A comparative study of satellite and ground-based phenology

Long time series of ground-based plant phenology, as well as more than two decades of satellite-derived phenological metrics, are currently available to assess the impacts of climate variability and trends on terrestrial vegetation. Traditional plant phenology provides very accurate information on individual plant species, but with limited spatial coverage. Satellite phenology allows monitoring of terrestrial vegetation on a global scale and provides an integrative view at the landscape level. Linking the strengths of both methodologies has high potential value for climate impact studies. We compared a multispecies index from ground-observed spring phases with two types (maximum slope and threshold approach) of satellite-derived start-of-season (SOS) metrics. We focus on Switzerland from 1982 to 2001 and show that temporal and spatial variability of the multispecies index correspond well with the satellite-derived metrics. All phenological metrics correlate with temperature anomalies as expected. The slope approach proved to deviate strongly from the temporal development of the ground observations as well as from the threshold-defined SOS satellite measure. The slope spring indicator is considered to indicate a different stage in vegetation development and is therefore less suited as a SOS parameter for comparative studies in relation to ground-observed phenology. Satellite-derived metrics are, however, very susceptible to snow cover, and it is suggested that this snow cover should be better accounted for by the use of newer satellite sensors.     

A temperature-precipitation based leafing model and its application in Northeast China

Plant phenology models, especially leafing models, play critical roles in evaluating the impact of climate change on the primary production of temperate plants. Existing models based on temperature alone could not accurately simulate plant leafing in arid and semi-arid regions. The objective of the present study was to test the suitability of the existing temperature-based leafing models in arid and semi-arid regions, and to develop a temperature-precipitation based leafing model (TP), based on the long-term (i.e., 12-27 years) ground leafing observation data and meteorological data in Northeast China. The better simulation of leafing for all the plant species in Northeast China was given by TP with the fixed starting date (TPn) than with the parameterized starting date (TPm), which gave the smallest average root mean square error (RMSE) of 4.21 days. Tree leafing models were validated with independent data, and the coefficient of determination (R(2)) was greater than 0.60 in 75% of the estimates by TP and the spring warming model (SW) with the fixed starting date. The average RMSE of herb leafing simulated by TPn was 5.03 days, much lower than other models (>9.51 days), while the average R(2) of TPn and TPm were 0.68 and 0.57, respectively, much higher than the other models (<0.22). It indicates that TPn is a universal model and more suitable for simulating leafing of trees and herbs than the prior models. Furthermore, water is an important factor determining herb leafing in arid and semi-arid temperate regions.     

河北省草本植物物候特征及其对气候变暖的响应

以1981—2006年河北省8个国家农业气象观测站的草本植物物候观测资料和47个气象站的地面观测资料为基础,运用EOF和REOF等统计学方法,研究了河北省草本植物物候期的变化特征及其对气候变暖的响应。结果表明:河北省草本植物展叶始期总体呈提前趋势,其中东部沿海平原提前趋势最大,中南部平原次之,西北部山区最小;黄枯始期主要表现出推迟趋势,生长季长度以延长趋势为主;春季气温对展叶始期的影响显著,河北省春季气温上升1℃,草本植物展叶始期提前4.1d;各站点生长季倾向率与年均温倾向率呈正相关,即年均温升幅大的站点,生长季延长的幅度也较大;草本植物物候变化特征及其对气候变暖的响应与木本植物基本一致,研究结果对丰富河北省物候与气候变化关系研究具有一定意义。     

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.     

