Chilling outweighs photoperiod in preventing precocious spring development

It is well known that increased spring temperatures cause earlier onset dates of leaf unfolding and flowering. However, a temperature increase in winter may be associated with delayed development when species' chilling requirements are not fulfilled. Furthermore, photosensitivity is supposed to interfere with temperature triggers. To date, neither the relative importance nor possible interactions of these three factors have been elucidated. In this study, we present a multispecies climate chamber experiment to test the effects of chilling and photoperiod on the spring phenology of 36 woody species. Several hypotheses regarding their variation with species traits (successional strategy, floristic status, climate of their native range) were tested. Long photoperiods advanced budburst for one‐third of the studied species, but magnitudes of these effects were generally minor. In contrast to prior hypotheses, photosensitive responses were not restricted to climax or oceanic species. Increased chilling length advanced budburst for almost all species; its effect greatly exceeding that of photoperiod. Moreover, we suggest that photosensitivity and chilling effects have to be rigorously disentangled, as the response to photoperiod was restricted to individuals that had not been fully chilled. The results indicate that temperature requirements and successional strategy are linked, with climax species having higher chilling and forcing requirements than pioneer species. Temperature requirements of invasive species closely matched those of native species, suggesting that high phenological concordance is a prerequisite for successful establishment. Lack of chilling not only led to a considerable delay in budburst but also caused substantial changes in the chronological order of species' budburst. The results reveal that increased winter temperatures might impact forest ecosystems more than formerly assumed. Species with lower chilling requirements, such as pioneer or invasive species, might profit from warming winters, if late spring frost events would in parallel occur earlier.     

The shifting phenological landscape: Within‐ and between‐species variation in leaf emergence in a mixed‐deciduous woodland

Many organisms rely on synchronizing the timing of their life‐history events with those of other trophic levels—known as phenological matching—for survival or successful reproduction. In temperate deciduous forests, the extent of matching with the budburst date of key tree species is of particular relevance for many herbivorous insects and, in turn, insectivorous birds. In order to understand the ecological and evolutionary forces operating in these systems, we require knowledge of the factors influencing leaf emergence of tree communities. However, little is known about how phenology at the level of individual trees varies across landscapes, or how consistent this spatial variation is between different tree species. Here, we use field observations, collected over 2 years, to characterize within‐ and between‐species differences in spring phenology for 825 trees of six species (Quercus robur, Fraxinus excelsior, Fagus sylvatica, Betula pendula, Corylus avellana, and Acer pseudoplatanus) in a 385‐ha woodland. We explore environmental predictors of individual variation in budburst date and bud development rate and establish how these phenological traits vary over space. Trees of all species showed markedly consistent individual differences in their budburst timing. Bud development rate also varied considerably between individuals and was repeatable in oak, beech, and sycamore. We identified multiple predictors of budburst date including altitude, local temperature, and soil type, but none were universal across species. Furthermore, we found no evidence for interspecific covariance of phenology over space within the woodland. These analyses suggest that phenological landscapes are highly complex, varying over small spatial scales both within and between species. Such spatial variation in vegetation phenology is likely to influence patterns of selection on phenology within populations of consumers. Knowledge of the factors shaping the phenological environments experienced by animals is therefore likely to be key in understanding how these evolutionary processes operate.     

Predicted changes in the synchrony of larval emergence and budburst under climatic warming

Summary The impact of climatic warming on the synchrony of insect and plant phenologies was modelled in the case of winter moth (Operophtera brumata) and Sitka spruce (Picea sitchensis) in the Scottish uplands. The emergence of winter moth larvae was predicted with a thermal time requirement model and the budburst of Sitka spruce was predicted from a previously published model (Cannell and Smith 1983) based on winter chilling and thermal time. The date of emergence of winter moth larvae was predicted to occur earlier under climatic warming but the date of budburst of Sitka spruce was not greatly changed, resulting in decreased synchrony between larval emergence and budburst. The general question of how a change of climate might affect phenological synchrony and insect abundance is discussed.     

