Extended season for northern butterflies

Butterflies are like all insects in that they are temperature sensitive and a changing climate with higher temperatures might effect their phenology. Several studies have found support for earlier flight dates among the investigated species. A comparative study with data from a citizen science project, including 66 species of butterflies in Sweden, was undertaken, and the result confirms that most butterfly species now fly earlier during the season. This is especially evident for butterflies overwintering as adults or as pupae. However, the advancement in phenology is correlated with flight date, and some late season species show no advancement or have even postponed their flight dates and are now flying later in the season. The results also showed that latitude had a strong effect on the adult flight date, and most of the investigated species showed significantly later flights towards the north. Only some late flying species showed an opposite trend, flying earlier in the north. A majority of the investigated species in this study showed a general response to temperature and advanced their flight dates with warmer temperatures (on average they advanced their flight dates by 3.8 days/°C), although not all species showed this response. In essence, a climate with earlier springs and longer growing seasons seems not to change the appearance patterns in a one-way direction. We now see butterflies on the wings both earlier and later in the season and some consequences of these patterns are discussed. So far, studies have concentrated mostly on early season butterfly–plant interactions but also late season studies are needed for a better understanding of long-term population consequences.     

The influence of life history traits on the phenological response of British butterflies to climate variability since the late‐19th century

Many species of plants and animals have advanced their phenology in response to climate warming in recent decades. Most of the evidence available for these shifts is based on data from the last few decades, a period coinciding with rapid climate warming. Baseline data is required to put these recent phenological changes in a long‐term context. We analysed the phenological response of 51 resident British butterfly species using data from 83 500 specimens in the collections of the Natural History Museum, London, covering the period 1880–1970. Our analysis shows that only three species significantly advanced their phenology between 1880 and 1970, probably reflecting the relatively small increase in spring temperature over this period. However, the phenology of all but one of the species we analysed showed phenological sensitivity to inter‐annual climate variability and a significant advancement in phenology in years in which spring or summer temperatures were warm and dry. The phenologies of butterfly species were more sensitive to weather if the butterfly species was early flying, southerly distributed, and a generalist in terms of larval diet. This observation is consistent with the hypothesis that species with greater niche breadth may be more phenologically sensitive than species with important niche constraints. Comparison of our results with post‐1976 data from the UK Butterfly Monitoring Scheme show that species flying early in the year had a greater rate of phenological advancement prior to the mid‐1970s. Additionally, prior to the mid‐1970s, phenology was influenced by temperatures in March or April, whereas since 1976, February temperature had a stronger influence on the phenology. These results suggest that early flying species may be approaching the limits of phenological advancement in response to recent climate warming.     

Butterfly phenology in Mediterranean mountains using space‐for‐time substitution

Inferring species' responses to climate change in the absence of long‐term time series data is a challenge, but can be achieved by substituting space for time. For example, thermal elevational gradients represent suitable proxies to study phenological responses to warming. We used butterfly data from two Mediterranean mountain areas to test whether mean dates of appearance of communities and individual species show a delay with increasing altitude, and an accompanying shortening in the duration of flight periods. We found a 14‐day delay in the mean date of appearance per kilometer increase in altitude for butterfly communities overall, and an average 23‐day shift for 26 selected species, alongside average summer temperature lapse rates of 3°C per km. At higher elevations, there was a shortening of the flight period for the community of 3 days/km, with an 8.8‐day average decline per km for individual species. Rates of phenological delay differed significantly between the two mountain ranges, although this did not seem to result from the respective temperature lapse rates. These results suggest that climate warming could lead to advanced and lengthened flight periods for Mediterranean mountain butterfly communities. However, although multivoltine species showed the expected response of delayed and shortened flight periods at higher elevations, univoltine species showed more pronounced delays in terms of species appearance. Hence, while projections of overall community responses to climate change may benefit from space‐for‐time substitutions, understanding species‐specific responses to local features of habitat and climate may be needed to accurately predict the effects of climate change on phenology.     

