Spatial variation in breeding phenology at small spatial scales: A stochastic effect of population size

Spatial variation in phenology can occur at small spatial scales over which individuals can disperse or forage within one generation. Previous studies have assumed that variations in phenological peaks are caused by differences in abiotic environmental characteristics. However, environments should generally be similar among local habitats over small spatial scales. When the local population size is small, the phenological peak of the local population should be strongly affected by the variation in timing expressed by individuals. If a regional population consists of small local subpopulations (e.g., a metapopulation), the stochastic processes regulated by population sizes may explain the spatial variation in phenology. In this study, we quantitatively evaluated the extent of the spatial and annual variations in the breeding phenology of the forest green tree frog, Rhacophorus arboreus habiting a small area (<10 km²). The spatial variation in phenological peaks among 25 breeding sites was large over 6 years. This spatial variation was not explained by differences in air temperature or water depth. Randomization tests revealed that a large portion of the spatial variation could be explained by differences in population size, without considering site-specific factors. Annual variations in phenological peaks tended to be greater for smaller populations. These results imply that the stochastic process might have caused the spatial and annual variations in the phenological peaks of R. arboreus observed in the study region. Understanding spatiotemporal variation in phenology determined by stochastic process would be important to better predict interspecific interactions and (meta)population dynamics at small spatial scales.     

Below-ground respiratory responses of sugar maple and red maple saplings to atmospheric CO2 enrichment and elevated air temperature

The research described in this paper represents a part of a much broader research project with the general objective of describing the effects of elevated [CO2] and temperature on tree growth, physiological processes, and ecosystem-level processes. The specific objective of this research was to examine the below-ground respiratory responses of sugar maple (Acer saccharum Marsh.) and red maple (Acer rubrum L.) seedlings to elevated atmospheric [CO2] and temperature. Red maple and sugar maple seedlings were planted in the ground in each of 12 open-top chambers and exposed from 1994 through 1997 to ambient air or air enriched with 30 Pa CO2,< in combination with ambient or elevated (+4 °C) air temperatures. Carbon dioxide efflux was measured around the base of the seedlings and from root-exclusion zones at intervals during 1995 and 1996 and early 1997. The CO2 efflux rates averaged 0.4 μmol CO2 m-2 s-1 in the root-exclusion zones and 0.75 μmol CO2 m-2 s-1 around the base of the seedlings. Mineral soil respiration in root-exclusion zones averaged 12% higher in the high temperature treatments than at ambient temperature, but was not affected by CO2 treatments. The fraction of total efflux attributable to root + rhizosphere respiration ranged from 14 to 61% in measurements made around red maple plants, and from 35 to 62% around sugar maple plants. Root respiration rates ranged from 0 to 0.94 μmol CO2 s-1 m-2 of soil surface in red maple and from 0 to 1.02 in sugar maple. In both 1995 and 1996 root respiration rates of red maple were highest in high-CO2 treatments and lowest in high temperature treatments. Specific red maple root respiration rates of excised roots from near the soil surface in 1996 were also highest under CO2 enrichment and lowest in high temperature treatments. In sugar maple the highest rates of CO2 efflux were from around the base of plants exposed to both high temperature and high-CO2, even though specific respiration rates were< lowest for this species under the high temperature and CO2 enrichment regime. In both species, patterns of response to treatments were similar in root respiration and root mass, indicating that the root respiration responses were due in part to differences in root mass. The results underscore the need for separating the processes occurring in the roots from those in the forest floor and mineral soil in order to increase our understanding of the effects of global climate change on carbon sequestration and cycling in the below-ground systems of forests.     

Are Genes Randomly Distributed Over Space in Mature Populations of Sugar Maple (Acer saccharum Marsh.)?

