Estimation of temperature and precipitation from morphological characters of dicotyledonous leaves

The utility of regression and correspondence models for deducing climate from leaf physiognomy was evaluated by the comparative application of different predictive models to the same three leaf assemblages. Mean annual temperature (MAT), mean annual precipitation (MAP), and growing season precipitation (GSP) were estimated from the morphological characteristics of samples of living leaves from two extant forests and an assemblage of fossil leaves. The extant forests are located near Gainesville, Florida, and in the Florida Keys; the fossils were collected from the Eocene Clarno Nut Beds, Oregon. Simple linear regression (SLR), multiple linear regression (MLR), and canonical correspondence analysis (CCA) were used to estimate temperature and precipitation. The SLR models used only the percentage of species having entire leaf margins as a predictor for MAT and leaf size as a predictor for MAP. The MLR models used from two to six leaf characters as predictors, and the CCA used 31 characters. In comparisons between actual and predicted values for the extant forests, errors in prediction of MAT were 0.6°-5.7°C, and errors in prediction of precipitation were 6-89 cm (=6-66%). At the Gainesville site, seven models underestimated MAT and only one overestimated it, whereas at the Keys site, all eight models overestimated MAT. Precipitation was overestimated by all four models at Gainesville, and by three of them at the Keys. The MAT estimates from the Clarno leaf assemblage ranged from 14.3° to 18.8°C, and the precipitation estimates from 227 to 363 cm for MAP and from 195 to 295 cm for GSP.     

Climatic reconstruction at the Miocene Shanwang basin, China, using leaf margin analysis, CLAMP, coexistence approach, and overlapping distribution analysis

The reconstruction of the climate in the Miocene Shanwang basin is an important link in understanding past climate and environmental changes in East Asia. A recent study showed that the mean annual temperature (MAT) estimates derived from leaf margin analysis (LMA) and the Climate Leaf Analysis Multivariate Program (CLAMP) conflicted with and were remarkably lower than those estimated by the coexistence approach (CA). Overlapping distribution analysis (ODA), a new method introduced here, is used to reconstruct the Shanwang Miocene climate based explicitly on local plant distribution data and associated meteorological stations. The Shanwang flora (17-15.2 Ma) suggests a MAT of 10.9-14.5°C and a mean annual precipitation (MAP) of 1107.3-1880.0 mm. This result is closer to the values derived from CLAMP and LMA than that obtained by CA. This report is the first comprehensive intercomparison of foliar physiognomic and nearest living relative climate proxies in a Chinese context and provides important cross validation of results.     

MuSICA, a CO2, water and energy multilayer, multileaf pine forest model: evaluation from hourly to yearly time scales and sensitivity analysis

The current emphasis on global climate studies has led the scientific community to set up a number of sites for measuring long‐term biospheric fluxes, and to develop a wide range of biosphere–atmosphere exchange models. This paper presents a new model of this type, which has been developed for a pine forest canopy. In most coniferous species the canopy layer is well separated from the understorey and several cohorts of needles coexist. It was therefore found necessary to distinguish several vegetation layers and, in each layer, several leaf classes defined not only by their light regime and wetness status but also by their age. This model, named MuSICA, is a multilayer, multileaf process‐based model. Each submodel is first independently parameterized using data collected at a EUROFLUX site near Bordeaux (Southwestern France). Particular care is brought to identify the seasonal variations in the various physiological parameters. The full model is then evaluated using a two‐year long data set, split up into 12 day‐type classes defined by the season, the weather type and the soil water status. Beyond the good overall agreement obtained between measured and modelled values at various time scales, several points of further improvement are identified. They concern the seasonal variations in the stomatal response of needles and the soil/litter respiration, as well as their interaction with soil or litter moisture. A sensitivity analysis to some of the model features (in‐canopy turbulent transfer scheme, leaf age classes, water retention, distinction between shaded and sunlit leaves, number of layers) is finally performed in order to evaluate whether significant simplifications can be brought to such a model with little loss in its predictive quality. The distinction between several leaf classes is crucial if one is to compute biospheric fluxes accurately. It is also evidenced that accounting for in‐canopy turbulent transfer leads to better estimates of the sensible heat flux.     

