的生长发育节点。长时间地面物候观测数据表明,近50年全球乔木、灌木、草本植物的春季物候期受温度升高、降水与辐射变化等影响,以每10年2 d到10 d的速率提前。但因植物物候响应气候因子的机制仍不清楚,导致对未来气候变化情景下的植物物候变化预测存在较大的不确定性。在此背景下,控制实验成为探究气候因子对植物物候影响机制的重要手段。综述了物候控制实验中不同气候因子（温度、水分、光照等）的控制方法。总结了目前为止控制实验在植物物候对气候因子响应方面得到的重要结论,发现植物春季物候期（展叶、开花等）主要受冷激、驱动温度与光周期的影响,秋季物候期（叶变色和落叶）主要受低温、短日照与水分胁迫的影响。提出未来物候控制实验应重点解决木本植物在秋季进入休眠的时间点确定、低温和短日照对木本植物秋季物候的交互作用量化、草本植物春秋季物候的影响因子识别等科学问题。     

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.     

Beyond gradual warming: extreme weather events alter flower phenology of European grassland and heath species

Shifts in the phenology of plant and animal species or in the migratory arrival of birds are seen as ‘fingerprints’ of global warming. However, even if such responses have been documented in large continent‐wide datasets of the northern hemisphere, all studies to date correlate the phenological pattern of various taxa with gradual climatic trends. Here, we report a previously unobserved phenomenon: severe drought and heavy rain events caused phenological shifts in plants of the same magnitude as one decade of gradual warming. We present data from two vegetation periods in an experimental setting containing the first evidence of shifted phenological response of 10 grassland and heath species to simulated 100‐year extreme weather events in Central Europe. Averaged over all species, 32 days of drought significantly advanced the mid‐flowering date by 4 days. The flowering length was significantly extended by 4 days. Heavy rainfall (170 mm over 14 days) had no significant effect on the mid‐flowering date. However, heavy rainfall reduced the flowering length by several days. Observed shifts were species‐specific, (e.g. drought advanced the mid‐flowering date for Holcus lanatus by 1.5 days and delayed the mid‐flowering date for Calluna vulgaris by 5.7 days, heavy rain advanced mid‐flowering date of Lotus corniculatus by 26.6 days and shortened the flowering length of the same species by 36.9 days). Interestingly, the phenological response of individual species was modified by community composition. For example, the mid‐flowering date of C. vulgaris was delayed after drought by 9.3 days in communities composed of grasses and dwarf shrubs compared with communities composed of dwarf shrubs only. This indicates that responses to extreme events are context specific. Additionally, the phenological response of experimental communities to extreme weather events can be modified by the functional diversity of a stand. Future studies on phenological response patterns related to climate change would profit from explicitly addressing the role of extreme weather events.     

Predictors of plant phenology in a diverse high‐latitude alpine landscape: growth forms and topography

Question: Different plant growth forms may have distinctly different functioning in ecosystems. Association of phenological patterns with growth form will therefore help elucidate the role of phenology in an ecosystem. We ask whether growth forms of common vascular plants differ in terms of vegetative and flowering phenology, and if such phenological differences are consistent across environmental gradients caused by landscape‐scale topography. Location: A high‐latitude alpine landscape in Finnmark County, Norway (70°N). Methods: We assessed vegetative and flowering phenology repeatedly in five growth forms represented by 11 common vascular plant species across an altitudinal gradient and among differing slope aspects. Results: Species phenology clustered well according to growth form, and growth form strongly explained variation in both flowering and vegetative phenology. Altitude and aspect were poor predictors of phenological variation. Vegetative phenology of the growth forms, ranked from slowest to fastest, was in the order evergreen shrubs <sedges‐deciduous shrubs <grasses <forbs, while a reverse ranking was found for flowering phenology. Conclusion: Growth form‐specific phenological patterns are associated with fundamentally different abilities for resource acquisition and resource conservation. The weak effect of landscape‐scale topographic factors indicates that variation within growth forms is mainly influenced by local environmental factors not accounted for in this study. On the basis of these results, we argue that growth forms should be considered as predictors of phenology together with the customary use of topography and normalized difference vegetation index, especially when assessing the role of phenology in an ecosystem.     

