Ecophysiological responses of two closely related Magnoliaceae genera to seasonal changes in subtropical China

Aims Plants use a variety of hydraulic strategies to adapt to seasonal drought that differ by species and environmental conditions. The early-diverging Magnoliaceae family includes two closely related genera with contrasting leaf habits, Yulania (deciduous) and Michelia (evergreen), which naturally inhabit temperate and tropical regions, respectively. Here, we evaluate the hydraulic strategy of species from both genera that have been ex situ conserved in a subtropical region to determine how they respond to the novel cool–dry season climatic pattern.Methods We measured ecophysiological traits in five Michelia and five Yulania species conserved in the South China Botanical Garden in both wet and dry season conditions and monitored the whole-year sap flow for four of these species.Important findings We found that Magnoliaceae species that have been ex situ conserved in a subtropical climate did not suffer from excessive water stress due to the mild drought conditions of the dry season and the ecophysiological adjustments the species made to avoid this stress, which differed by leaf habit. Specifically, deciduous species completely shed their leaves during the dry season, while evergreen species decreased their turgor loss points, dry mass based photosynthetic rates, stomatal conductance and specific leaf areas (SLAs) compared to wet season measurements. In comparing the two distinct leaf habits during the wet season, the leathery-leaved evergreen species had higher leaf hydraulic conductance and leaf to sapwood area ratios than the papery-leaved deciduous species, while the deciduous species had greater hydraulic conductivity calculated on both a stem and leaf area basis, dry mass based photosynthetic rates, leaf nutrients, SLAs and stomatal sizes than the evergreen species. Interestingly, species from both genera maintained similar sap flow in the wet season. Both photosynthetically active radiation and vapour pressure deficit affected the diurnal patterns of sap flow in the wet season, while only vapour pressure deficit played a dominant role in the dry season. This study reveals contrasting hydraulic strategies in Yulania and Michelia species under subtropical seasonal conditions, and suggests that these ecophysiological adjustments might be affected more by leaf habit than seasonality, thus reflecting the divergent evolution of the two closely related genera. Furthermore, we show that Magnoliaceae species that are ex situ conserved in a subtropical climate are hydraulically sound, a finding that will inform future conservation efforts of this ancient family under the threat of climatic change.     

Leaf habit does not predict leaf functional traits in cerrado woody species

Plant species with a high leaf life span (LLS) commonly have a low specific leaf area (SLA), leaf nitrogen per unit mass (N), and phosphorous concentration (P), whereas species with low LLS have a high SLA, N and P. However, LLS tends to be longer in species growing in low-nutrient soils and, therefore, differences in LLS and other leaf traits may not be consistent with a plant classification according to leaf habit. Here we investigated whether leaf habit is consistent with leaf economic spectrum trade-offs in cerrado (a Neotropical savanna) woody species. We analyzed the SLA, N and P of 125 woody species with a distinct leaf habit (deciduous, semideciduous, brevideciduous or evergreen). We also gathered data on the LLS (33 species), maximum net photosynthesis per leaf area (A_area, 56 species) and per leaf mass (A_mass, 31 species), comprising the most extensive database analyzed so far for the cerrado. Differences among leaf habit groups were tested using generalized linear mixed models and ANOVA. We did not find differences in SLA and N among species with a distinct leaf habit, but deciduous species had a higher leaf P concentration than evergreens. Species did not differ in LLS and A_mass, but A_area varied among groups. Semideciduous species had higher A_area values than deciduous and brevideciduous species, but all other groups had similar A_area values. Because of the small difference in the LLS, SLA, leaf N, leaf P and maximum net photosynthesis, we argue that deciduous, brevideciduous, semideciduous and evergreen species may not constitute different functional groups in cerrado woody species.     

Deciduous and evergreen trees differ in juvenile biomass allometries because of differences in allocation to root storage

Background and Aims

Biomass partitioning for resource conservation might affect plant allometry, accounting for a substantial amount of unexplained variation in existing plant allometry models. One means of resource conservation is through direct allocation to storage in particular organs. In this study, storage allocation and biomass allometry of deciduous and evergreen tree species from seasonal environments were considered. It was expected that deciduous species would have greater allocation to storage in roots to support leaf regrowth in subsequent growing seasons, and consequently have lower scaling exponents for leaf to root and stem to root partitioning, than evergreen species. It was further expected that changes to root carbohydrate storage and biomass allometry under different soil nutrient supply conditions would be greater for deciduous species than for evergreen species.