Estimation of deciduous forest leaf area index using direct and indirect methods

This study evaluated one semi-direct and three indirect methods for estimating leaf area index (LAI) by comparing these estimates with direct estimates derived from litter collection. The semi-direct method uses a thin metallic needle to count a number of contacts across fresh litter layers. One indirect method is based on the penetration of diffuse global radiation measured over the course of a day. The second indirect method uses the LAI-2000 plant canopy analyser (PCA) which measures diffuse light penetration from five different sky sectors simultaneously. The third indirect method uses the Demon portable light sensor to measure the penetration of direct beam sunlight at different zenith angles over the course of half a day. The Poisson model of gap frequency was applied to estimate plant area index (PAI) from observed transmittances using the second and third methods. Litter collection from 11 temperate decidous forests gave values of LAI ranging from 1.7 to 7.5. Estimates based on the needle method showed a significant linear relationship with LAI values obtained from litter collections but were systematically lower (by 6–37%). PAI estimates using all three indirect techniques (fixed light sensor system, LAI-2000 and Demon) showed a strong linear relationship with LAI derived from litter collection. Differences, averaged over all forest stands, between PAI estimates from each of the three indirect methods and LAI from litter collections were below 2%. If we consider that LAI=PAI–WAI (wood area index) then, all three indirect methods underestimated LAI by an additional factor close to the value of WAI. One reason could be a local clumping of architectural canopy components: in particular, the spatial dispositions of branchlets and leaves are not independent, leading to a non-random relationship between the distributions of these two canopy components.     

Towards a predictive description of forest canopies from litter properties

1. Specific leaf area (SLA), carbon isotope ratio (δ¹³C) and leaf nitrogen content (N) were measured in litter in 1993 and in the litter layer and canopy in 1996 in a pedunculate Oak ( Quercus robur ) wood at Wytham, Oxford.
2. The frequency distribution of SLA of leaves falling through the late summer and autumn of 1993 changed with a greater contribution of leaves of high SLA earlier in the period. For the same SLA leaves falling earlier in the season had less negative δ¹³C and higher N content than leaves falling later.
3. The frequency distribution of SLA of litter leaves was similar in both years but litter leaves had lower N and less negative δ¹³C in 1996.
4. In the canopy, there was a highly significant negative relationship between height in the canopy and SLA in green and senescent leaves. The relationship between SLA and both δ¹³C ratio and leaf N was different between green, senescent and litter leaves.
5. The frequency distribution of foliage in the canopy estimated from the frequency of SLA in the litter and the vertical pattern of SLA in the canopy was similar to the distribution measured by the 'drop-line' method.     

Fungal species diversity in juvenile and adult leaves of Eucalyptus globulus from plantations affected by Mycosphaerella leaf disease

In recent years, Mycosphaerella leaf disease (MLD) has become very common in Eucalyptus globulus plantations in Galicia, northwest Spain. The aetiology of MLD is complex and is associated with several species of Mycosphaerella and Teratosphaeria. A survey of the fungal mycobiota associated with juvenile and adult leaves and with leaf litter of the same trees in MLD‐affected plantations was made. The goal was to identify pathogens and endophytes, to determine whether the mycobiota of each leaf type differed and whether leaf litter might be a reservoir of MLD inoculum. Fungi belonging to 113 different species were isolated from the leaves of juvenile and adult trees sampled at 10 locations; 81 species occurred in juvenile and 65 in adult leaves. The average number of species obtained from juvenile leaves was significantly greater (P > 0.01) compared to adult leaves. This difference suggested that juvenile leaves are not only more susceptible to a group of pathogens, but to a wide range of fungi. Therefore, a general resistance mechanism might be lacking or be less effective in juvenile than in adult leaves. Several pathogenic species were identified in both leaf types. Leaf litter and living leaf mycobiotas were very different. However, some of the species they shared were MLD pathogens, suggesting that leaf litter could contribute to the inoculum of MLD.     