Partly decoupled tree-ring width and leaf phenology response to 20th century temperature change in Sweden

The recent warming trend, and associated shifts in growing season length, challenge the principle of uniformitarianism, i.e., that current relations are persistent over time, and complicates the uncritical inferences of past climate from tree-ring data. Here we conduct a comparison between tree-ring width chronologies of Pinus sylvestris L. (Scots pine), Picea abies (L.) Karst. (Norway spruce) and Betula pubescens Ehrh. (Downy birch) and phenological observations (budburst and leaf senescence) of Fagus sylvatica L. (European beech), Quercus robur L. (European oak), Betula sp. (Birch), Norway spruce and Scots pine) in Sweden to assess to what extent the tree-ring width–temperature relationship and the timing of phenological phases are affected by increased temperature. Daily meteorological observations confirm a prolongation of the thermal growing season, most consistently observed as an earlier onset of around 1–2 weeks since the beginning of the 20th century. Observations of budburst closely mimic this pattern, with budburst of the deciduous trees occurring 1–2.5 weeks earlier. In contrast to the changes seen in phenology and observational temperature data, the tree-ring width–temperature relationships remain surprisingly stable throughout the 20th century. Norway spruce, Scots pine and Downy birch all show consistently significant correlations with at least one 30 day-long window of temperature starting in late June–early July season. Norway spruce displays the largest degree of stability, with a consistent 60 day-long temperature window with significant correlation starting around Julian calendar day 150. Thus, our results suggest that the principle of uniformitarianism is not violated during the period covered by modern meteorological observations. Further research is needed to determine at what thresholds the temperature sensitivity of these species may alter or deteriorate as a consequence of the ongoing climate change.     

Climatic determinants of budburst seasonality in four temperate-zone tree species

Several physiological processes controlling tree phenology remain poorly understood and in particular bud dormancy. Many studies have emphasised the action of chilling temperatures in breaking dormancy. However, the effect of the preceding summer temperatures has rarely been investigated although there is some evidence that they may be involved in the settlement and intensity of dormancy as well as cold acclimation. In this paper, thermal time to budburst in relation to the duration of chilling outdoors, preceding summer temperatures and forcing temperatures was studied by outdoors experiments in seedlings of Platanus acerifolia , Vitis vinifera , Quercus pubescens and Castanea sativa . Results showed that temperatures of the preceding summer had no significant effect on the timing of budburst, P. acerifolia and Q. pubescens showed a very weak response to the duration of chilling, and the phenological characteristics of each species were found to be adapted to the climate conditions of its own geographical area. The phenological model used in this study explained 82–100% of the variance of the data without taking into account summer temperatures. Thus, although summer temperatures may be well involved in the intensity of dormancy and cold hardiness, they do not significantly affect budburst and therefore may not need to be considered in phenological models for predicting budburst.     

Spatial heterogeneity in the timing of birch budburst in response to future climate warming in Ireland

In order to predict the impact of future climate warming on trees it is important to quantify the effect climate has on their development. Our understanding of the phenological response to environmental drivers has given rise to various mathematical models of the annual growth cycle of plants. These models simulate the timing of phenophases by quantifying the relationship between development and its triggers, typically temperature. In addition, other environmental variables have an important role in determining the timing of budburst. For example, photoperiod has been shown to have a strong influence on phenological events of a number of tree species, including Betula pubescens (birch). A recently developed model for birch (DORMPHOT), which integrates the effects of temperature and photoperiod on budburst, was applied to future temperature projections from a 19-member ensemble of regional climate simulations (on a 25 km grid) generated as part of the ENSEMBLES project, to simulate the timing of birch budburst in Ireland each year up to the end of the present century. Gridded temperature time series data from the climate simulations were used as input to the DORMPHOT model to simulate future budburst timing. The results showed an advancing trend in the timing of birch budburst over most regions in Ireland up to 2100. Interestingly, this trend appeared greater in the northeast of the country than in the southwest, where budburst is currently relatively early. These results could have implications for future forest planning, species distribution modeling, and the birch allergy season.     

Within-canopy variation in photosynthetic capacity,SLA and foliar N in temperate broad-leaved trees with contrasting shade tolerance

Key message

The relative shade tolerance of T. cordata , F. sylvatica , and C. betulus in mature stands is based on different species-specific carbon and nitrogen allocation patterns.