Methods

Root carbohydrate storage and organ biomass allometries were compared for juveniles of 20 savanna tree species of different leaf habit (nine evergreen, 11 deciduous) grown in two nutrient treatments for periods of 5 and 20 weeks (total dry mass of individual plants ranged from 0·003 to 258·724 g).

Key Results

Deciduous species had greater root non-structural carbohydrate than evergreen species, and lower scaling exponents for leaf to root and stem to root partitioning than evergreen species. Across species, leaf to stem scaling was positively related, and stem to root scaling was negatively related to root carbohydrate concentration. Under lower nutrient supply, trees displayed increased partitioning to non-structural carbohydrate, and to roots and leaves over stems with increasing plant size, but this change did not differ between leaf habits.

Conclusions

Substantial unexplained variation in biomass allometry of woody species may be related to selection for resource conservation against environmental stresses, such as resource seasonality. Further differences in plant allometry could arise due to selection for different types of biomass allocation in response to different environmental stressors (e.g. fire vs. herbivory).     

木兰科常绿与落叶物种叶片构建策略的差异

为探究木兰科(Magnoliaceae)常绿与落叶物种叶片构建的生理生态策略,选取黄山木兰(Yulaniacylindrica)、玉兰(Y.denudata)和鸡公山玉兰(Y. jigongshanensis) 3种落叶物种,以及荷花玉兰(Magnolia grandiflora)、含笑花(Michelia figo)、石碌含笑(M. shiluensis) 3种常绿物种,对其叶片构建成本和叶片寿命相关的性状进行比较。结果表明,木兰科3落叶种的单位叶片面积成本(CCarea)显著低于3常绿种,但落叶和常绿物种的叶片质量成本(CCmass)差异不显著。落叶物种的叶氮、磷含量(Nmass,Pmass)和比叶面积(SLA)均显著高于常绿物种,而叶片寿命(LLS)显著低于常绿物种。CCarea与LLS呈显著正相关,Nmass、Pmass和SLA均与LLS呈显著负相关。这说明木兰科玉兰属落叶物种单位面积叶片构建成本小于常绿物种;落叶物种叶片寿命短,但采取低成本构建策略,提高比叶面积获得更多光资源,增加营养积累,也揭示了玉兰属落叶物种适应北亚热带较短的生长季和较低水热条件的生理生态策略。     

Leaf litter carbon,nitrogen, and phosphorus stoichiometric patterns as related to climatic factors and leaf habits across Chinese broad-leaved tree species

Our understanding of how climate and leaf habit (evergreen vs. deciduous) drive leaf litter carbon (C), nitrogen (N), and phosphorus (P) stoichiometric patterns is largely limited but is particularly important for broad-leaved forests, since the forest is sensitive to climate change. Here, we investigated leaf litter C, N, and P stoichiometric ratios of broad-leaved tree species in relation to climate and leaf habit using previous publications and our additional samplings across China. We found that mean leaf litter C:N across Chinese broad-leaved tree species was within the range of the global flora, whereas C:P was lower and N:P higher. Evergreen species displayed higher leaf litter C:N, C:P, and N:P than their deciduous counterparts. Both leaf litter C:P and N:P for all species pooled were negatively correlated with latitude, driven by mean annual precipitation (MAP) and mean annual temperature, respectively, while leaf litter C:N displayed no clear latitudinal trend. The direction and magnitude of leaf litter C, N, and P stoichiometric ratios in response to climate diverged between leaf habits. For example, evergreen leaf litter C:N was negatively correlated with MAP, while deciduous counterparts did not respond significantly to MAP. We conclude that leaf litter C, N, and P stoichiometric ratios shifted along the climatic gradient, and the strength of such shifts differed between leaf habits. Therefore, leaf litter stoichiometric patterns across leaf habits suggest that any climate change-driven shift in species distribution may potentially alter the ecosystem’s nutrient cycling processes of evergreen- and deciduous-dominated broad-leaved forests differentially.     