Decomposition and nitrogen release from the foliage litter of fir (Abies sachalinensis) and oak (Quercus crispula) under different forest canopies in Hokkaido, Japan

When two tree species co-occur, decomposition and nitrogen (N) release from the foliage litter depend on two factors: the forest floor conditions under each canopy type and the species composition of the litter. We conducted an experiment using fir and oak to answer several questions regarding decomposition beneath canopies of the two species and the effects of litter species composition on decomposition. We compared the rates of decomposition and N release from three different litters (fir needle, oak leaf, and a mixture of the two) in 1-mm-mesh litterbags on the forest floor under three different canopies (a 40-year-old fir plantation, large oak trees, and mixed fir and oak trees) in Hokkaido, Japan, over a 2-year period. Beneath each of these canopy types, the litter decomposition rate and percentage of N remaining in the litterbags containing a mixture of fir and oak litter were not significantly different from the expected values calculated for litterbags containing litter from a single tree species. Oak leaf litter decomposed significantly faster than fir needle litter beneath each canopy type. The litter decomposition rate was significantly higher beneath the fir canopy than under the oak canopy, and was intermediate under the mixed canopy of fir and oak. No net N release, that is, a decrease in the total N compared to the original amount, was detected from fir litter under each canopy type or from oak leaf litter beneath the oak canopy. N increased over the original amount in the fir litter beneath the oak canopy and the mixed canopy of fir and oak, but N was released from the oak litter under the fir canopy and the mixed canopy of fir and oak. These results suggest that oak leaf litter blown onto fir forest floor enhances nutrient cycling, and this might be a positive effect of a mixed stand of conifer and broad-leaved trees.     

Productivity and mineral cycling in an oak coppice forest 2. Annual net production of the forest

Annual net production was estimated in the secondary coppice forest near Tokyo, which was dominated by a deciduous oak,Quercus serrata Thunb. Lateral growth of stems and old branches was directly estimated by examining the annual rings for 35 shoots in a clear-cut quadrat of 10m×10m. Phytomasses of current organs were also weighed in the quadrat. Preharvest losses of current organs were determined by twelve 0.5 m² litter traps for fine litter and twelve 6 m² quadrats for woody litter. Branch production was also assessed indirectly by use of the stem-branch allometry and death of branches. The results of the indirect method were in sufficient agreement with the result of the direct one. Grazing loss of leaves from the canopy was estimated directly from the loss in leaf area and indirectly from the animal faeces caught by the litter traps. Net production of the canopy trees was 149 kg a⁻¹ year⁻¹, in which leaf production was 36.9 kg. Animals grazed about 14% of the leaf area by the end of the growing season. True consumption of leaves by animals was 7.6% of leaf production or 10% of leaf mass. Production of undergrowth, mainly a dwarf bamboo,Pleioblastus chino Makino, was 28 kg a⁻¹ year⁻¹, being 15% of the total stand production. Productivity of this forest was significantly higher than that of cool-temperate deciduous broadleaf forests.     