Abstract

The leaf morphology and photosynthetic capacity of trees are remarkably plastic in response to intra-canopy light gradients. While most studies examined seedlings, it is not well understood how plasticity differs in mature trees among species with contrasting shade tolerance. We studied light-saturated net photosynthesis (A _max), maximum carboxylation rate (V _cmax), electron transport capacity (J _max) and leaf dark respiration (R _d) along natural light gradients in the canopies of 26 adult trees of five broad-leaved tree species in a mixed temperate old-growth forest (Fraxinus excelsior, Acer pseudoplatanus, Carpinus betulus, Tilia cordata and Fagus sylvatica), representing a sequence from moderately light-demanding to highly shade-tolerant species. We searched for species differences in the dependence of photosynthetic capacity on relative irradiance (RI), specific leaf area (SLA) and nitrogen per leaf area (N _a ). The three shade-tolerant species (C. betulus, T. cordata, F. sylvatica) differed from the two more light-demanding species by the formation of shade leaves with particularly high SLA but relatively low N _a and consequently lower area-based A _max, and a generally higher leaf morphological and functional plasticity across the canopy. Sun leaf morphology and physiology were more similar among the two groups. The three shade-tolerant species differed in their shade acclimation strategies which are primarily determined by the species’ plasticity in SLA. Under low light, T. cordata and F. sylvatica increased SLA, mass-based foliar N and leaf size, while C. betulus increased solely SLA exhibiting only low intra-crown plasticity in leaf morphology and N allocation patterns. This study with mature trees adds to our understanding of tree species differences in shade acclimation strategies under the natural conditions of a mixed old-growth forest.     

Are polyphagous geometrid moths with flightless females adapted to budburst phenology of local host species?

The majority of studies demonstrating local adaptation of insect herbivores involve sessile species, particularly those with a parthenogentic phase to their life history or endophagous "parasites" of plants. Current arguments suggest the strength of selection determines whether local adaptation can or cannot take place. Therefore local adaptation should not be limited to species with such traits. We studied the ability of three polyphagous geometrid moths with flightless adult females ( Erannis defoliaria , Operophtera brumata and O. fagata ) to synchronise their egg hatching with the budburst of a local host species in north east Scotland. A strong selection for hatching time is expected among generalist moths given the large variation in budburst phenology and an inability to hatch in synchrony with budburst decreases moth fitness substantially. In two successive seasons, we trapped emerging females from patches of five host species and recorded the temperature sum needed for 50% egg hatch of each brood laid by the trapped females. The hatching times of broods were compared against the average budburst time of the maternal host species in the study area. In addition, the trapping dates of each female were recorded. Only O. brumata showed synchrony with egg hatch and budburst which suggests local phenological adaptation to different host species . This could be maintained by selection and partial reproductive isolation between populations dwelling on different host species. No phenological adaptation was found in the other common geometrids of the study area.     

Photosynthetic traits of Sphagnum and feather moss species in undrained,drained and rewetted boreal spruce swamp forests

In restored peatlands, recovery of carbon assimilation by peat‐forming plants is a prerequisite for the recovery of ecosystem functioning. Restoration by rewetting may affect moss photosynthesis and respiration directly and/or through species successional turnover. To quantify the importance of the direct effects and the effects mediated by species change in boreal spruce swamp forests, we used a dual approach: (i) we measured successional changes in moss communities at 36 sites (nine undrained, nine drained, 18 rewetted) and (ii) photosynthetic properties of the dominant Sphagnum and feather mosses at nine of these sites (three undrained, three drained, three rewetted). Drainage and rewetting affected moss carbon assimilation mainly through species successional turnover. The species differed along a light‐adaptation gradient, which separated shade‐adapted feather mosses from Sphagnum mosses and Sphagnum girgensohnii from other Sphagna, and a productivity and moisture gradient, which separated Sphagnum riparium and Sphagnum girgensohnii from the less productive S. angustifolium, S. magellanicum and S. russowii. Undrained and drained sites harbored conservative, low‐production species: hummock‐Sphagna and feather mosses, respectively. Ditch creation and rewetting produced niches for species with opportunistic strategies and high carbon assimilation. The direct effects also caused higher photosynthetic productivity in ditches and in rewetted sites than in undrained and drained main sites.     

Arrested succession in logging gaps: is tree seedling growth and survival limiting?

Thirty years after selective timber harvest in the Kibale National Park, Uganda, many abandoned logging gaps are dominated by Acanthus pubescens, and show little forest recovery. To examine if this arrested successional state was caused by limited tree seedling growth and survival, we planted seedlings of four forest tree species (Albizia grandibracteata, Mimusops bagshawei, Prunus africana and Uvariopsis congensis) in A. pubescens‐dominated logging gaps and in control areas of adjacent forest. To assess if clearing A. pubescens facilitates forest regeneration, we planted seedlings of two species (A. grandibracteata and U. congensis) in small clearings cut within the logging gaps. We examined mortality, growth, herbivory and site characteristics among the treatments. Finally, we described the physical attributes of the A. pubescens‐dominated gaps. Seedlings of all the four species survived and grew equally well in A. pubescens and forest treatments, and most site characteristics were also similar. Seedlings planted in clearings grew more than in either forest or A. pubescens sites. Very few established trees were found in A. pubescens sites, and most of these were near the forest edges. We also discussed the role of elephants (Loxodonta africana) and collapsing A. pubescens canopies in the maintenance of an arrested successional state in these logging gaps.     