Seasonal occurrence and distribution of myxomycetes on different types of leaf litter in a warm temperate forest of western Japan

The seasonal occurrence and distribution of myxomycetes on different types of newly defoliated leaf litter were examined in a secondary forest in a warm temperate region of western Japan. The two types of leaf litter (deciduous trees, Prunus verecunda and Quercus variabilis, and evergreen trees, Q. glauca and Cinnamomum camphora) were incubated in trays on the forest floor. A total of 45 myxomycete species were recorded from 3021 collected samples that occurred at the July peak during the warmest and humidest season from April to November. The occurrence of species was significantly related to the changes in mean temperature and minimum temperature on both leaf types under humid conditions. Myxomycete assemblages were divided into three seasonal phases. Most of the species occurred in June–September, while a few species demonstrated characteristic distributions; i.e., Didymium melanospermum appeared in April– May and Diderma umbilicatum appeared in October– November. The respective leaf types supported the reproduction of myxomycetes with high species richness and diversity, with 34 species and H’ = 2.59 on deciduous trees and similarly 30 species and H’ = 2.49 on evergreen trees. Several species, however, exhibited a preference for either the deciduous tree or evergreen tree leaves. Thus, a mixed forest that defoliates during different two seasons yields a greater species diversity of myxomycete assemblage.     

Distinct Leaf‐trait Syndromes of Evergreen and Deciduous Trees in a Seasonally Dry Tropical Forest

In seasonally dry tropical forests, tree species can be deciduous, remaining without leaves throughout the dry season, or evergreen, retaining their leaves throughout the dry season. Deciduous and evergreen trees specialize in habitats that differ in water availability (hillside and riparian forest, respectively) and in their exposure to herbivore attack (seasonal and continuous, respectively). We asked whether syndromes of leaf traits in deciduous and evergreen trees were consistent with hypothesized abiotic and biotic selective pressures in their respective habitat. We measured seven leaf traits in 19 deciduous and 11 evergreen tree species in a dry tropical forest in Western Mexico, and measured rates of herbivory on 23 of these species. We investigated the covariance of leaf traits in syndromes related to phenology and associated physiology, and to anti‐herbivory defense. We found evidence for syndromes that separated phenological strategies among four traits: toughness, water content, specific leaf area, and carbon:nitrogen (C:N) ratios. We found a trade‐off between two other traits: trichomes and latex. Overall, evergreen species exhibited lower rates of herbivory than deciduous species. Lower rates of herbivory were explained by a syndrome of higher toughness, lower water content, and higher C:N ratios, which are traits representative of evergreen trees. Phenology and trait syndromes did not exhibit significant phylogenetic signal, consistent with the hypothesis of evolutionary convergence among phenologies and associated leaf‐trait syndromes. Our results suggest that deciduous and evergreen trees could respond to differential water availability and herbivory in their respective habitats by converging on distinct leaf‐trait syndromes. Abstract in Spanish is available at http://www.blackwell‐synergy.com/loi/btp .     

Plant diversity patterns in semi-natural grasslands along an elevational gradient in southern Norway

Leaf demography and growth of six common, co-occurring woody plant species that varied in stature (tree vs. shrub) and leaf texture (sclerophyllous, coriaceous, malacophyllous) were examined in a subtropical savanna parkland in southern Texas, USA. We tested the hypotheses that, (a) leaves of plants with evergreen canopies would have longer life spans than those of deciduous species; (b) supplementation of soil moisture would decrease leaf life span in both evergreen and deciduous species; (c) species responses to increased soil moisture availability would be inversely related to leaf longevity; and (d) deciduous growth forms would exhibit a greater growth response to increased soil moisture availability than their evergreen counterparts.A variety of seasonal leaf habits (evergreen, winter-deciduous and summer-deciduous canopies) and leaf life spans (median = 66 to 283 days) were represented by the targeted species, but there was no clear relationship between seasonal leaf habit and leaf longevity. Among species with evergreen canopies, median leaf longevity ranged from short (Zanthoxylum fagara = 116 days; Condalia hookeri = 158 days) to long (Berberis trifoliolata = 283 days) but did not exceed 1 yr. In fact, leaf longevity in evergreen shrubs was often comparable to, or shorter than, that of species with deciduous canopies (Ziziphus obtusifolia = 66 days; Diospyros texana = 119 days; Prosopis glandulosa = 207 days). Augmentation of surface soil moisture had no detectable effect on median leaf life span in any species and there was no clear relationship between leaf longevity and species growth responses to irrigation. Contrary to expectations, species with evergreen canopies responded to irrigation by producing more leaf biomass, longer shoots and more leaf cohorts/year than did deciduous species.Species differences in the annual cycle of leaf initiation, leaf longevity and canopy development, combined with contrasts in root distributions and a highly variable climate, may allow for spatial and temporal partitioning of resources and hence, woody species coexistence and diversity in this system. However, the lack of expected relationships between leaf longevity, leaf habit and plant responses to resource enhancement suggests that structure-function relationships and functional groupings developed in strongly seasonal environments cannot be applied with confidence to these subtropical savannas and thorn woodlands.     