Tell me what you eat and I'll tell you when you fly: diet can predict phenological changes in response to climate change

Changes in phenology are correlated with climate change. However, we still struggle to understand the traits making species susceptible to climate change, and the implications of species' reactions for communities and food webs. Butterflies and moths are an ecologically important group that have shown pronounced phenological changes over the last decades. Tests using a > 150-year dataset from 566 European butterfly and moth species demonstrated that variation in phenological change was strongly related to traits describing plant-herbivore interactions (larval diet breadth, diet composition), and the life cycle. The results indicate that climate change related shifts in phenology are correlated with the seasonal availability and palatability of food plants. Lepidopterans feeding on herbaceous plants showed smaller shifts in flight periods but larger increases in voltinism than lepidopterans feeding on woody plants. Consequently, the effect of herbivorous lepidopterans may increase in herb-rich grassland ecosystems under warmer conditions, and not in forest ecosystems.     

Who flies first? – habitat‐specific phenological shifts of butterflies and orthopterans in the light of climate change: a case study from the south‐east Mediterranean

1. Insects undergo phenological change at different rates, showing no consistent trend between habitats, time periods, species or groups. Understanding how and why this variability occurs is crucial. 2. Phenological patterns of butterflies and Orthoptera were analysed using a novel approach of standardised major axis (SMA) analysis. It was investigated whether: (i) phenology (the mean date and duration of flight) of butterflies and Orthoptera changed from one survey (1998 and 1999 respectively) to another (2011), (ii) the rate at which phenology changed differed between taxa and (iii) phenological change was significantly different across habitat types (agriculture fields, grasslands, and forests). Using the 2011 dataset, we investigated relationships between habitat‐specific variables and species phenology. 3. For both groups, late‐emerging species had an advanced onset on the second survey while the duration showed no consistent trend for butterflies and did not change for Orthoptera. Although the rate at which phenology changed was consistent between the two groups, at the habitat level, a longer duration of flight period emerged for butterflies in agriculture fields while Orthoptera showed no differentiation in flight duration between habitats. We found an earlier emergence of butterflies in grasslands compared to forests, attributed to habitat‐specific temperature, whereas spatial variation in humidity had a significantly lower effect on butterflies' phenology in grasslands compared to forests. A gradual delay of butterfly appearances as the canopy cover increased was also found. 4. The utility of SMA analysis was demonstrated in phenological studies and evidence was detected that both habitat type and habitat‐specific variables refine species' phenological responses.     

Similarities in butterfly emergence dates among populations suggest local adaptation to climate

Phenology shifts are the most widely cited examples of the biological impact of climate change, yet there are few assessments of potential effects on the fitness of individual organisms or the persistence of populations. Despite extensive evidence of climate‐driven advances in phenological events over recent decades, comparable patterns across species' geographic ranges have seldom been described. Even fewer studies have quantified concurrent spatial gradients and temporal trends between phenology and climate. Here we analyse a large data set (~129 000 phenology measures) over 37 years across the UK to provide the first phylogenetic comparative analysis of the relative roles of plasticity and local adaptation in generating spatial and temporal patterns in butterfly mean flight dates. Although populations of all species exhibit a plastic response to temperature, with adult emergence dates earlier in warmer years by an average of 6.4 days per °C, among‐population differences are significantly lower on average, at 4.3 days per °C. Emergence dates of most species are more synchronised over their geographic range than is predicted by their relationship between mean flight date and temperature over time, suggesting local adaptation. Biological traits of species only weakly explained the variation in differences between space‐temperature and time‐temperature phenological responses, suggesting that multiple mechanisms may operate to maintain local adaptation. As niche models assume constant relationships between occurrence and environmental conditions across a species' entire range, an important implication of the temperature‐mediated local adaptation detected here is that populations of insects are much more sensitive to future climate changes than current projections suggest.     

Local environment,not local adaptation,drives leaf‐out phenology in common gardens along an elevational gradient in Acadia National Park,Maine

Premise of the Study

Climate‐driven changes in phenology are substantially affecting ecological relationships and ecosystem processes. The role of variation among species has received particular attention; for example, variation among species’ phenological responses to climate can disrupt trophic interactions and can influence plant performance. Variation within species in phenological responses to climate, however, has received much less attention, despite its potential role in ecological interactions and local adaptation to climate change.

Methods

We constructed three common gardens across an elevation gradient on Cadillac Mountain in Acadia National Park, Maine, to test population‐level responses in leaf‐out phenology in a reciprocal transplant experiment. The experiment included three native species: low bush blueberry (Vaccinium angustifolium), sheep's laurel (Kalmia angustifolia), and three‐toothed cinquefoil (Sibbaldiopsis tridentata).