Spatial autocorrelation analysis was employed to assess thegenetic structure in sugar maple (Acer saccharum). Nine allozymesfound at three isozyme loci (Fle1, Fle2, Pgi2) were analysedwithin six natural, mature populations. Based on the Gabrielnetwork, most autocorrelation coefficients (Moran's I) werenot significant. In two populations, however, the geographicdistribution of Fle2-107, Pgi2-100, and Pgi2-087 was positivelyautocorrelated. Non-random divergence of allozymes was alsoindicated in some populations by positive low order (short distance)and negative high order (long distance) Moran's I. On average,the spatial structure within the populations was weakly distinct.Copyright1993, 1999 Academic Press Spatial autocorrelation, Moran's I, Gabriel network, correlogram, allozymes, maple     

Climate change and extension of the Ginkgo biloba L. growing season in Japan

To understand the effects of climate change on the growing season of plants in Japan, we conducted trend analysis of phenological phases and examined the relationship between phenology and air temperatures. We used phenological data for Ginkgo biloba L., collected from 1953 to 2000. We defined the beginning and the end of the growing season (BGS and EGS) as the dates of budding and leaf fall, respectively. Changes in the air temperature in the 45 days before the date of BGS affected annual variation in BGS. The annual variation in air temperature over the 85 days before EGS affected the date of EGS. The average annual air temperature in Japan has increased by 1.3°C over the last four decades (1961–2000), and this increase has caused changes in ginkgo phenology. In the last five decades (1953–2000), BGS has occurred approximately 4 days earlier than previously, and EGS has occurred about 8 days later. Consequently, since 1953 the length of the growing season (LGS) has been extended by 12 days. Since around 1970, LGS and air temperatures have shown increasing trends. Although many researchers have stated that phenological events are not affected by the air temperature in the fall, we found high correlations not only between budding dates and air temperatures in spring but also between leaf‐fall dates and air temperatures in autumn. If the mean annual air temperature increases by 1°C, LGS could be extended by 10 days. We also examined the spatial distribution of the rate of LGS extension, but we did not find an obvious relationship between LGS extension and latitude.     

Topographic microclimates drive microhabitat associations at the range margin of a butterfly

The habitat associations of individuals underpin the dynamics of species distributions. Broad‐scale gradients in climate can alter habitat associations across species’ geographic ranges, but topographic heterogeneity creates local microclimates which could generate variation in habitat use at finer spatial scales. We examined the selection of microhabitats for egg‐laying by populations of a thermally‐constrained butterfly, the skipper Hesperia comma, across 16 sites with different regional temperatures and topographic microclimates. Using models of thermal microclimate, we examined how the association between eggs and warm bare ground microhabitats varied with ambient temperature, and predicted bare ground associations in 287 existing H. comma populations, to investigate the relative impacts of regional temperatures and topographic microclimates on microhabitat use. Eggs were most strongly associated with bare ground in relatively cool sites, indicating climate‐driven changes in microhabitat use. The majority of temperature variation between study sites was attributable to topographic microclimates rather than regional temperature differences, such that changes in microhabitat associations occurred principally between north‐ and south‐facing slopes within the same region. Predicted microhabitat associations across the UK distribution of H. comma showed that, due to the large temperature differences generated by topography, most of the between‐population variation in microhabitat use occurs locally within 5 km grid squares, with a smaller proportion occurring at a regional level between 5 km squares. Our findings show how microclimatic variation generated by topography alters the habitat associations of populations at fine spatial scales, suggesting that microclimate‐driven changes in habitat suitability could shape species’ distribution dynamics and their responses to environmental change.     

The effect of segregation of flowering time on fine-scale spatial genetic structure in an alpine-snowbed herb Primula cuneifolia

The flowering phenology of alpine-snowbed plants varies widely depending on the time of snowmelt. This variation may cause spatial and temporal heterogeneity in pollen dispersal, which in turn may influence genetic structure. We used spatial autocorrelation analyses to evaluate relative effect of segregation in flowering time and physical distance on fine-scale spatial genetic structure (SGS) of a snowbed herb Primula cuneifolia sampled in 10-m grids within a continuous snow patch (110 x 250 m) using nine allozyme loci. Although the individual flower lasts for 相似文献   

Dendroclimatology of sugar maple (Acer saccharum): Climate-growth response in a late-successional species

Sugar maple (Acer saccharum) is a shade-tolerant, late successional dominant species in the North American eastern deciduous forest. The objective of this study was to quantify the relationship between climate and radial growth in sugar maple and to identify spatial and temporal patterns in dendroclimatic response. We used a combination of archived sugar maple tree-ring chronologies and newly sampled sites to calculate dendroclimatic response of sugar maple at 13 sites in the United States and Canada. At all sites, sugar maple growth was significantly correlated to monthly temperature, precipitation, or Palmer Drought Severity Index. However, there were no generalizable patterns in sugar maple’s growth response to climate. Individual sites had unique dendroclimatic responses with respect to: a) which climatic variables were correlated to radial growth; b) what months had significant correlations between climate and radial growth; and c) what years had significant correlations between climate and radial growth. The individualistic dendroclimatic response of sugar maple appears to reflect a plastic response of the species to changes in climate perhaps related to its status as a strong competitor in late-successional forests. This ability to survive a wide range of environmental conditions may bode well for the species persistence under variable future climatic conditions. It also points to the need for more research on late-successional species in examining forest response to potential climate change scenarios because these species may be more resilient than early-successional species.     