Relationships between plant traits and climate in the Mediterranean region: A pollen data analysis

Question: What are the correlations between the degree of drought stress and temperature, and the adoption of specific adaptive strategies by plants in the Mediterranean region? Location: 602 sites across the Mediterranean region. Method: We considered 12 plant morphological and phenological traits, and measured their abundance at the sites as trait scores obtained from pollen percentages. We conducted stepwise regression analyses of trait scores as a function of plant available moisture (α) and winter temperature (MTCO). Results: Patterns in the abundance for the plant traits we considered are clearly determined by α, MTCO or a combination of both. In addition, trends in leaf size, texture, thickness, pubescence and aromatic leaves and other plant level traits such as thorniness and aphylly, vary according to the life form (tree, shrub, forb), the leaf type (broad, needle) and phenology (evergreen, summer‐green). Conclusions: Despite conducting this study based on pollen data we have identified ecologically plausible trends in the abundance of traits along climatic gradients. Plant traits other than the usual life form, leaf type and leaf phenology carry strong climatic signals. Generally, combinations of plant traits are more climatically diagnostic than individual traits. The qualitative and quantitative relationships between plant traits and climate parameters established here will help to provide an improved basis for modelling the impact of climate changes on vegetation and form a starting point for a global analysis of pollen‐climate relationships.     

Contrasting effects of warming and increased snowfall on Arctic tundra plant phenology over the past two decades

Recent changes in climate have led to significant shifts in phenology, with many studies demonstrating advanced phenology in response to warming temperatures. The rate of temperature change is especially high in the Arctic, but this is also where we have relatively little data on phenological changes and the processes driving these changes. In order to understand how Arctic plant species are likely to respond to future changes in climate, we monitored flowering phenology in response to both experimental and ambient warming for four widespread species in two habitat types over 21 years. We additionally used long‐term environmental records to disentangle the effects of temperature increase and changes in snowmelt date on phenological patterns. While flowering occurred earlier in response to experimental warming, plants in unmanipulated plots showed no change or a delay in flowering over the 21‐year period, despite more than 1 °C of ambient warming during that time. This counterintuitive result was likely due to significantly delayed snowmelt over the study period (0.05–0.2 days/yr) due to increased winter snowfall. The timing of snowmelt was a strong driver of flowering phenology for all species – especially for early‐flowering species – while spring temperature was significantly related to flowering time only for later‐flowering species. Despite significantly delayed flowering phenology, the timing of seed maturation showed no significant change over time, suggesting that warmer temperatures may promote more rapid seed development. The results of this study highlight the importance of understanding the specific environmental cues that drive species’ phenological responses as well as the complex interactions between temperature and precipitation when forecasting phenology over the coming decades. As demonstrated here, the effects of altered snowmelt patterns can counter the effects of warmer temperatures, even to the point of generating phenological responses opposite to those predicted by warming alone.     

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.     

Phenology and polyploidy in annual Brachypodium species (Poaceae) along the aridity gradient in Israel

Local adaptation of plants along environmental gradients provides strong evidence for clinal evolution mediated by natural selection. Plants have developed diverse strategies to mitigate stress, for example, drought escape is a phenological strategy to avoid drought stress, while polyploidy was proposed as a genomic adaptation to stress. Polyploidy as an adaptation to aridity (an environmental parameter integrating temperature and precipitation) was previously documented in annual Brachypodium spp. (Poaceae) in the Western Mediterranean. Here, we examined whether polyploidy or phenology are associated with aridity in annual Brachypodium spp. along the aridity gradient in the Eastern Mediterranean. Using flow cytometry, we determined ploidy levels of plants from natural populations along the Israeli gradient, spanning ∼424 km from mesic Mediterranean to extreme desert climates. In a common garden we recorded time of seedling emergence, flowering and senescence. We tested whether the proportion of allotetraploids in the populations and phenological traits were associated with aridity. Contrary to a previous study in the Western Mediterranean, we found no effect of aridity on the proportion of allotetraploids and diploids within populations. Interestingly, phenology was associated with aridity: time of emergence was later, while flowering and senescence were earlier in desert plants. Our results indicate that in the Eastern Mediterranean, adaptation of Brachypodium to aridity is mediated mainly by phenology, rather than ploidy level. Therefore, we suggest that genome duplication is not the main driver of adaptation to environmental stress; rather, phenological change as a drought escape mechanism may be the major adaptation.     