Short-term effects of simulated environmental change on phenology, leaf traits, and shoot growth of alpine plants on a temperate mountain, northern Japan

In order to assess the responses of circumpolar and semicircumpolar plants growing around their southern distribution margins to artificial warming, we set up 11 open-top chambers (OTCs) on a fell-field (1680 m a.s.l.) in the Taisetsu Mountains, northern Japan. The OTCs increased mean air temperature by 1.3°C through the growing season (June–September) and extended the length of the growing season. We examined phenology and leaf traits of plants in the OTCs and control plots during the first season under artificial warming treatment using two deciduous and three evergreen species. Ledum palustre (evergreen shrub), Vaccinium uliginosum , and Arctous alpinus (deciduous shrubs) showed earlier leaf emergence and/or flowering in the OTCs. Deciduous shrubs had longer individual leaf longevity and an extended foliage period in the OTCs than in the control plots. There were no significant differences in specific leaf area and leaf size for many species between the OTCs and the control plots. Vaccinium vitis-idaea (evergreen shrub), L. palustre, A. alpinus , and Empetrum nigrum (evergreen shrub) had lower leaf nitrogen concentration in the OTCs than in the control plots, whereas it was higher in V. uliginosum . Only E. nigrum showed larger annual shoot growth in the OTCs. No clear differences in response to the warming effect were detected between evergreen and deciduous species in the first season. Circumpolar plants growing in temperate alpine regions may be more affected by season length rather than temperature itself.     

Annual photosynthetic activities of temperate evergreen and deciduous broadleaf tree species with simultaneous and successive leaf emergence in response to altitudinal air temperature

To explain why the composition of evergreen and deciduous forests changes along air temperature gradients, we measured several traits of single leaves from temperate deciduous and evergreen broadleaf trees with simultaneous and successive leaf emergence growing at different altitudes in the field. The parameters included seasonal net photosynthetic rate, longevity, mass per area, nitrogen content, and photosynthetic nitrogen-use efficiency. With decreasing altitude, the leaf longevity of deciduous broadleaf trees increased, whereas the maximum net photosynthetic rate decreased. In contrast, leaf longevity of evergreen broadleaf trees decreased, whereas the minimum net photosynthetic rate in winter increased. Along the air temperature gradient, the annual production of deciduous trees with simultaneous leaf emergence may be constant, because the integrated lifetime net photosynthetic rate (ILNPR) of a single leaf changed little. In comparison, deciduous trees with successive leaf emergence may show enhanced annual production with increasing air temperature, by increasing the total leaf number per branch and tree under an extended growing season. Temperate evergreen broadleaf tree species may also show increased annual production with increasing air temperature by sufficiently raising the number of the first-year leaves to the total leaves of branch and tree, which is accelerated by raising the integrated first-year net photosynthetic rate of the single leaf, despite little change in the ILNPR. With increasing air temperature from cool-temperate to warm-temperate zones, evergreen broadleaf tree species gain an advantage of the annual production over deciduous broadleaf tree species with simultaneous leaf emergence.     

Responses of alpine shrubs to simulated environmental change during three years in the mid-latitude mountain, northern Japan