Key Results

Evidence for local adaptation of phenological response to temperature varied among the species, but was weak for all three. Rather, variation in phenological response to temperature appeared to be driven by local microclimate at each garden site and year‐to‐year variation in temperature.

Conclusions

Population‐level adaptations in leaf‐out phenology appear to be relatively unimportant for these species in Acadia National Park, perhaps a reflection of strong genetic mixing across elevations, or weak differences in selection on phenological response to spring temperatures at different elevations. These results concur with other observational data in Acadia and highlight the utility of experimental approaches to understand the importance of annual and local site variation in affecting phenology both among and within plant species.     

Temperature‐related shifts in butterfly phenology depend on the habitat

Many species are becoming active earlier in the season as the climate becomes warmer. In parallel to phenological responses to climate change, many species have also been affected by habitat changes due to anthropogenic land use. As habitat type can directly affect microclimatic conditions, concurrent changes in climate and habitat could have interacting effects on the phenology of species. Temperature‐related shifts in phenology, however, have mostly been studied independent of habitat types. Here, I used long‐term data from a highly standardized monitoring program with 519 transects to study how phenology of butterflies is affected by ambient temperature and habitat type. I compared forests, agricultural areas and settlements, reflecting three major land use forms, and considered butterfly species that were observed in all three of these habitats. Seasonal appearance of the butterflies was affected both by the ambient temperature and the habitat type. As expected, warmer temperatures led to an overall advancement of the appearance and flight period of most species. Surprisingly, however, phenology of species was delayed in settlement habitats, even though this habitat type is generally associated with higher temperatures. A possible explanation is dispersal among habitat types, such that source–sink effects affect local phenology. When there is little productivity in settlement areas, observed butterflies may have immigrated from forest or agricultural habitats and thus appear later in settlements. My findings suggest that a spillover of individuals among habitats may affect phenology trends and indicate that phenological studies need to be interpreted in the context of habitat types. This becomes especially important when defining strategies to prevent or mitigate effects of climate and land‐use changes on phenology and abundance of species.     

Divergent phenological response to hydroclimate variability in forested mountain watersheds

Mountain watersheds are primary sources of freshwater, carbon sequestration, and other ecosystem services. There is significant interest in the effects of climate change and variability on these processes over short to long time scales. Much of the impact of hydroclimate variability in forest ecosystems is manifested in vegetation dynamics in space and time. In steep terrain, leaf phenology responds to topoclimate in complex ways, and can produce specific and measurable shifts in landscape forest patterns. The onset of spring is usually delayed at a specific rate with increasing elevation (often called Hopkins' Law; Hopkins, 1918), reflecting the dominant controls of temperature on greenup timing. Contrary with greenup, leaf senescence shows inconsistent trends along elevation gradients. Here, we present mechanisms and an explanation for this variability and its significance for ecosystem patterns and services in response to climate. We use moderate‐resolution imaging spectro‐radiometer (MODIS) Normalized Difference Vegetation Index (NDVI) data to derive landscape‐induced phenological patterns over topoclimate gradients in a humid temperate broadleaf forest in southern Appalachians. These phenological patterns are validated with different sets of field observations. Our data demonstrate that divergent behavior of leaf senescence with elevation is closely related to late growing season hydroclimate variability in temperature and water balance patterns. Specifically, a drier late growing season is associated with earlier leaf senescence at low elevation than at middle elevation. The effect of drought stress on vegetation senescence timing also leads to tighter coupling between growing season length and ecosystem water use estimated from observed precipitation and runoff generation. This study indicates increased late growing season drought may be leading to divergent ecosystem response between high and low elevation forests. Landscape‐induced phenological patterns are easily observed over wide areas and may be used as a unique diagnostic for sources of ecosystem vulnerability and sensitivity to hydroclimate change.     