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

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

Base-cation Cycling by Individual Tree Species in Old-growth Forests of Upper Michigan,USA

The influence of individual tree species on base-cation (Ca, Mg, K, Na) distribution and cycling was examined in sugar maple (Acer saccharum Marsh.), basswood (Tilia americana L.), and hemlock (Tsuga canadensis L.) in old-growth northern hardwood – hemlock forests on a sandy, mixed, frigid, Typic Haplorthod over two growing seasons in northwestern Michigan. Base cations in biomass, forest floor, and mineral soil (0–15 cm and 15–40 cm) pools were estimated for five replicated trees of each species; measured fluxes included bulk precipitation, throughfall, stemflow, litterfall, forest-floor leachate, mineralization + weathering, shallow-soil leachate, and deep-soil leachate. The three species differed in where base cations had accumulated within the single-tree ecosystems. Within these three single-tree ecosystems, the greatest quantity of base cations in woody biomass was found in sugar maple, whereas hemlock and basswood displayed the greatest amount in the upper 40 cm of mineral soil. Base-cation pools were ranked: sugar maple > basswood, hemlock in woody biomass; sugar maple, basswood > hemlock in foliage; hemlock > sugar maple, basswood in the forest floor, and basswood > sugar maple, hemlock in the mineral soil. Base-cation fluxes in throughfall, stemflow, the forest-floor leachate, and the deep-soil leachate (2000 only) were ranked: basswood > sugar maple > hemlock. Our measurements suggest that species-related differences in nutrient cycling are sufficient to produce significant differences in base-cation contents of the soil over short time intervals (<65 years). Moreover, these species-mediated differences may be important controls over the spatial pattern and edaphic processes of northern hardwood-hemlock ecosystems in the upper Great Lakes region.     

Soil Base Saturation Combines with Beech Bark Disease to Influence Composition and Structure of Sugar Maple-Beech Forests in an Acid Rain-Impacted Region

Sugar maple, an abundant and highly valued tree species in eastern North America, has experienced decline from soil calcium (Ca) depletion by acidic deposition, while beech, which often coexists with sugar maple, has been afflicted with beech bark disease (BBD) over the same period. To investigate how variations in soil base saturation combine with effects of BBD in influencing stand composition and structure, measurements of soils, canopy, subcanopy, and seedlings were taken in 21 watersheds in the Adirondack region of NY (USA), where sugar maple and beech were the predominant canopy species and base saturation of the upper B horizon ranged from 4.4 to 67%. The base saturation value corresponding to the threshold for Al mobilization (16.8%) helped to define the species composition of canopy trees and seedlings. Canopy vigor and diameter at breast height (DBH) were positively correlated (P < 0.05) with base saturation for sugar maple, but unrelated for beech. However, beech occupied lower canopy positions than sugar maple, and as base saturation increased, the average canopy position of beech decreased relative to sugar maple (P < 0.10). In low-base saturation soils, soil-Ca depletion and BBD may have created opportunities for gap-exploiting species such as red maple and black cherry, whereas in high-base saturation soils, sugar maple dominated the canopy. Where soils were beginning to recover from acidic deposition effects, sugar maple DBH and basal area increased progressively from 2000 to 2015, whereas for beech, average DBH did not change and basal area did not increase after 2010.     