Asymmetric effects of cooler and warmer winters on beech phenology last beyond spring

In temperate trees, the timings of plant growth onset and cessation affect biogeochemical cycles, water, and energy balance. Currently, phenological studies largely focus on specific phenophases and on their responses to warming. How differently spring phenology responds to the warming and cooling, and affects the subsequent phases, has not been yet investigated in trees. Here, we exposed saplings of Fagus sylvatica L. to warmer and cooler climate during the winter 2013–2014 by conducting a reciprocal transplant experiment between two elevations (1,340 vs. 371 m a.s.l., ca. 6°C difference) in the Swiss Jura mountains. To test the legacy effects of earlier or later budburst on the budset timing, saplings were moved back to their original elevation shortly after the occurrence of budburst in spring 2014. One degree decrease in air temperature in winter/spring resulted in a delay of 10.9 days in budburst dates, whereas one degree of warming advanced the date by 8.8 days. Interestingly, we also found an asymmetric effect of the warmer winter vs. cooler winter on the budset timing in late summer. Budset of saplings that experienced a cooler winter was delayed by 31 days compared to the control, whereas it was delayed by only 10 days in saplings that experienced a warmer winter. Budburst timing in 2015 was not significantly impacted by the artificial advance or delay of the budburst timing in 2014, indicating that the legacy effects of the different phenophases might be reset during each winter. Adapting phenological models to the whole annual phenological cycle, and considering the different response to cooling and warming, would improve predictions of tree phenology under future climate warming conditions.     

Sap flux of five co-occurring tree species in a temperate broad-leaved forest during seasonal soil drought

In an old-growth forest in Central Germany, sap flux was studied in five broad-leaved tree species that were assumed to differ in drought sensitivity. Under moist soil conditions, average daily sap flux density (J _s) in the outermost xylem varied by a factor of 2.3 among the species (67–152 g cm⁻² per day, n=5 trees per species), and declined in the sequence Fagus sylvatica > Acer pseudoplatanus > Tilia cordata > Carpinus betulus > Fraxinus excelsior. Decreasing soil moisture content (Θ) resulted in linearly reduced J _s in four of the species. During a dry period, J _s was reduced by 44% in T. cordata, 39% in F. sylvatica, 37% in A. pseudoplatanus and 31% in C. betulus compared to sap flux at equal vapour pressure deficit (D) in the wet period. F. excelsior, the only ring-porous species studied, lacked a significant response in J _s to D and Θ. The relative reduction in water use during the dry period was not related to the assumed drought sensitivity of the species as inferred from their abundance in natural woodlands. J _s was positively correlated with tree diameter at breast height (DBH) in three species but decreased with DBH in two species. Dyeing experiments revealed that DBH accounted for 94% of the variation in sapwood area found in a bulk sample of all diffuse-porous trees. This suggests that DBH is a reliable estimator of sapwood area of temperate diffuse-porous species irrespective of species identity. In contrast, sap flux density was found to be greatly dependent on tree species. The estimated whole-plant water use for diffuse-porous trees of a given diameter (49 cm) ranged between 74 and 168 kg per day per species under moist soil conditions. Thus, in temperate mixed forests, species-specific differences in water use can result in a considerable spatial heterogeneity of canopy transpiration.     

Temperature alone does not explain phenological variation of diverse temperate plants under experimental warming

Anthropogenic climate change has altered temperate forest phenology, but how these trends will play out in the future is controversial. We measured the effect of experimental warming of 0.6–5.0 °C on the phenology of a diverse suite of 11 plant species in the deciduous forest understory (Duke Forest, North Carolina, USA) in a relatively warm year (2011) and a colder year (2013). Our primary goal was to dissect how temperature affects timing of spring budburst, flowering, and autumn leaf coloring for functional groups with different growth habits, phenological niches, and xylem anatomy. Warming advanced budburst of six deciduous woody species by 5–15 days and delayed leaf coloring by 18–21 days, resulting in an extension of the growing season by as much as 20–29 days. Spring temperature accumulation was strongly correlated with budburst date, but temperature alone cannot explain the diverse budburst responses observed among plant functional types. Ring‐porous trees showed a consistent temperature response pattern across years, suggesting these species are sensitive to photoperiod. Conversely, diffuse‐porous species responded differently between years, suggesting winter chilling may be more important in regulating budburst. Budburst of the ring‐porous Quercus alba responded nonlinearly to warming, suggesting evolutionary constraints may limit changes in phenology, and therefore productivity, in the future. Warming caused a divergence in flowering times among species in the forest community, resulting in a longer flowering season by 10‐16 days. Temperature was a good predictor of flowering for only four of the seven species studied here. Observations of interannual temperature variability overpredicted flowering responses in spring‐blooming species, relative to our warming experiment, and did not consistently predict even the direction of flowering shifts. Experiments that push temperatures beyond historic variation are indispensable for improving predictions of future changes in phenology.     