Best environmental predictors of breeding phenology differ with elevation in a common woodland bird species

Temperatures in mountain areas are increasing at a higher rate than the Northern Hemisphere land average, but how fauna may respond, in particular in terms of phenology, remains poorly understood. The aim of this study was to assess how elevation could modify the relationships between climate variability (air temperature and snow melt‐out date), the timing of plant phenology and egg‐laying date of the coal tit (Periparus ater). We collected 9 years (2011–2019) of data on egg‐laying date, spring air temperature, snow melt‐out date, and larch budburst date at two elevations (~1,300 m and ~1,900 m asl) on a slope located in the Mont‐Blanc Massif in the French Alps. We found that at low elevation, larch budburst date had a direct influence on egg‐laying date, while at high‐altitude snow melt‐out date was the limiting factor. At both elevations, air temperature had a similar effect on egg‐laying date, but was a poorer predictor than larch budburst or snowmelt date. Our results shed light on proximate drivers of breeding phenology responses to interannual climate variability in mountain areas and suggest that factors directly influencing species phenology vary at different elevations. Predicting the future responses of species in a climate change context will require testing the transferability of models and accounting for nonstationary relationships between environmental predictors and the timing of phenological events.     

Greater phenological sensitivity to temperature on higher Scottish mountains: new insights from remote sensing

Mountain plants are considered among the species most vulnerable to climate change, especially at high latitudes where there is little potential for poleward or uphill dispersal. Satellite monitoring can reveal spatiotemporal variation in vegetation activity, offering a largely unexploited potential for studying responses of montane ecosystems to temperature and predicting phenological shifts driven by climate change. Here, a novel remote‐sensing phenology approach is developed that advances existing techniques by considering variation in vegetation activity across the whole year, rather than just focusing on event dates (e.g. start and end of season). Time series of two vegetation indices (VI), normalized difference VI (NDVI) and enhanced VI (EVI) were obtained from the moderate resolution imaging spectroradiometer MODIS satellite for 2786 Scottish mountain summits (600–1344 m elevation) in the years 2000–2011. NDVI and EVI time series were temporally interpolated to derive values on the first day of each month, for comparison with gridded monthly temperatures from the preceding period. These were regressed against temperature in the previous months, elevation and their interaction, showing significant variation in temperature sensitivity between months. Warm years were associated with high NDVI and EVI in spring and summer, whereas there was little effect of temperature in autumn and a negative effect in winter. Elevation was shown to mediate phenological change via a magnification of temperature responses on the highest mountains. Together, these predict that climate change will drive substantial changes in mountain summit phenology, especially by advancing spring growth at high elevations. The phenological plasticity underlying these temperature responses may allow long‐lived alpine plants to acclimate to warmer temperatures. Conversely, longer growing seasons may facilitate colonization and competitive exclusion by species currently restricted to lower elevations. In either case, these results show previously unreported seasonal and elevational variation in the temperature sensitivity of mountain vegetation activity.     

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.     

Phenological plasticity is a poor predictor of subalpine plant population performance following experimental climate change

Phenological shifts, changes in the seasonal timing of life cycle events, are among the best documented responses of species to climate change. However, the consequences of these phenological shifts for population dynamics remain unclear. Population growth could be enhanced if species that advance their phenology benefit from longer growing seasons and gain a pre-emptive advantage in resource competition. However, it might also be reduced if phenological advances increase exposure to stresses, such as herbivores and, in colder climates, harsh abiotic conditions early in the growing season. We exposed subalpine grasslands to ~3 K of warming by transplanting intact turfs from 2000 m to 1400 m elevation in the eastern Swiss Alps, with turfs transplanted within the 2000 m site acting as a control. In the first growing season after transplantation, we recorded species’ flowering phenology at both elevations. We also measured species’ cover change for three consecutive years as a measure of plant performance. We used models to estimate species’ phenological plasticity (the response of flowering time to the change in climate) and analysed its relationship with cover changes following climate change. The phenological plasticity of the 18 species in our study varied widely but was unrelated to their changes in cover. Moreover, early- and late-flowering species did not differ in their cover response to warming, nor in the relationship between cover changes and phenological plasticity. These results were replicated in a similar transplant experiment within the same subalpine community, established one year earlier and using larger turfs. We discuss the various ecological processes that can be affected by phenological shifts, and argue why the population-level consequences of these shifts are likely to be species- and context-specific. Our results highlight the importance of testing assumptions about how warming-induced changes in phenotypic traits, like phenology, impact population dynamics.     