Leaf canopy as a dynamic system: ecophysiology and optimality in leaf turnover

BACKGROUND AND AIMS: In a leaf canopy, there is a turnover of leaves; i.e. they are produced, senesce and fall. These processes determine the amount of leaf area in the canopy, which in turn determines canopy photosynthesis. The turnover rate of leaves is affected by environmental factors and is different among species. This mini-review discusses factors responsible for leaf dynamics in plant canopies, focusing on the role of nitrogen. SCOPE: Leaf production is supported by canopy photosynthesis that is determined by distribution of light and leaf nitrogen. Leaf nitrogen determines photosynthetic capacity. Nitrogen taken up from roots is allocated to new leaves. When leaves age or their light availability is lowered, part of the leaf nitrogen is resorbed. Resorbed nitrogen is re-utilized in new organs and the rest is lost with dead leaves. The sink-source balance is important in the regulation of leaf senescence. Several models have been proposed to predict response to environmental changes. A mathematical model that incorporated nitrogen use for photosynthesis explained well the variations in leaf lifespan within and between species. CONCLUSION: When leaf turnover is at a steady state, the ratio of biomass production to nitrogen uptake is equal to the ratio of litter fall to nitrogen loss, which is an inverse of the nitrogen concentration in dead leaves. Thus nitrogen concentration in dead leaves (nitrogen resorption proficiency) and nitrogen availability in the soil determine the rate of photosynthesis in the canopy. Dynamics of leaves are regulated so as to maximize carbon gain and resource-use efficiency of the plant.     

Leaf litter inputs reinforce islands of nitrogen fertility in a lowland tropical forest

The role of lowland tropical forest tree communities in shaping soil nutrient cycling has been challenging to elucidate in the face of high species diversity. Previously, we showed that differences in tree species composition and canopy foliar nitrogen (N) concentrations correlated with differences in soil N availability in a mature Costa Rican rainforest. Here, we investigate potential mechanisms explaining this correlation. We used imaging spectroscopy to identify study plots containing 10–20 canopy trees with either high or low mean canopy N relative to the landscape mean. Plots were restricted to an uplifted terrace with relatively uniform parent material and climate. In order to assess whether canopy and soil N could be linked by litterfall inputs, we tracked litter production in the plots and measured rates of litter decay and the carbon and N content of leaf litter and leaf litter leachate. We also compared the abundance of putative N fixing trees and rates of free-living N fixation as well as soil pH, texture, cation exchange capacity, and topographic curvature to assess whether biological N fixation and/or soil properties could account for differences in soil N that were, in turn, imprinted on the canopy. We found no evidence of differences in legume communities, free-living N fixation, or abiotic properties. However, soils beneath high canopy N assemblages received ~ 60% more N via leaf litterfall due to variability in litter N content between plot types. The correlation of N in canopy leaves, leaf litter, and soil suggests that, under similar abiotic conditions, litterfall-mediated feedbacks can help maintain soil N differences among tropical tree assemblages in this diverse tropical forest.

     

The Contribution of Leaves from Different Levels within a Tomato Crop to Canopy Net Photosynthesis: An Experimental Examination of Two Canopy Models

The rates of net photosynthesis per unit ground area by a closedcanopy of tomato plants were measured over a range of naturallight flux densities. The canopy, of leaf area index 8.6, wasdivided into three horizontal layers of equal depth. On successivedays the canopy was progressively defoliated in layers fromthe ground upwards, allowing the photosynthetic contributionfrom individual leaf layers to be determined. The uppermostlayer, 23% of the total leaf area, assimilated 66% of the netCO₂ fixed by the canopy and accounted for a similar percentageof the total leaf respiration. Net photosynthesis versus light response curves for individualleaves from different positions within the canopy were alsoobtained. Leaf conductances to CO₂ transfer and the dark respirationrates of leaves from the uppermost leaf layer were approximatelyten times those from the lowest layer. The canopy data were analysed using a simple model which assumedthat the canopy was composed of leaves with identical photosyntheticand respiratory characteristics. The model fitted the data andallowed the characteristics of an ‘idealized’ leafto be estimated. The estimated values of the leaf light utilizationefficiency, ,and the leaf conductance CO₂ transfer, , were similarto values directly determined for individual leaves in the uppermostleaf layer and the estimated rate of leaf dark respiration,R_d, corresponded to measured rates for leaves much lower inthe canopy. The simple model may be used to examine gross effectsof crop environment on the leaf photosynthetic characteristicof an ‘idealized’ leaf, but cannot be used to predictaccurately canopy net photosynthesis from the photosyntheticand respiratory characteristics of any single real leaf. A moredetailed model, developed to allow explicitly for the observedvariation in and R_d within the canopy is appropriate for thispurpose.     