Effects of artificial warming on phenology, individual leaf traits, vegetative growth, and reproduction of five alpine species (two deciduous and three evergreen shrubs) were investigated during three years in the mid-latitude alpine, northern Japan. Eleven open-top chambers (OTCs) were set up on a fellfield (1680 m a. s. l.) in the Taisetsu Mountains by which air temperature at plant height was increased by ca 2°C. Vaccinium uliginosum (deciduous shrub) showed earlier leaf emergence in every season and earlier flowering only in the first season in the OTCs. By contrast, acceleration of leaf emergence in the OTCs was not clear for other species, i.e. Arctous alpinus (deciduous shrub). Ledum palustre. V. vitis-idaea , and Empetrum nigrum (evergreen shrub). Both deciduous species showed longer leaf life-span in the OTCs every season. All evergreen species had higher leaf survival rates in the OTCs. indicating extension of leaf life-span. Leaf nitrogen concentration and leaf mass per unit leaf area (mg cm ⁻²) generally tended to decrease in the OTCs. Relationships between the individual leaf traits and cumulative air temperature during the leaf developing period were not clear. Total leaf production during the three seasons increased in the OTCs in A. alpinus. L. palustre. V. vitis-idaea , and E. nigrum. All evergreen shrubs showed larger shoot growth in the OTCs but both deciduous shrubs did not show significant changes. In contrast to the vegetative growth, deciduous shrubs produced more flowers in the OTCs. Fruit production was not influenced by the OTCs for all species. The extension of photosynthetic period in the OTCs may contribute to the larger vegetative growth or flower production.     

重庆石灰岩地区主要木本植物叶片性状及养分再吸收特征

以重庆石灰岩地区15种常绿木本植物和14种落叶木本植物为研究对象,对两种生活型植物叶片衰老前后叶干物质含量(LDMC)、比叶面积(SLA)和叶片厚度(LT)进行了比较,并采用不同的计算方法(单位质量叶片养分含量、单位面积叶片养分含量)分析了两类植物叶片衰老前后养分含量及再吸收特征,最后对养分再吸收效率与其他叶性状因子之间的关系进行了相关分析。结果表明:常绿植物成熟叶LDMC、LT及衰老叶LT显著低于落叶植物,落叶植物成熟叶和衰老叶SLA均显著高于常绿植物(P0.05);基于单位质量叶片计算的养分含量,常绿植物成熟和衰老叶N、P量均低于落叶植物,而基于单位面积叶片计算的N、P含量则表现出相反的趋势;基于不同方法计算的N、P再吸收效率差异不明显,其中常绿植物基于单位质量叶片养分含量计算的N、P平均再吸收效率为39.42%、43.79%,落叶植物的为24.08%、33.59%;常绿和落叶植物N、P再吸收效率与LDMC、SLA、LT和成熟叶N、P含量之间没有显著相关性,但与衰老叶养分含量存在显著负相关(P0.05)。研究发现,无论是常绿植物还是落叶植物,衰老叶N、P含量均较低,表明石灰岩地区植物具有较高的养分再吸收程度。     

Variation in evergreen and deciduous species leaf phenology in Assam, India

In the present study phenological activities such as leaf and shoot growth, leaf pool size and leaf fall were observed for 3 years (March 2007–March 2010) in 19 tree species (13 evergreen and 6 deciduous species) in a wet tropical forest in Assam, India. The study area receives total annual average rainfall of 2,318 mm of which most rain fall (>70 %) occurs during June–September. Both the plant groups varied significantly on most of the shoot and leaf phenology parameters. In general, growth in deciduous species initiated before the evergreen species and showed a rapid shoot growth, leaf recruitment and leaf expansion compared to evergreen species. Leaf recruitment period was significantly different between evergreen (4.2 months) and deciduous species (6.8 months). Shoot elongation rate was also significantly different for evergreen and deciduous species (0.09 vs. 0.14 cm day^?1 shoot^?1). Leaf number per shoot was greater for deciduous species than for evergreen species (34 vs. 16 leaves). The average leaf life span of evergreen species (328 ± 32 days) was significantly greater than that of deciduous species (205 ± 16 days). The leaf fall in deciduous species was concentrated during the winter season (Nov–Feb), whereas evergreens retained their leaves until the next growing season. Although the climate of the study area supports evergreen forests, the strategies of the deciduous species such as faster leaf recruitment rate, longer leaf recruitment time, faster shoot elongation rate during favorable growing season and short leaf life span perhaps allows them to coexist with evergreen species that have the liberty to photosynthesize round the year. Variations in phenological strategies perhaps help to reduce the competition among evergreen and deciduous species for resources in these forests and enable the coexistence of both the groups.     