Springtime in the city: exotic shrubs promote earlier greenup in urban forests

Despite the widespread recognition that urban areas are frequently dominated by exotic and invasive plants, the consequences of these changes in community structure have not been explicitly considered as an explanation for the pattern of advanced leaf phenology, or early greenup, reported in many urban areas. As such, we evaluated two hypotheses that could account for advanced greenup in forests along an urban to rural gradient: advanced phenology within individual species or differences in woody plant community. We monitored the spring leafing phenology of Aesculus glabra (Ohio buckeye), Lonicera maackii (Amur honeysuckle), and Acer negundo (box elder) in 11 forests spanning an urban to rural gradient in central Ohio, USA. From February to April 2006, we monitored these species, recorded woody plant composition, and documented daily minimum and maximum temperatures at each site. We found a weak but general trend of advanced phenology within species in more urban landscapes. Monthly average minimum temperatures were higher with increasing urbanization while monthly average maximum temperatures were similar across the urban to rural gradient. We also found evidence for shifts in woody plant communities along the urbanization gradient, mainly driven by the abundance of L. maackii, an invasive exotic species, in the more urban forests. Because L. maackii leafs out weeks earlier than native woody species and is very abundant in urban forests, we suggest that the invasion of forests by this species can generate earlier greenup of urban forests.     

Altered geographic and temporal variability in phenology in response to climate change

Aim In response to recent climate warming, numerous studies have reported an earlier onset of spring and, to a lesser degree, a later onset of autumn, both determined from phenological observations. Here, we examine whether these reported changes have affected the synchronization of events on a regional level by examining temporal and spatial variability in phenology. In particular, we study whether years with earlier springs are associated with an altered spatial variability in phenology. Location Germany and the United Kingdom. Methods Plant phenological observations of 35 different phases (events such as flowering and leafing) collected by the German Weather Service (1951–2002) and butterfly phenological records of 29 species collected by the UK Butterfly Monitoring Scheme (1976–2003) are used. In these long‐term records, we examine the temporal (year‐to‐year) variability and the spatial (geographic or between site) variability with particular emphasis on how they vary with time of the year and with earliness or lateness of the phase. Results Early phenological events (i.e. spring) are more variable than later events, both in time and in space, although the pattern is clearer for plants than for butterflies. Confirming previous results, we find a clear relationship between the mean date of spring and summer phases and the degree to which they have become earlier. The spatial variability of spring events is greater in warmer years that have faster plant development. However, late spring and summer events do not show a consistent relationship. Autumn events are somewhat more spatially variable in years characterized by later seasons. Main conclusions This is the first examination of spatial variability of plant and animal phenological events at a multinational scale. Earlier spring events are likely to be associated with increased spatial variability in plants, although this is unlikely to also be true for summer events. If species experience differential changes in geographic variation this may disrupt interactions among them, e.g. in food webs. On the other hand, these may offer advantages for mobile species. Further research on linked species is recommended.     

From observations to experiments in phenology research: investigating climate change impacts on trees and shrubs using dormant twigs

Background and Aims Climate change is advancing the leaf-out times of many plant species and mostly extending the growing season in temperate ecosystems. Laboratory experiments using twig cuttings from woody plant species present an affordable, easily replicated approach to investigate the relative importance of factors such as winter chilling, photoperiod, spring warming and frost tolerance on the leafing-out times of plant communities. This Viewpoint article demonstrates how the results of these experiments deepen our understanding beyond what is possible via analyses of remote sensing and field observation data, and can be used to improve climate change forecasts of shifts in phenology, ecosystem processes and ecological interactions.Scope The twig method involves cutting dormant twigs from trees, shrubs and vines on a single date or at intervals over the course of the winter and early spring, placing them in containers of water in controlled environments, and regularly recording leaf-out, flowering or other phenomena. Prior to or following leaf-out or flowering, twigs may be assigned to treatment groups for experiments involving temperature, photoperiod, frost, humidity and more. Recent studies using these methods have shown that winter chilling requirements and spring warming strongly affect leaf-out and flowering times of temperate trees and shrubs, whereas photoperiod requirements are less important than previously thought for most species. Invasive plant species have weaker winter chilling requirements than native species in temperate ecosystems, and species that leaf-out early in the season have greater frost tolerance than later leafing species.Conclusions This methodology could be extended to investigate additional drivers of leaf-out phenology, leaf senescence in the autumn, and other phenomena, and could be a useful tool for education and outreach. Additional ecosystems, such as boreal, southern hemisphere and sub-tropical forests, could also be investigated using dormant twigs to determine the drivers of leaf-out times and how these ecosystems will be affected by climate change.     