A comparison of direct and indirect methods for measuring leaf and surface areas of individual bushes

Indirect estimates of leaf area from measurements with three commercially available instruments (DEMON, LAI-2000 and Sunfleck Ceptometer) were compared with directly measured areas of individual Retama sphaerocarpa bushes. The three indirect methods gave good estimates of the total surface area of individual bushes. For the DEMON, the method of log-linear averaging of transmitted radiation gave estimates closer to directly measured surface area than the method of averaging transmission linearly. For the LAI-2000, estimated surface area index multiplied by canopy projected area gave the best agreement with directly measured values. For measurements with the Sunfleck Ceptometer, values of surface area estimated from the transmission of photosynthetic quantum flux density, without correcting for diffuse radiation, gave the best agreement with directly measured values. Surface areas estimated by the three instruments were not significantly different from directly measured total (leaf + branch + stem) surface areas. Leaf surface area could be calculated from estimated total surface area minus directly measured branch surface area. Measured branch surface area was linearly related to canopy projected area.     

Canopy photosynthetic production in a Japanese larch forest. II. Estimation of the canopy photosynthetic production

Seasonal changes and yearly gross canopy photosynthetic production were estimated for an 18 year old Japanese larch (Larix leptolepis) forest between 1982 and 1984. A canopy photosynthesis model was applied for the estimation, which took into account the effect of light interception by the non-photosynthetic organs. Seasonal changes in photosynthetic ability, amount of canopy leaf area and light environment within the canopy were also taken into account. Amount of leaf area was estimated by the leaf area growth of a single leaf. The change of light environment within the canopy during the growing season was estimated with a light penetration model and the leaf increment within the canopy. Canopy respiration and surplus production were calculated as seasonal and yearly values for the three years studied. Mean yearly estimates of canopy photosynthesis, canopy respiration and surplus production were 37, 13 and 23 tCO₂ ha⁻¹ year⁻¹, respectively. Vertical trend, seasonal changes and yearly values of the estimates were analyzed in relation to environmental and stand factors.     

Species-dependent responses of soil microbial properties to fresh leaf inputs in a subtropical forest soil in South China

Aims Forest disturbance from extreme weather events due to climate change could increase the contribution of fresh green leaves to the litter layer of soil and subsequently alter the composition and activity of the soil microbial properties and soil carbon cycling. The objective of this study was to compare the effect of naturally fallen litter and fresh leaves on the soil microbial community composition and their activities.Methods Fresh leaves and normal fallen litter were collected from four tree species (Pinus elliottii, Schima superba, Acacia mangium, A. auriculaeformis) in subtropical China and mixed with soil. Soil microbial community composition was determined using PLFAs, and its activity was quantified by soil respiration. During a 12-month period, the decomposition rate of litter was measured bimonthly using a litterbag method. Soil microbial samples were collected after 6 and 12 months. Soil respiration was measured monthly.Important findings We found that fresh leaves decomposed faster than their conspecific fallen litter. Although total microbial biomass and bacterial biomass were similar among treatments, soil fungal biomass was higher in fresh leaf than fallen litter treatments, resulting in greater values of the Fungal phospholipid fatty acids (PLFAs)/Bacterial PLFAs ratio. Fungal PLFA values were greater for Schima superba than the other species. The effect of litter type on soil respiration was species-dependent. Specifically, fallen litter released 35% more CO₂ than fresh leaves of the conifer P. elliottii. The opposite pattern was observed in the broadleaf species whose fresh leaf treatments emitted 17%–32% more CO₂ than fallen litter. Given future predictions that global climate change will cause more disturbances to forests, these results indicate that conifer and broadleaf forests in subtropical China may respond differently to increased fresh litter inputs, with net soil microbial respiration decreasing in conifer forests and increasing in broadleaf forests.     