Frost sensitivity of Fagus sylvatica and co-occurring deciduous tree species at exposed sites

In Central-European forest stands on southerly to south-westerly exposed slopes of hills with shallow soil, European beech (Fagus sylvatica L.) loses its predominance over other deciduous tree species such as oak (Quercus spp.) or whitebeam (Sorbus aria [L.] Cr.) due to limited availability of water. To test whether winter frost is an additional factor affecting the performance of beech at such exposed sites, we measured the minimum air temperatures between the sampling dates, and determined the frost sensitivity of F. sylvatica and of co-occurring tree species (Quercus petraea [Matt.] Liebl., Quercus pubescens Willd., Quercus robur L., S. aria [L.] Cr.) eight times during the winter months, from October to March, at two sites in the low mountain ranges of Central Germany, which share the same climatic, topographic and edaphic conditions. At each site, two localities (stand margins and the interior of the stands) were compared. The frost sensitivity was determined in twigs of the trees using the electrolyte leakage method.Throughout the winter, frost sensitivity of F. sylvatica was higher at the stand margin than in the forest interior at one of the sites. At the other one, the frost sensitivity of S. aria was lower at the stand margin than within the stand. No consistent differences between the localities were detected in the Quercus species. Generally, the differences in frost sensitivity between the localities were only small; this was probably due to the slight differences in the minimum air temperature. Thus, it appears unlikely that, in exposed localities, the performance of the investigated tree species is limited by winter frost. More consistent differences were found among species, with F. sylvatica proving to be the most frost-sensitive, and S. aria, the least frost-sensitive species, particularly at the stand margins. The role of frost for the occurrence and performance of F. sylvatica should be further studied at the limit of its distribution along the boundary to the semihumid areas of Central Europe.     

Effects of constant and fluctuating temperature on time to budburst in Betula pubescens and its relation to bud respiration

 Effects of fluctuating and constant temperatures on budburst time, and respiration in winter buds were studied in Betula pubescens Ehrh. Dormant seedlings were chilled at 0°C for 4 months and then allowed to sprout in long days (LD, 24 h) at constant temperatures of 6, 9, 12, 15, 18 and 21°C, and at diurnally fluctuating temperatures (12/12 h, LD 24 h) with means of 9, 12, 15 and 18°C. No difference in thermal time requirements for budburst was found between plants receiving constant and fluctuating temperatures. The base temperature for thermal time accumulation was estimated to 1°C. Respiration in post-dormant (dormancy fully released) excised winter buds from an adult tree increased exponentially with temperature and was 20 times as high at 30°C than at 0°C. However, respiration in buds without scales was 30% higher at 0°C, and it was 2.7 times higher at 24°C than in intact buds. Thus, the tight bud scales probably constrain respiration and growth and are likely to delay budburst in spring. Arrhenius plots of the respiration data were biphasic with breaks at 13–15°C. However, this phase transition is unlikely to be associated with chilling sensitivity since the present species is hardy and adapted to a boreal climate. Received: 10 January 1997 / Accepted: 23 June 1997     

Do the capacity and kinetics for modification of xanthophyll cycle pool size depend on growth irradiance in temperate trees?