Is climate warming more consequential towards poles? The phenology of Lepidoptera in Finland

The magnitude and direction of phenological shifts from climate warming could be predictably variable across the planet depending upon the nature of physiological controls on phenology, the thermal sensitivity of the developmental processes and global patterns in the climate warming. We tested this with respect to the flight phenology of adult nocturnal moths (3.33 million captures of 334 species) that were sampled at sites in southern and northern Finland during 1993–2012 (with years 2005–2012 treated as an independent model validation data set). We compared eight competing models of physiological controls on flight phenology to each species and found strong support for thermal controls of phenology in 66% of the species generations. Among species with strong thermal control of phenology in both the south and north, the average development rate was higher in northern vs. southern populations at 10 °C, but about the same at 15 and 20 °C. With a 3 °C increase in temperature (approximating A2 scenario of IPPC for 2090–2099 relative to 1980–1999) these species were predicted to advance their phenology on average by 17 (SE ± 0.3) days in the south vs. 13 (±0.4) days in the north. The higher development rates at low temperatures of poleward populations makes them less sensitive to climate warming, which opposes the tendency for stronger phenological advances in the north from greater increases in temperature.     

Diversity increases with elevation: empidine dance flies (Diptera,Empididae) challenge a predominant pattern

Tropical mountain forests are important reservoirs of biodiversity. They are usually species‐rich and often support endemic species making them prime targets for conservation effort. The aim of this study was to investigate elevational patterns of species diversity and phenology to provide a meaningful understanding of insects’ spatio‐temporal distributions along a tropical gradient of elevation. Our study focuses on the Empidinae communities (Diptera, Empididae) from Doi Inthanon (north Thailand, gradient 400–2556 m asl), sampled during two entire years (2006–2007, 2014). This group of insects is more diverse in temperate localities than in the tropics and we found that: (1) increase in altitude and latitude has a similar effect, so that the diversity of our model increases with elevation; (2) the phenology is strongly influenced by seasonality with a peak of diversity occurring during the transition between the dry and rainy seasons; (3) there is no phenological shift in the diversity peak with elevations; and (4) the species composition changes along the altitudinal gradient and through the year (high beta diversity). The increase in diversity with increasing elevation and the peak of diversity occurring at the transitional period both strongly coincide with abiotic factors, the decreasing temperature and the arrival of the monsoon, respectively.     

The phenology ofProcladius crassinervis (Zett),Procladius signatus (Zett) andProcladius choreus (Mg) (Diptera,Chironomidae) at Lake Hald,Denmark

In an investigation of the phenology of the chironomid species of Lake Hald, Denmark, a very late flight activity (September–November) of the univoltine speciesProcladius choreus (Mg.) was observed. Among observations of the phenological patterns of univoltine chironomids inhabiting lakes of the Baltic type in Denmark, the phenological pattern ofProcladius choreus stands alone (JÓNSSON, 1987; LARSEN, 1991). Further two species of the genusProcladius Skuse,viz. Procladius crassinervis (Zett) andProcladius signatus (Zett), were found in the investigation and the phenological pattern of the three congeneric species was clearly allochronic, withProcladius crassinervis flying in spring,Procladius signatus flying in summer andProcladius choreus flying in autumn.     

Elevational adaptation and plasticity in seedling phenology of temperate deciduous tree species

Phenological events, such as the initiation and the end of seasonal growth, are thought to be under strong evolutionary control because of their influence on tree fitness. Although numerous studies highlighted genetic differentiation in phenology among populations from contrasting climates, it remains unclear whether local adaptation could restrict phenological plasticity in response to current warming. Seedling populations of seven deciduous tree species from high and low elevations in the Swiss Alps were investigated in eight common gardens located along two elevational gradients from 400 to 1,700 m. We addressed the following questions: are there genetic differentiations in phenology between populations from low and high elevations, and are populations from the upper elevational limit of a species’ distribution able to respond to increasing temperature to the same extent as low-elevation populations? Genetic variation of leaf unfolding date between seedlings from low and high populations was detected in six out of seven tree species. Except for beech, populations from high elevations tended to flush later than populations from low elevations, emphasizing that phenology is likely to be under evolutionary pressure. Furthermore, seedlings from high elevation exhibited lower phenological plasticity to temperature than low-elevation provenances. This difference in phenological plasticity may reflect the opposing selective forces involved (i.e. a trade-off between maximizing growing season length and avoiding frost damages). Nevertheless, environmental effects were much stronger than genetic effects, suggesting a high phenological plasticity to enable tree populations to track ongoing climate change, which includes the risk of tracking unusually warm springs followed by frost.     