LITTER FALL PRODUCTION IN RELATION TO ENVIRONMENTAL FACTORS IN NORTHEAST CHINA'S FORESTS

 在中国东北长白山、帽儿山、凉水、根河的主要森林类型中设置27个样地, 连续3年(2004~2006年)观测森林凋落物的生产量, 以研究我国东北地区森林凋落物产量及其与环境因子的关系。结果表明, 不同森林类型凋落物年产量存在显著差异, 针阔叶混交林显著高于落叶针叶林和常绿针叶林, 落叶针叶林、常绿针叶林、落叶阔叶林和针阔叶混交林的年平均产量分别为2 337、2 472、3 130和4 146 kg&;#8226;hm^–2; 树叶、枝条、繁殖器官和其它组分占总凋落量的平均比例为71%、22%、6%和1%, 不同森林类型凋落物组分的比例差异较大。森林凋落物产量主要受温度限制, 降水、森林类型和群落结构无显著影响。不同组分凋落物量的影响因素不同: 树叶凋落量主要受温度和森林类型的影响; 枝条凋落量主要受降水和蓄积量的影响; 而繁殖器官凋落量则与树种的繁殖特性以及年降水有关。各组分占总凋落量的比例主要受降水影响, 树叶占凋落物比例随降水增加而下降, 枝条所占比例很小, 表现出与叶相反的变化趋势。     

Sensitivity of leaf size and shape to climate within Acer rubrum and Quercus kelloggii

* Variation in the size and shape (physiognomy) of leaves has long been correlated to climate, and paleobotanists have used these correlations to reconstruct paleo-climate. Most studies focus on site-level means of largely nonoverlapping species sets. The sensitivity of leaf shape to climate within species is poorly known, which limits our general understanding of leaf-climate relationships and the value of intraspecific patterns for paleoclimate reconstructions. * The leaf physiognomy of two species whose native North American ranges span large climatic gradients (Acer rubrum and Quercus kelloggii) was quantified and correlated to mean annual temperature (MAT). Quercus kelloggii was sampled across a wide elevation range, but A. rubrum was sampled in strictly lowland areas. * Within A. rubrum, leaf shape correlates with MAT in a manner that is largely consistent with previous site-level studies; leaves from cold climates are toothier and more highly dissected. By contrast, Q. kelloggii is largely insensitive to MAT; instead, windy conditions with ample plant-available water may explain the preponderance of small teeth at high elevation sites, independent of MAT. * This study highlights the strong correspondence between leaf form and climate within some species, and demonstrates that intraspecific patterns may contribute useful information towards reconstructing paleoclimate.     

Limited evidence that mixing leaf litter accelerates decomposition or increases diversity of decomposers in streams of eastern Canada

Most studies of terrestrial litter decomposition in streams and rivers have used leaves from a single tree species, but leaf packs in streams in eastern North America are usually mixtures of two or more species. Litter mixtures may decay more quickly than either of the component species. If so, estimates of stream energy and nutrient budgets may be inaccurate. In northern Nova Scotia, Canada, we measured mass loss from binary mixtures (1:1 mass ratio) of leaf litter in mesh bags, using freshly fallen or air-dried litter from five species of canopy trees. We repeated the experiment eight times, in summer and fall, in two streams and a small river, over 3 years. In some trials we enumerated benthic invertebrate and fungal colonization of decaying litter. Although there were marked differences in mass loss rates among litter types, decomposition was accelerated in mixtures relative to the mean of the component species in only three of eight trials, and only in mixtures containing N-rich speckled alder leaves. Mixing yellow birch and red maple leaves inhibited decomposition. Diversity (Shannon–Weaver Index), species richness, and abundance of aquatic hyphomycete fungi, as indexed by conidial production, were never greater (and sometimes less) on litter mixtures than on the component species. Total numbers, taxonomic richness and diversity of benthic invertebrates generally, and litter-feeding species in particular, were not augmented by mixing litter types. Litter mixtures appear to dilute a preferred substrate with patches of a less preferred substrate. Our results provide only weak support for the contention that combining two litter types leads to acceleration of decomposition rates. Handling editor: K. Martens     