Fine-scale spatial genetic structure of an endangered marsh herb, Caldesia grandis (Alismataceae)

The endangered marsh herb, Caldesia grandis, is native to China. We investigated the spatial structure of the genetic variation of three populations of C. grandis using RAPD markers and spatial autocorrelation analysis, based on the method of equal distance interval. A total of 157 individuals were sampled from four patches collected from the region of Hunan and Yunnan Provinces, China. Among the polymorphic bands generated by seven selective primers, polymorphic bands with frequencies ranging from 20 to 80% were used to calculate Moran's I spatial autocorrelation coefficient for each patch. We found significant spatial structure of genetic variation in the three patches in Bei Hai (BH) (patches BH-1 and BH-2) and Guai Hu (GH) (patch GH-1) populations of C. grandis (with significant positive autocorrelation within the short distance class). In contrast, the genetic variation in the Lang Pan Hu (LPH) population (patch LPH-1) was found to be randomly distributed. The different spatial distribution patterns may be attributed to environment differences. These results have implications for the conservation and management of this species, especially for sampling strategies for ex situ conservation.     

Spatial variability in prey phenology determines predator movement patterns and prey survival

Species have phenological variation among local habitats that are located at relatively small spatial scales. However, less studies have tested how this spatial variability in phenology can mediate intra-/inter-specific interactions. When predators track phenological variation of prey among local habitats, survival of prey within a local habitat strongly influenced by phenological synchrony with their conspecifics in adjacent habitats. Theory predicts that phenological synchrony among local habitats increases prey survival in local habitat within spatially structured environments because the predators have to make a habitat choice for foraging. Consequently, total survival of prey at regional scale should be higher. By using a spatially explicit field experiment, we tested above hypothesis using a prey–predator interaction between tadpole (Rhacophorus arboreus) and newt (Cynops pyrrhogaster). We established enclosures (≈regional scale) consisting of two tanks (≈local habitat scale) with different degree of prey phenological synchrony. We found that phenological synchrony of prey between tanks within each enclosure decreased the mean residence time of the predator in each tank, which resulted in higher survival of prey at a local habitat scale, supporting the theoretical prediction. Furthermore, individual-level variation in predator residence time explained the between-tank variation in prey survival in enclosures with phenological synchrony, implying that movement patterns of the predator can mediate variation in local population dynamics of their prey. However, total survival at each enclosure was not higher under phenological synchrony. These results suggest the importance of relative timing of prey phenology, not absolute timing, among local habitats in determining prey–predator interactions.     

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.     

Phenology of a northern hardwood forest canopy

While commonplace in other parts of the world, long‐term and ongoing observations of the phenology of native tree species are rare in North America. We use 14 years of field survey data from the Hubbard Brook Experimental Forest to fit simple models of canopy phenology for three northern hardwood species, sugar maple (Acer saccharum), American beech (Fagus grandifolia), and yellow birch (Betula alleghaniensis). These models are then run with historical meteorological data to investigate potential climate change effects on phenology. Development and senescence are quantified using an index that ranges from 0 (dormant, no leaves) to 4 (full, green canopy). Sugar maple is the first species to leaf out in the spring, whereas American beech is the last species to drop its leaves in the fall. Across an elevational range from 250 to 825 m ASL, the onset of spring is delayed by 2.7±0.4 days for every 100 m increase in elevation, which is in reasonable agreement with Hopkin's law. More than 90% of the variation in spring canopy development, and just slightly less than 90% of the variation in autumn canopy senescence, is accounted for by a logistic model based on accumulated degree‐days. However, degree‐day based models fit to Hubbard Brook data appear to overestimate the rate at which spring development occurs at the more southerly Harvard Forest. Autumn senescence at the Harvard Forest can be predicted with reasonable accuracy in sugar maple but not American beech. Retrospective modeling using five decades (1957–2004) of Hubbard Brook daily mean temperature data suggests significant trends (P≤0.05) towards an earlier spring (e.g. sugar maple, rate of change=0.18 days earlier/yr), consistent with other studies documenting measurable climate change effects on the onset of spring in both North America and Europe. Our results also suggest that green canopy duration has increased by about 10 days (e.g. sugar maple, rate of change=0.21 days longer/yr) over the period of study.     

Within-stand variation in silver birch (Betula pendula Roth) phenology

Key message

Within a local population genotypes differ in the timing of bud burst, but genotypes with early bud burst unfold their leaves slower, resulting in an equal period of carbon gain.