Predicting the sensitivity of butterfly phenology to temperature over the past century

Studies to date have documented substantial variation among species in the degree to which phenology responds to temperature and shifts over time, but we have a limited understanding of the causes of such variation. Here, we use a spatially and temporally extensive data set (ca. 48 000 observations from across Canada) to evaluate the utility of museum collection records in detecting broad‐scale phenology‐temperature relationships and to test for systematic differences in the sensitivity of phenology to temperature (days °C^?1) of Canadian butterfly species according to relevant ecological traits. We showed that the timing of flight season predictably responded to temperature both across space (variation in average temperature from site to site in Canada) and across time (variation from year to year within each individual site). This reveals that collection records, a vastly underexploited resource, can be applied to the quantification of broad‐scale relationships between species' phenology and temperature. The timing of the flight season of earlier fliers and less mobile species was more sensitive to temperature than later fliers and more mobile species, demonstrating that ecological traits can account for some of the interspecific variation in species' phenological sensitivity to temperature. Finally, we found that phenological sensitivity to temperature differed across time and space implying that both dimensions of temperature will be needed to translate species' phenological sensitivity to temperature into accurate predictions of species' future phenological shifts. Given the widespread temperature sensitivity of flight season timing, we can expect long‐term temporal shifts with increased warming [ca. 2.4 days °C^?1 (0.18 SE)] for many if not most butterfly species.     

Flowering phenology of Campanula on Mt Olynipos, Greece

We studied the fowering phenology of the genus Campanula (represented by nine species) along the elevation gradient of Mt Olympos (Greece) in order to assess whether there are elevation patterns at the genus level and whether these relate to patterns previously observed along such gradients at the community level. The traits examined were time and duration of the flowering period, flower life span, and duration of male and female flower phases. Floral attributes, such as number of flowers per plant, flower biomass, flower size, were also studied in order to examine whether they change with elevation or influence flower phenology. Flowering of Campanula species started in mid-May and ended in late September. The average duration of flowering of the genus was ca 27 d and the average floral longevity 4 d. In all but one species, the female phase lasted longer than the male. Campanula versicolor differed remarkably from all others in flower phenology and other floral trails. Nearly all Campanula populations studied had right or positively skewed flowering distributions indicating that flowering begins more abruptly than it ends. At the genus level, the time of flowering increased with elevation by 2 3d for every 100 m. Floral longevity also increased with elevation, by 0,2 d for every 100 m. Neither duration of flowering nor duration of the flower phases showed any consistent change with elevation. The same is true of the non–phenological floral traits examined. No trade–off between duration of flowering and flower life span or between structure and maintenance of flowers was apparent. The pattern of increasing floral longevity along the elevation gradient suggests a mechanism of compensation for reduecd pollinator availability.     

Phylogenetic conservatism and climate factors shape flowering phenology in alpine meadows

The study of phylogenetic conservatism in alpine plant phenology is critical for predicting climate change impacts; currently we have a poor understanding of how phylogeny and climate factors interactively influence plant phenology. Therefore, we explored the influence of phylogeny and climate factors on flowering phenology in alpine meadows. For two different types of alpine plant communities, we recorded phenological data, including flowering peak, first flower budding, first flowering, first fruiting and the flowering end for 62 species over the course of 5 years (2008–2012). From sequences in two plastid regions, we constructed phylogenetic trees. We used Blomberg’s K and Pagel’s lambda to assess the phylogenetic signal in phenological traits and species’ phenological responses to climate factors. We found a significant phylogenetic signal in the date of all reproductive phenological events and in species’ phenological responses to weekly day length and temperature. The number of species in flower was strongly associated with the weekly day lengths and followed by the weekly temperature prior to phenological activity. Based on phylogenetic eigenvector regression (PVR) analysis, we found a highly shared influence of phylogeny and climate factors on alpine species flowering phenology. Our results suggest the phylogenetic conservatism in both flowering and fruiting phenology may depend on the similarity of responses to external environmental cues among close relatives.