Field study of beech (t Fagus sylvatica L.) and melojo oak (t Quercus pyrenaica Willd) leaf litter decomposition in the centre of the Iberian Peninsula

The effect of the canopy on leaf decomposition of beech (t Fagus sylvatica) and melojo oak (t Quercus pyrenaica) was studied during a period of 660 days in a mountain forest of central Spain; response of leaves to leaching was also studied to determine the effects of rainfall after leaf fall. Beech leaves lost 5.8% of their weight by leaching, whereas melojo oak leaves lost 13%. Under both types of canopy, beech leaves showed almost no difference in their decomposition patterns, with decay constants of 0.31 and 0.32 respectively. Melojo oak leaves showed quite a different behaviour under both canopies; decay constant was 0.47 under t Quercus pyrenaica and 0.77 under beech canopy. Total immobilization of nitrogen was less in the melojo oak forest. Effects of summer dryness were sharper in the melojo oak forest, where decomposition stopped during the summer. This delay in the decomposition might have been due to the lower canopy density in the melojo oak forest. The decomposition patterns of the leaves of both species, under the canopy of the other species, suggests what might happen in mixed stands. Thus, the presence of melojo oaks in beech forest would increase decay and decrease nitrogen immobilization. The presence of beech trees in melojo oak forests would improve microclimatic conditions and increase decay, whereas beech litter on the soil would immobilizate more nitrogen.     

Field Evaluation of Some Models Estimating the Seasonal Pattern of Airborne Ascospores of Venturia inaequalis

A 6-year study was carried out to evaluate the accuracy of some models in estimating airborne ascospores of Venturia inaequalis . The proportion of the season's ascospores trapped on each discharge event was compared with the proportion of mature ascospores, estimated by the New Hampshire model or by some related models. The models differed from each other in the degree-day cumulation, accounting or not for the leaf litter wetness caused by rainfall or by deposition of atmospheric humidity. The New Hampshire model did not fit spore trappings well: 59% of the actual values fell outside the range of the estimates, and 83% of them were overestimates. The wide discrepancy between reality and estimates resulted from the effect of dryness: when many consecutive rainless days occurred, the proportion of ascospores trapped was constantly lower than the model estimates, due to a slowed spore maturation. The effect of dryness was evident during the greater part of the ascospore maturity season, irrespective of the proportion of the season's ascospores that had just matured when the dry period began. Models accounting for leaf litter wetness significantly improved estimates. Therefore, in the Po Valley, the accuracy of the New Hampshire model can be improved by accumulating degree-days only when leaf litter is wet.     

Sensitivity of leaf size and shape to climate: global patterns and paleoclimatic applications

? Paleobotanists have long used models based on leaf size and shape to reconstruct paleoclimate. However, most models incorporate a single variable or use traits that are not physiologically or functionally linked to climate, limiting their predictive power. Further, they often underestimate paleotemperature relative to other proxies. ? Here we quantify leaf-climate correlations from 92 globally distributed, climatically diverse sites, and explore potential confounding factors. Multiple linear regression models for mean annual temperature (MAT) and mean annual precipitation (MAP) are developed and applied to nine well-studied fossil floras. ? We find that leaves in cold climates typically have larger, more numerous teeth, and are more highly dissected. Leaf habit (deciduous vs evergreen), local water availability, and phylogenetic history all affect these relationships. Leaves in wet climates are larger and have fewer, smaller teeth. Our multivariate MAT and MAP models offer moderate improvements in precision over univariate approaches (± 4.0 vs 4.8°C for MAT) and strong improvements in accuracy. For example, our provisional MAT estimates for most North American fossil floras are considerably warmer and in better agreement with independent paleoclimate evidence. ? Our study demonstrates that the inclusion of additional leaf traits that are functionally linked to climate improves paleoclimate reconstructions. This work also illustrates the need for better understanding of the impact of phylogeny and leaf habit on leaf-climate relationships.