Abstract

The ability of local populations to cope with disturbances like adverse weather events or a changing climate depends on the genotypic richness of such populations, emphasising the importance of differences between genotypes in traits related to growth and survival at this scale. Due to their longevity, these differences are of special importance in trees, yet for trees, differences between genotypes within local populations remain unexplored. The phenological cycle is important in this respect, since a correct timing of phenological events is critical for growth and survival of trees, especially in environments with strong seasonality and changes in the timing of phenological events has consequences for, among others, net ecosystem productivity and the climate system as a whole. In this light accounting for differences in the timing of phenological events within species is currently identified as a research challenge. This study contributes to the knowledge of differences between genotypes on the small spatial scale of a local population. We examined the timing of phenological events of 15 micropropagated silver birch (Betula pendula Roth) genotypes representing a natural population. Measurements covered bud burst (7 years) and leaf unfolding in spring and chlorophyll degradation in autumn (2 years for both). These data were used to estimate the period of carbon gain. Differences between genotypes in the temperature sum required for bud burst were present, with genotypes showing ‘early’ (i.e. a low temperature sum requirement for bud burst) and ‘late’ bud burst across the 7-year study period. Differences were small in most years (i.e. 3 days), but differences of 16 days were recorded within the 7-year study period as well. Genotypes with ‘early’ bud burst were less sensitive to variations in environmental conditions in spring compared to genotypes with ‘late’ bud burst. Differences in bud burst were not carried over to the estimated period of carbon gain. Due to faster leaf expansion in genotypes with ‘late’ bud burst and the lack of differences between genotypes in autumn senescence the estimated period of carbon gain was similar among genotypes.     

Functional differences between summer and winter season rain assessed with MODIS‐derived phenology in a semi‐arid region

Questions: We asked several linked questions about phenology and precipitation relationships at local, landscape, and regional spatial scales within individual seasons, between seasons, and between year temporal scales. (1) How do winter and summer phenological patterns vary in response to total seasonal rainfall? (2) How are phenological rates affected by the previous season rainfall? (3) How does phenological variability differ at landscape and regional spatial scales and at season and inter‐annual temporal scales? Location: Southern Arizona, USA. Methods: We compared satellite‐derived phenological variation between 38 distinct 625‐km² landscapes distributed in the northern Sonoran Desert region from 2000 to 2007. Regression analyses were used to identify relationships between landscape phenology dynamics in response to precipitation variability across multiple spatial and temporal scales. Results: While both summer and winter seasons show increases of peak greenness and peak growth with more precipitation, the timing of peak growth was advanced with more precipitation in winter, while the timing of peak greenness was advanced with more precipitation in summer. Surprisingly, summer maximum growth was negatively affected by winter precipitation. The spatial variations between summer and winter phenology were similar in magnitude and response. Larger‐scale spatial and temporal variation showed strong differences in precipitation patterns; however the magnitudes of phenological spatial variability in these two seasons were similar. Conclusions: Vegetation patterns were clearly coupled to precipitation variability, with distinct responses at alternative spatial and temporal scales. Disaggregating vegetation into phenological variation, spanning value, timing, and integrated components revealed substantial complexity in precipitation‐phenological relationships.     

Local climate mediates spatial and temporal variation in carabid beetle communities in three forests in Mount Odaesan,Korea

1. Global environmental change can dramatically alter the composition of floral and faunal communities, and elucidating the mechanisms underlying this process is important for predicting its outcomes. Studies on global climate change have mostly focused on statistical summaries within wide spatial and temporal scales; less attention has been paid to variability in microclimates at narrower spatial and temporal scales. 2. The microclimate is the suite of climatic conditions measured in a local area. Environmental variables at the microclimatic scale can be critical for the ecology of organisms inhabiting each area. The effect of spatial and temporal changes in the microclimate on the ecology of carabid beetle communities in three sites on Mount Odaesan, Korea was examined. 3. Carabid beetle communities and quantified site‐specific environmental factors from measurements of air temperature, air humidity, light intensity and soil temperature over 5 years (2010–2015) were surveyed. 4. It was found that microclimatic variables and the patterns of temporal changes in carabid beetle communities differed between the three sites within the single mountain system. Microclimatic variables influencing temporal changes in beetle communities also differed between the sites. Therefore, it is suggested that variation in local microclimates affects spatial and temporal variation in carabid beetle communities at a local scale. 5. The present results demonstrate the importance of regular surveys of communities at local scales. Such surveys are expected to reveal an additional fraction of variation in communities and underlying processes that have been overlooked in studies of global community patterns and change.     