Convergence in leaf phenology traits of two understory ferns in the northwestern Iberian Peninsula

Aims Plants control leaf phenology to maximize annual photosynthetic product. Although ferns play an important ecological role in many habitats, especially forests, their phenology traits have been poorly studied. Here, we investigate the leaf phenology of two ferns of the forest understorey and analyse the relationship between the timing of leaf emergence and spore dispersal and the effect of between-year climatic variation.Methods We compared the leafing and sporing phenologies of two ferns with very large (>2 m), overwintering leaves: Culcita macrocarpa and Woodwardia radicans. We regularly monitored individuals of six populations in the northwestern Iberian Peninsula during a 3-year study. We studied eight phenology variables: leafing start date, leafing end date, leaf expansion time, number of new leaves per individual, between-individual synchrony, within-individual synchrony, percentage of fertile leaves and spore release date. We also determined leaf mass per area (LMA) and gathered data on air temperature and humidity.Important findings Both C. macrocarpa and W. radicans produce few leaves (~2 leaves individual^-1 year^-1), which expand simultaneously for a very long period (from midwinter to early summer), are retained for more than 1 year (37 and 19 months, respectively) and have relatively high LMAs. Such traits, together with large leaf size, have also been found in seed plants from the forest understorey and represent adaptations to this light-limited environment. Spores of both study ferns are simultaneously released in late winter, with little between-year variation caused by differences in air humidity. This remarkable similarity between species suggests that the convergence in timing of leaf emergence favours the convergence in timing of spore dispersal.     

Leaf phenology in relation to canopy closure in southern Appalachian trees

Leaf phenology varies markedly across tree species of temperate deciduous forests. Early leafing in spring may increase light capture and carbon gain prior to canopy closure, allowing saplings to survive in understory sites deeply shaded in midsummer. We quantified sapling leaf phenology for 18 tree species and seasonal variation in understory light availability at three sites along a ridge-slope-cove landform gradient in the Great Smoky Mountains National Park. Early leafing species (e.g., Aesculus flava, Carpinus caroliniana) broke bud an average of 24 d before late leafers (e.g., Magnolia fraseri, Nyssa sylvatica). Canopy closure occurred 14-18 d earlier and summer understory light was on average 63-74% lower on intermediate and mesic sites than on the xeric site. Early leafing species intercepted 45-80% of their growing season photon flux before canopy closure vs. 8-15% for late leafers. However, earlier leafing increased exposure to freezing temperatures by 5.5% per week near the mean time of bud break. Early leafing is strongly correlated with midsummer shade, risk of freezing temperatures, and distribution on mesic sites across a "main spectrum" of 15 deciduous species. Differences in leaf phenology and resultant impacts on spring carbon gain may help determine tree shade tolerance and distribution in southern Appalachian forests.     

Temperature sensitivity of spring vegetation phenology correlates to within-spring warming speed over the Northern Hemisphere