广西猫儿山不同海拔常绿和落叶树种的营养再吸收模式

土壤养分供给性大小是否影响植物氮和磷再吸收效率仍存在争议。调查了广西猫儿山不同海拔常绿和落叶树种成熟和衰老叶片的氮和磷含量,探讨营养再吸收是否受到叶片习性和海拔的影响。所有树种氮和磷再吸收效率的平均值分别为56.5%和52.1%。常绿树种比落叶树种有显著较高的氮再吸收效率(P0.001)和磷再吸收效率(P0.01),这与前者有较低的衰老叶片氮和磷含量密切相关。随着海拔的上升,氮再吸收效率显著下降(P0.01),磷再吸收效率显著提高(P0.05)。氮再吸收效率与土壤氮:磷比(r=-0.41,P0.05)和成熟叶片氮:磷比(r=-0.37,P0.05)负相关,磷再吸收效率与土壤氮:磷比(r=0.44,P0.05)和成熟叶片氮:磷比(r=0.47,P0.01)正相关,表明了树种对低海拔氮限制的适应逐渐转变为对高海拔磷限制的适应。此外,氮再吸收效率与年均温正相关(r=0.43,P0.05)而磷再吸收效率与年均温负相关(r=-0.45,P0.01),这表明气温也是调节树木营养再吸收格局的重要影响因素。不同海拔树种氮和磷再吸收模式的差异可能是引起广西猫儿山常绿树种沿海拔形成双峰分布的原因之一。     

4种阔叶树种叶中氮和磷的季节动态及其转移

从叶完全展开到生长季结束 ,对常绿阔叶树种日本米槠 (Castanopsis cuspidata(Thunb.) Schottky)和具柄冬青 (Ilexpedunculosa Miq)及落叶阔叶树种栎 (Quercus serrata Murr.)和栓皮栎 (Quercus variabilis Blume)叶片的 N和 P浓度、含量和养分转移进行了测定 .在生长期中日本米槠新叶的 N浓度在 5月为 36 .6 g/ kg,然后降到 15 .5和 17.5 g/ kg之间 ,其老叶的N浓度波动于 10 .4和 13.1g/ kg的范围内 ,而具柄冬青新叶的 N浓度从 2 7.3下降到 16 .0 g/ kg,接着上升到 18.3g/ kg,其老叶的 N浓度在 12 .0到 15 .5 g/ kg的范围内。栎和栓皮栎的叶 N浓度分别从 2 8.8下降到 18.1g/ kg和从 2 8.5下降到17.5 g/ kg。日本米槠新叶的 N含量从 1.5 4下降到 1.35 g/ m2 ,接着上升到 1.5 0 g/ m2 ,其老叶 N含量从 1.36下降到1.0 0 g/ m2 ,接着上升到 1.2 1g/ m2 ,而具柄冬青新叶的 N含量从 2 .2 5下降到 1.6 0 g/ m2 ,接着上升到 2 .2 0 g/ m2 ,其老叶的 N含量从 2 .13下降到 1.6 5 g/ m2。栎和栓皮栎的叶 N含量分别从 2 .10下降到 1.2 8g/ m2和从 2 .95下降到 2 .13g/ m2。日本米槠新叶的 P浓度由 3.39g/ kg降到 1.12和 1.15 g/ kg之间 ,其老叶的 P浓度变化于 0 .6 6和 0 .88g/ kg的范围内 ,而具柄冬青新叶的     

Leaf traits and water relations of 12 evergreen species in Costa Rican wet and dry forests: patterns of intra-specific variation across forests and seasons

This study examined variation in leaf traits and water relations in 12 evergreen and semideciduous woody species that occur in both seasonal wet and dry forests in Costa Rica and compared intra-specific leaf–trait correlations to those found in inter-specific global studies. The following traits were measured in both forests across seasons for 2 years: leaf nitrogen (N), leaf carbon (C), specific leaf area (SLA), toughness, cuticle thickness, leaf thickness, and leaf lifespan (LLS). Leaf water potential (LWP) and water content (LWC) were measured as indices of plant available water. Canopy openness, soil moisture, and herbivory were also measured to compare environmental variation across sites. Although species contributed the greatest amount to variation in traits, season, forest, and their interaction had a large influence on patterns of intra-specific leaf–trait variation. Leaf traits that contributed most to variation across sites were C, LWP, leaf thickness, and SLA. Traits that contributed most to variation across seasons were leaf toughness, LWP, and LWC. Furthermore, leaf traits were more correlated (i.e., number and strength of correlations) in the dry than in the wet forest. In contrast to results from global literature syntheses, there was no correlation between LLS and N, or LLS and SLA. Both light and water availability vary seasonally and may be causing variation in a number of leaf traits, specifically those that relate to water relations and leaf economics. Strong seasonality may cause leaf–trait relationships at the local scale to differ from those documented in continental and global-scale studies.     