The influence of space in genetic-environmental relationships when environmental heterogeneity and seed dispersal occur at similar scale

We tested the importance of microenvironmental topographic parameters as predictors of emmer wheat genetic variation using three classes of single-locus (or at most several-loci) genetic markers (allozymes, glutenins, and qualitative traits) and two classes of markers of polygenic inheritance (phenological and morphological traits). Canonical correspondence analysis (CCA) and redundancy analysis (RDA) detected a significant effect of spatially structured environmental variation on genetic differences between plants for allozymes, glutenins, and quantitative morphological and phenological traits. However, after removing a spatial component of variation in partial CCA and partial RDA, the relationship of the remaining environmental variation with these genetic markers could be explained by chance alone, allowing us to rule out microniche topographic specialization in emmer wheat. Topographic autocorrelation exhibited a certain degree of similarity with genetic marker autocorrelation, indicating similar scales of environmental heterogeneity and seed flow. The detected population genetic structure agrees with one expected under isolation by distance as a result of limited gene flow. A negative relationship of genetic similarity with the logarithm of distance between plants was detected for both molecular markers and quantitative traits, which differed in the strength but not the pattern of association.     

The influence of altitude and urbanisation on trends and mean dates in phenology (1980–2009)

Long-term studies on urban phenology using network data are commonly limited by the small number of observation sites within city centres. Moreover, cities are often located on major rivers and consequently at lower altitudes than their rural surroundings. For these reasons, it is important (1) to go beyond a plain urban–rural comparison by taking the degree of urbanisation into account, and (2) to evaluate urbanisation and altitudinal effects simultaneously. Temporal phenological trends (1980–2009) for nine phenological spring events centred on the German cities of Frankfurt, Cologne and Munich were analysed. Trends of phenological onset dates were negative (i.e. earlier onset in phenology) for 96% of the 808 time series and significantly negative for 56% of the total number. Mean trends for the nine phenological events ranged between −0.23 days year⁻¹ for beech and −0.50 days year⁻¹ for hazel. The dependence of these trends and of mean dates on altitude and on the degree of urbanisation was explored. For mean dates, we demonstrated an earlier phenological onset at lower altitude and with a higher degree of urbanisation: altitude effects were highly significant and ranged between 1.34 days (100 m)⁻¹ (beech) and 4.27 days (100 m)⁻¹ (hazel). Coefficients for the log-transformed urban index were statistically significant for five events and varied greatly between events (coefficients from −1.74 for spruce to −5.08 for hazel). For trends in phenology, altitude was only significant for Norway maple, and no urban effects were significant. Hence, trends in phenology did not change significantly with higher altitudes or urbanised areas.     

Comparison of phenological characteristics for several woody plants in urban climates

Phenological properties of woody species were compared between two urban climates during 1997 and 1998. The study areas were Chungdam Park, Chungdam-dong, Kangnam-gu, Seoul (the urban center, 43 species) and Namhan-sansung Area, Sansung-ri, Joongbu-myon, Kwangju Gun, Kyonggi Province (the urban periphery, 16 species). Distance between these sites was 13.5 km. The differences of budding, foliation, and flowering times (1997 versus 1998) were 10.9, 3.2, and 7.4 days, respectively. Species that budded and flowered earlier were strongly influenced by Nuttonson’s Index (Tn) of February and March, but those with later dates were only weakly influenced. Unlike for budding and flowering times, foliation time was determined by air temperature or other factors in the leaf-growing season rather than by Tn. The Tn influence over phenology was stronger in shrubs and lianas than in trees. Phenophases in Chungdam Park appeared earlier than those in the Namhansansung area. The phenological differences between the two areas were 7.3 days in budding time, 8.3 days in foliation time, and 10.2 days in flowering time in mean values, with variations among species. Based on flowering-time data, the phenological variation between the two areas was equivalent to a 2.5° latitude difference. Budding time varied the most (20 days) inZelkova serrate, compared with only 3 days forPrunus padus. Differences in foliation time ranged from 15 days (inAlnus hirsute andStyrax obassia) to 0 days (P. padus). Flowering time differences were largest (24 days) inRhododendron mucronulatum and smallest (2 days) inP. padus. One can conclude that heat pollution in the urban center in Seoul severely changed phenology, and that sensitivity to that pollution differed among plant species.