The inter-annual shift of spring vegetation phenology relative to per unit change of preseason temperature, referred to as temperature sensitivity (days °C⁻¹), quantifies the response of spring phenology to temperature change. Temperature sensitivity was found to differ greatly among vegetation from different environmental conditions. Understanding the large-scale spatial pattern of temperature sensitivity and its underlying determinant will greatly improve our ability to predict spring phenology. In this study, we investigated the temperature sensitivity for natural ecosystems over the North Hemisphere (north of 30°N), based on the vegetation phenological date estimated from NDVI time-series data provided by the Advanced Very High Resolution Radiometer (AVHRR) and the corresponding climate dataset. We found a notable longitudinal change pattern with considerable increases of temperature sensitivity from inlands to most coastal areas and a less obvious latitudinal pattern with larger sensitivity in low latitude area. This general spatial variation in temperature sensitivity is most strongly associated with the within-spring warming speed (WWS; r = 0.35, p < 0.01), a variable describing the increase speed of daily mean temperature during spring within a year, compared with other factors including the mean spring temperature, spring precipitation and mean winter temperature. These findings suggest that the same magnitude of warming will less affect spring vegetation phenology in regions with higher WWS, which might partially reflect plants’ adaption to local climate that prevents plants from frost risk caused by the advance of spring phenology. WWS accounts for the spatial variation in temperature sensitivity and should be taken into account in forecasting spring phenology and in assessing carbon cycle under the projected climate warming.     

北京地区木本春季物候特征与生物学特性的关系

植物物候是植物生活史中的重要性状,也是指示气候与自然环境变化的重要指标,现已成为全球变化领域的研究热点之一。传统物候研究多假设物候由气候因素决定,如气温、降水、光照等,并主要从植物物候的年际变化角度探讨了气候因素对物候特征的影响。然而,不同物种的物候存在较大差异表明植物物候还与自身生物学特性（如系统发育和功能性状）有关,但植物生物学特性如何影响植物物候仍缺乏深入研究。基于北京地区44种木本植物1965-2018年的展叶始期和开花始期观测资料,以展叶始期和开花始期的3类物候特征（平均物候期、物候对温度的响应敏感度和物候期的积温需求）为例,探究植物物候特征与系统发育和功能性状的关系。首先,利用系统发育信号Blomberg’s K和进化模型检验植物物候特征是否具有系统发育保守性,并通过系统发育信号表征曲线直观表达植物物候特征的进化模式;之后,利用广义估计方程分析植物生活型、传粉型与物候特征的关系,以揭示不同植物的资源利用方式及生存策略的差异。研究发现：（1）除展叶始期的温度敏感度外,其余物候特征的进化均受随机遗传漂变和自然选择力的共同作用,可推断物候特征具有系统发育保守性,即亲缘关系越近的物种物候特征越相似。（2）开花始期的系统发育信号强度比展叶始期更大,表明繁殖物候的系统发育可能比生长物候更保守。（3）植物展叶始期及其积温需求与生活型密切相关。灌木比乔木的展叶时间早、积温需求少。植物开花始期与传粉型相关,风媒植物开花显著早于虫媒植物。研究成果有助于深入理解物候变化的生物学机制,对于丰富物候学的理论研究有重要意义,同时对植物保护也具有重要的指导价值。     

Forecasting phenology under global warming

As a consequence of warming temperatures around the world, spring and autumn phenologies have been shifting, with corresponding changes in the length of the growing season. Our understanding of the spatial and interspecific variation of these changes, however, is limited. Not all species are responding similarly, and there is significant spatial variation in responses even within species. This spatial and interspecific variation complicates efforts to predict phenological responses to ongoing climate change, but must be incorporated in order to build reliable forecasts. Here, we use a long-term dataset (1953–2005) of plant phenological events in spring (flowering and leaf out) and autumn (leaf colouring and leaf fall) throughout Japan and South Korea to build forecasts that account for these sources of variability. Specifically, we used hierarchical models to incorporate the spatial variability in phenological responses to temperature to then forecast species'' overall and site-specific responses to global warming. We found that for most species, spring phenology is advancing and autumn phenology is getting later, with the timing of events changing more quickly in autumn compared with the spring. Temporal trends and phenological responses to temperature in East Asia contrasted with results from comparable studies in Europe, where spring events are changing more rapidly than are autumn events. Our results emphasize the need to study multiple species at many sites to understand and forecast regional changes in 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。     

Climatic effects on the breeding phenology and reproductive success of an arctic-nesting goose species