The present study investigated the interaction of growth irradiance (Q_int) with leaf capacity for and kinetics of adjustment of the pool size of xanthophyll cycle carotenoids (sum of violaxanthin, antheraxanthin and zeaxanthin; VAZ) and photosynthetic electron transport rate (J_max) after changes in leaf light environment. Individual leaves of lower‐canopy/lower photosynthetic capacity species Tilia cordata Mill. and upper canopy/higher photosynthetic capacity species Populus tremula L. were either illuminated by additional light of 500–800 µmol m^?2 s^?1 for 12 h photoperiod or enclosed in shade bags. The extra irradiance increased the total amount of light intercepted by two‐fold for the upper and 10–15‐fold for the lower canopy leaves, whereas the shade bags transmitted 45% of incident irradiance. In control leaves, VAZ/area, VAZ/Chl and J_max were positively associated with leaf growth irradiance (Q_int). After 11 d extra illumination, VAZ/Chl increased in all cases due to a strong reduction in foliar chlorophyll, but VAZ/area increased in the upper canopy leaves of both species, and remained constant or decreased in the lower canopy leaves of T. cordata. The slope for VAZ/area changes with cumulative extra irradiance was positively associated with Q_int only in T. cordata, but not in P. tremula. Nevertheless, all leaves of P. tremula increased VAZ/area more than the most responsive leaves of T. cordata. Shading reduced VAZ content only in P. tremula, but not in T. cordata, again demonstrating that P. tremula is a more responsive species. Compatible with the hypothesis of the role of VAZ in photoprotection, the rates of photosynthetic electron transport declined less in P. tremula than in T. cordata after the extra irradiance treatment. However, foliar chlorophyll contents of the exposed leaves declined significantly more in the upper canopy of P. tremula, which is not consistent with the suggestion that the leaves with the highest VAZ content are more resistant to photoinhibition. This study demonstrates that previous leaf light environment may significantly affect the adaptation capacity of foliage to altered light environment, and also that species differences in photosynthetic capacity and acclimation potentials importantly alter this interaction.     

A Lichenological Excursion to the West of Scotland

ABSTRACT

Background: Hybridisation can be a threat for the survival of a rare species because, in the case of insufficient numbers of appropriate mates, a rare form is much more likely to cross with a widespread taxon.

Aims: In the present study, we tested hypotheses concerning the level of hybridisation between endangered Betula humilis and its widespread congeners: B. pendula and B. pubescens as a function of habitat conditions.

Methods: We genotyped 312 individuals of three species using AFLP markers. B. humilis specimens were sampled in populations with low and high groundwater levels. Morphological identification of B. pubescens and B. pendula was verified using the Atkinson discriminant function.

Results: Altogether, 15 individuals (4.8%) were indicated as putative hybrids. The B. humilis hybrids were found in dry habitats and they could be classified as F1 or F2 generation. Tree hybrids could represent backcrosses to either B. pendula or B. pubescens.

Conclusions: Genetic analyses contradicted the idea that hybridisation between B. humilis and its close relatives was extensive. On the other hand, the presence of introgressed individuals in the populations in areas with low groundwater levels implied that pollen swamping might be a threat for declining B. humilis stands.     

Contrasting road effect signals in reproduction of long- versus short-lived amphibians

Despite an increasing understanding of the effects of roadways on amphibian populations, no studies have examined road effects on demographic traits other than survival. We predicted that road mortality could exert a disproportionate effect on fecundity in long-lived species due to shifts in population age structures to younger individuals of smaller size that produce commensurately smaller egg masses. To test this hypothesis, we assessed egg mass sizes of a long-lived amphibian (spotted salamander, Ambystoma maculatum) and short-lived one (wood frog, Rana sylvatica) in wetlands near and far from highways. Egg mass sizes of A. maculatum were smaller in wetlands near highways. In contrast, those of R. sylvatica were similar among wetlands regardless of the distance from highways. We conclude that paved highways with moderate traffic volume may be having important effects on populations of long-lived amphibians through mortality-mediated depression of reproduction.     

Leaf morphological variation in beech (Fagus sylvatica L.) populations in Greece and its relation to their post‐glacial origin

Diversity in leaf morphological traits was assessed in 38 Greek Fagus sylvatica populations, covering the distribution of the species in the country. The relationship between the post‐glacial origin of these populations and leaf morphology was investigated. The results showed a complex and geographically continuous morphological diversity pattern, influenced mainly by traits expressing leaf size, leaf shape and petiole length. Two simultaneous trends appeared to be responsible for the existing diversity pattern. One was geographical, with leaf types resembling F. sylvatica subsp. sylvatica occurring in the western part of the distribution of beech in Greece and types resembling F. sylvatica subsp. orientalis being dominant in the eastern part. A second trend seemed to be connected with the post‐glacial origin of the populations, as described by previous plastid DNA haplotype studies of the same trees. The genetic background and the possible adaptation of beech populations to different environmental conditions have resulted in a complex morphological pattern, especially in areas in which different post‐glacial lineages appear to meet. © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2011, 165 , 422–436.