Responses of green-up dates of grasslands in China and woody plants in Europe to air temperature and precipitation: Empirical evidences based on survival analysis

长期以来, 气候与植物物候关系的研究大多基于线性模型, 但植被物候对气候变化的响应可能是非线性的。该文利用非线性模型——生存分析模型来分析时间序列中过去事件(气候因子)对目的变量(物候)的作用: 用生存分析模型分析了春季气温和降水量对内蒙古草地、青藏高原草甸和欧洲地区木本植物返青期的影响。其中, 内蒙古与青藏高原的物候信息来自遥感数据, 欧洲地区物候信息为实测数据。蒙特卡洛方法用于拟合模型参数。结果表明: 生存分析模型适合对上述不同研究对象的物候影响因素进行分析, 并能模拟非线性效应; 在内蒙古草地, 春季气温和降水对春季返青有很大的影响, 而青藏高原草甸和欧洲地区木本植物返青期对春季降水响应较小, 对春季气温变化的响应随Holdridge干燥度指数上升而下降; 在预测返青期时发现: 春季平均气温提高1 ℃会导致上述地区返青期提前1-6天; 而春季气温与降水的增加会导致返青期发生明显的非线性变化, 这种非线性效应无法基于线性模型模拟出来。结果说明生存分析模型既能用于分析不同尺度下植物物候与气候的关系, 也能用于模型预测, 尤其适合探讨大幅度气候变化对物候的非线性影响。     

Year clustering analysis for modelling olive flowering phenology

It is now widely accepted that weather conditions occurring several months prior to the onset of flowering have a major influence on various aspects of olive reproductive phenology, including flowering intensity. Given the variable characteristics of the Mediterranean climate, we analyse its influence on the registered variations in olive flowering intensity in southern Spain, and relate them to previous climatic parameters using a year-clustering approach, as a first step towards an olive flowering phenology model adapted to different year categories. Phenological data from Cordoba province (Southern Spain) for a 30-year period (1982–2011) were analysed. Meteorological and phenological data were first subjected to both hierarchical and “K-means” clustering analysis, which yielded four year-categories. For this classification purpose, three different models were tested: (1) discriminant analysis; (2) decision-tree analysis; and (3) neural network analysis. Comparison of the results showed that the neural-networks model was the most effective, classifying four different year categories with clearly distinct weather features. Flowering-intensity models were constructed for each year category using the partial least squares regression method. These category-specific models proved to be more effective than general models. They are better suited to the variability of the Mediterranean climate, due to the different response of plants to the same environmental stimuli depending on the previous weather conditions in any given year. The present detailed analysis of the influence of weather patterns of different years on olive phenology will help us to understand the short-term effects of climate change on olive crop in the Mediterranean area that is highly affected by it.     

Nonlinear flowering responses to climate: are species approaching their limits of phenological change?

Many alpine and subalpine plant species exhibit phenological advancements in association with earlier snowmelt. While the phenology of some plant species does not advance beyond a threshold snowmelt date, the prevalence of such threshold phenological responses within plant communities is largely unknown. We therefore examined the shape of flowering phenology responses (linear versus nonlinear) to climate using two long-term datasets from plant communities in snow-dominated environments: Gothic, CO, USA (1974–2011) and Zackenberg, Greenland (1996–2011). For a total of 64 species, we determined whether a linear or nonlinear regression model best explained interannual variation in flowering phenology in response to increasing temperatures and advancing snowmelt dates. The most common nonlinear trend was for species to flower earlier as snowmelt advanced, with either no change or a slower rate of change when snowmelt was early (average 20% of cases). By contrast, some species advanced their flowering at a faster rate over the warmest temperatures relative to cooler temperatures (average 5% of cases). Thus, some species seem to be approaching their limits of phenological change in response to snowmelt but not temperature. Such phenological thresholds could either be a result of minimum springtime photoperiod cues for flowering or a slower rate of adaptive change in flowering time relative to changing climatic conditions.     

Adaptation to climate through flowering phenology: a case study in Medicago truncatula

Local climatic conditions likely constitute an important selective pressure on genes underlying important fitness‐related traits such as flowering time, and in many species, flowering phenology and climatic gradients strongly covary. To test whether climate shapes the genetic variation on flowering time genes and to identify candidate flowering genes involved in the adaptation to environmental heterogeneity, we used a large Medicago truncatula core collection to examine the association between nucleotide polymorphisms at 224 candidate genes and both climate variables and flowering phenotypes. Unlike genome‐wide studies, candidate gene approaches are expected to enrich for the number of meaningful trait associations because they specifically target genes that are known to affect the trait of interest. We found that flowering time mediates adaptation to climatic conditions mainly by variation at genes located upstream in the flowering pathways, close to the environmental stimuli. Variables related to the annual precipitation regime reflected selective constraints on flowering time genes better than the other variables tested (temperature, altitude, latitude or longitude). By comparing phenotype and climate associations, we identified 12 flowering genes as the most promising candidates responsible for phenological adaptation to climate. Four of these genes were located in the known flowering time QTL region on chromosome 7. However, climate and flowering associations also highlighted largely distinct gene sets, suggesting different genetic architectures for adaptation to climate and flowering onset.