Climate warming is pronounced in the Arctic and migratory birds are expected to be among the most affected species. We examined the effects of local and regional climatic variations on the breeding phenology and reproductive success of greater snow geese ( Chen caerulescens atlantica ), a migratory species nesting in the Canadian Arctic. We used a long-term dataset based on the monitoring of 5447 nests and the measurements of 19 234 goslings over 16 years (1989–2004) on Bylot Island. About 50% of variation in the reproductive phenology of individuals was explained by spring climatic factors. High mean temperatures and, to a lesser extent, low snow cover in spring were associated with an increase in nest density and early egg-laying and hatching dates. High temperature in spring and high early summer rainfall were positively related to nesting success. These effects may result from a reduction in egg predation rate when the density of nesting geese is high and when increased water availability allows females to stay close to their nest during incubation recesses. Summer brood loss and production of young at the end of the summer increased when values of the summer Arctic Oscillation (AO) index were either very positive (low temperatures) or very negative (high temperatures), indicating that these components of the breeding success were most influenced by the regional summer climate. Gosling mass and size near fledging were reduced in years with high spring temperatures. This effect is likely due to a reduced availability of high quality food in years with early spring, either due to food depletion resulting from high brood density or a mismatch between hatching date of goslings and the timing of the peak of plant quality. Our analysis suggests that climate warming should advance the reproductive phenology of geese, but that high spring temperatures and extreme values of the summer AO index may decrease their reproductive success up to fledging.     

Leafing phenology and timing of egg laying in great tits Parus major and blue tits P. caeruleus

To evaluate the importance of tree leafing for the start of laying and clutch size of birds, we compared the breeding phenology of great tits Parus major and blue tits P. caeruleus between one coastal and two inland sites in the same geographical region. Because of the cooling influence of the sea, trees at the coastal site were known to initiate budburst about a week later than at the inland sites. During 5 years, breeding by the tits and the leaf phenology of birch Betula pendula , and oak Quercus robur were monitored. The leaf phenology of birch and oak explained a significant part of the between-year variation in the start of egg laying in blue and great tits, respectively. The tits started laying earlier at the sites with an early budburst, i.e. normally inland. However, leaf phenology was not an absolute cue to the start of laying, since blue tits laid earlier relative to leafing at the inland site than at the coastal site, and both tit species laid eggs earlier relative to leafing during late springs. In neither species was clutch size affected by leafing phenology. However, great tit females at the coastal site consistently produced fewer eggs than did those at the inland site. No such difference was found in the blue tits. Although leafing phenology may predict the start of laying in tits, other factors also influence its timing. These factors might include other cues, or differing life-history trade-offs depending on site or general climatic factors during the spring.     

An examination of the relationship between flowering times and temperature at the national scale using long-term phenological records from the UK

This paper examines the mean flowering times of 11 plant species in the British Isles over a 58-year period, and the flowering times of a further 13 (and leafing time of an additional 1) for a reduced period of 20 years. Timings were compared to Central England temperatures and all 25 phenological events were significantly related (P<0.001 in all but 1 case) to temperature. These findings are discussed in relation to other published work. The conclusions drawn from this work are that timings of spring and summer species will get progressively earlier as the climate warms, but that the lower limit for a flowering date is probably best determined by examining species phenology at the southern limit of their distribution. Received: 21 October 1999 / Revised: 27 January 2000 / Accepted: 27 January 2000     

Is water availability really the main environmental factor controlling the phenology of woody vegetation in the central Sahel?