Greater deciduous shrub abundance extends tundra peak season and increases modeled net CO2 uptake

Satellite studies of the terrestrial Arctic report increased summer greening and longer overall growing and peak seasons since the 1980s, which increases productivity and the period of carbon uptake. These trends are attributed to increasing air temperatures and reduced snow cover duration in spring and fall. Concurrently, deciduous shrubs are becoming increasingly abundant in tundra landscapes, which may also impact canopy phenology and productivity. Our aim was to determine the influence of greater deciduous shrub abundance on tundra canopy phenology and subsequent impacts on net ecosystem carbon exchange (NEE) during the growing and peak seasons in the arctic foothills region of Alaska. We compared deciduous shrub‐dominated and evergreen/graminoid‐dominated community‐level canopy phenology throughout the growing season using the normalized difference vegetation index (NDVI). We used a tundra plant‐community‐specific leaf area index (LAI) model to estimate LAI throughout the green season and a tundra‐specific NEE model to estimate the impact of greater deciduous shrub abundance and associated shifts in both leaf area and canopy phenology on tundra carbon flux. We found that deciduous shrub canopies reached the onset of peak greenness 13 days earlier and the onset of senescence 3 days earlier compared to evergreen/graminoid canopies, resulting in a 10‐day extension of the peak season. The combined effect of the longer peak season and greater leaf area of deciduous shrub canopies almost tripled the modeled net carbon uptake of deciduous shrub communities compared to evergreen/graminoid communities, while the longer peak season alone resulted in 84% greater carbon uptake in deciduous shrub communities. These results suggest that greater deciduous shrub abundance increases carbon uptake not only due to greater leaf area, but also due to an extension of the period of peak greenness, which extends the period of maximum carbon uptake.     

The effect of increased nutrient availability on leaf turnover and aboveground productivity of two evergreen ericaceous shrubs

Summary Leaf turnover and aboveground productivity in relation to nutrient availability were studied in the evergreen shrubs Erica tetralix and Calluna vulgaris. In monospecific stands of these species four levels of nutrient (NPK) availability were created during three growing seasons. Percentage survival and life expectancy of Erica leaves decreased with increasing nutrient availability. For Calluna there was no effect. Winter mortality of Erica leaves was smaller than growing season mortality. These was no difference for Calluna. The timing of leaf mortality of both species was not affected by nutrient treatment. At the end of the experimental period current year leaf biomass, total biomass and current year second year and third year biomass of both species showed a significant increase with increasing nutrient availability. The relative increase was greater for Calluna, except for second and third year biomass. Stem production and stem mortality of both species increased with increasing nutrient availability. The increased stem mortality resulted also for Calluna in an increased leaf turnover (per unit ground area) with increasing nutrient availability. Nutrient cycling in ecosystems dominated by these species will increase with increasing nutrient availability, because of increased leaf and stem turnover and productivity. This phenotypic effect is similar to the effect of the shift in dominance between different species which occurs along natural gradients of nutrient availability.     

Evidence for facultative deciduousness in Colophospermum mopane in semi‐arid African savannas

Leaf phenology dictates the time available for carbon assimilation, transpiration and nutrient uptake in plants. Understanding the environmental cues that control phenology is therefore vital for predicting climate‐related changes to plant and ecosystem function. In contrast to temperate systems, and to a lesser degree, tropical forest systems, the cues initiating leaf drop in tropical savannas are poorly studied. We investigated the cues for leaf fall in a tropical monodominant arid savanna species, Colophospermum mopane, using an irrigation experiment. We tracked soil moisture, solar radiation, air temperature, leaf water status, leaf health and leaf carbon balance through the dry season in both irrigated and control plants. Water was the primary cue driving leaf loss of C. mopane rather than temperature or light. Trees watered throughout the dry season retained their canopies. These leaves remained functional and continued photosynthesis throughout the dry season. Leaf carbon acquisition rates did not decline with leaf age but were affected by soil moisture availability and temperature. Leaf loss did not occur when leaf carbon gain was zero, or when a particular leaf carbon threshold was reached. Colophospermum mopane is facultatively deciduous as water availability determines leaf drop in this widespread arid savanna species. Obligate deciduosity is not the only successful strategy in climates with a long dry season.