Rainfall distribution and the soil moisture regime have been recognized to be the key drivers of the phenological rhythms in Sahelian woody plants, although different climate triggers have been assumed to be involved in determining the date of the onset of the phenophase. However, almost no comparisons have been made of the actual relative predictive power of these environmental factors. The aim of our study was to quantify the ability of several factors to predict phenophase occurrence in the dominant woody populations of northern Mali. Canopy leafing, flowering and fruiting were monitored from May 2005 to July 2007. Multiple logistic regressions were used to test the predictive power of cumulative rainfall, soil moisture, air temperature, air humidity and day length, with time lags of up to 2 months. Artificial variables derived from time lags observed in phenophases were included as predictors to account for possible auto-correlation and cross-correlation among phenophases. Surprisingly, a decrease in temperature associated with different time lags was most often found to be the strongest predictor of both leafing and reproductive phenophases. In Sahelian shrubs, morphological and physiological adaptations strongly contribute to the relative independence of their activity from water availability, leaf phenology being a way to adjust the plant water balance to current water availability and atmospheric water content. This study provides insight towards the development of a mechanistic understanding of phenological control in the Sahel, which is becoming increasingly important in the context of expected climate changes.     

31种木本植物开花物候与系统发育的关系

植物物候通常被认为是由环境因素,如降水、温度和日照长度所决定,然而环境因素是否是物候唯一的决定因素仍然存在很大争议。谱系结构表征了植物在进化上的顺序,该发育时序是否对物候产生影响,当前仍然未知。在调查2016年春季新疆乌鲁木齐市最常见的31种木本植物的初始开花时间、败花时间和开花持续时间的基础上,通过分析植物开花物候的分布特征、开花物候在乔灌木间的差别、以及植物谱系距离与开花物候距离间的关系,试图揭示植物的开花物候和物种谱系(进化)顺序间的关系。结果表明:(1)新疆乌鲁木齐市31种木本植物的初始开花时间为4月18日±9d、败花时间为5月5日±12d、开花持续时间为(16±8)d;(2)乔木的初始开花时间和败花时间的标准差分别均低于灌木,乔木开花物候相对灌木更稳定;(3)乔木的初始开花和败花时间均显著早于灌木(P0.05),但开花持续时间在两者间未有显著性差异(P0.05);(3)31种木本植物间的初始开花时间距离、败花时间距离和开花持续时间距离均与物种谱系距离存在显著线性回归关系(P0.05)。综上可知:乔灌木在垂直空间上的分化使得木本植物的开花物候在植物生活型间存在不同。对植物的开花物候,除已被证明的降水、温度和日照长度等环境因素的影响外,物种进化顺序也可能造成了它在植物种间、时间和空间上的变异。     

Land Surface Phenology in the Tropics: The Role of Climate and Topography in a Snow-Free Mountain

Leaf phenology represents a major temporal component of ecosystem functioning, and understanding the drivers of seasonal variation in phenology is essential to understand plant responses to climate change. We assessed the patterns and drivers of land surface phenology, a proxy for leafing phenology, for the meridional Espinhaço Range, a South American tropical mountain comprising a mosaic of savannas, dry woodlands, montane vegetation and moist forests. We used a 14-year time series of MODIS/NDVI satellite images, acquired between 2001 and 2015, and extracted phenological indicators using the TIMESAT algorithm. We obtained precipitation data from the Tropical Rainfall Measuring Mission, land surface temperature from the MODIS MOD11A2 product, and cloud cover frequency from the MODIS MOD09GA product. We also calculated the topographic wetness index and simulated clear-sky radiation budgets based on the SRTM elevation model. The relationship between phenology and environmental drivers was assessed using general linear models. Temporal displacement in the start date of the annual growth season was more evident than variations in season length among vegetation types, indicating a possible temporal separation in the use of resources. Season length was inversely proportional to elevation, decreasing 1.58 days per 100 m. Green-up and senescence rates were faster where annual temperature amplitude was higher. We found that water and light availability, modulated by topography, are the most likely drivers of land surface phenology in the region, determining the start, end and length of the growing season. Temperature had an important role in determining the rates of leaf development and the strength of vegetation seasonality, suggesting that tropical vegetation is also sensitive to latitudinal temperature changes, regardless of the elevational gradient. Our work improves the current understanding of phenological strategies in the seasonal tropics and emphasizes the importance of topography in shaping light and water availability for leaf development in snow-free mountains.