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.     

Inter-species variation of photosynthetic and xylem hydraulic traits in the deciduous and evergreen Euphorbiaceae tree species from a seasonally tropical forest in south-western China

The objective of the present study was to examine the functional coordination among hydraulic traits, xylem characteristics and gas exchange rates across three deciduous Euphorbiaceae tree species (Hevea brasiliensis, Macaranga denticulata and Bischofia javanica) and three evergreen Euphorbiaceae tree species (Drypetes indica, Aleurites moluccana and Codiaeum variegatum) from a seasonally tropical forest in south-western China. The deciduous tree species were more vulnerable to water stress-induced embolism than the evergreen tree species. However, the deciduous tree species generally had higher maximal rates of sapwood and leaf-specific hydraulic conductivity (K _S and K _L), respectively. Compared with the evergreen tree species, the deciduous tree species, however, possessed a lower density of sapwood and a wider diameter of xylem vessels. Regardless of leaf phenology, the hydraulic vulnerability and conductivity were significantly correlated with sapwood density and mean vessel diameter. Furthermore, the hydraulic vulnerability was positively correlated with water transport efficiency. In addition, the deciduous tree species exhibited higher maximal photosynthetic rates (A _max) and stomatal conductance (g _max), but lower water use efficiency (WUE). Interestingly, the A _max, g _max and WUE were strongly correlated with K _S and K _L across the deciduous and evergreen tree species. These results suggest that xylem structure, rather than leaf phenology, accounts for the difference in hydraulic traits between the deciduous tree species and the evergreen tree species. Meanwhile, our results show that there is a significant trade-off between hydraulic efficiency and safety, and a strong functional correlation between the hydraulic capacity and gas exchange rates across the deciduous and evergreen tree species.     

Leaf morphology of 40 evergreen and deciduous broadleaved subtropical tree species and relationships to functional ecophysiological traits

We explored potential of morphological and anatomical leaf traits for predicting ecophysiological key functions in subtropical trees. We asked whether the ecophysiological parameters stomatal conductance and xylem cavitation vulnerability could be predicted from microscopy leaf traits. We investigated 21 deciduous and 19 evergreen subtropical tree species, using individuals of the same age and from the same environment in the Biodiversity‐Ecosystem Functioning experiment at Jiangxi (BEF‐China). Information‐theoretic linear model selection was used to identify the best combination of morphological and anatomical predictors for ecophysiological functions. Leaf anatomy and morphology strongly depended on leaf habit. Evergreen species tended to have thicker leaves, thicker spongy and palisade mesophyll, more palisade mesophyll layers and a thicker subepidermis. Over 50% of all evergreen species had leaves with multi‐layered palisade parenchyma, while only one deciduous species (Koelreuteria bipinnata) had this. Interactions with leaf habit were also included in best multi‐predictor models for stomatal conductance (g_s) and xylem cavitation vulnerability. In addition, maximum g_s was positively related to log ratio of palisade to spongy mesophyll thickness. Vapour pressure deficit (vpd) for maximum g_s increased with the log ratio of palisade to spongy mesophyll thickness in species having leaves with papillae. In contrast, maximum specific hydraulic conductivity and xylem pressure at which 50% loss of maximum specific xylem hydraulic conductivity occurred (Ψ₅₀) were best predicted by leaf habit and density of spongy parenchyma. Evergreen species had lower Ψ₅₀ values and lower maximum xylem hydraulic conductivities. As hydraulic leaf and wood characteristics were reflected in structural leaf traits, there is high potential for identifying further linkages between morphological and anatomical leaf traits and ecophysiological responses.     

Summer and winter sensitivity of leaves and xylem to minimum freezing temperatures: a comparison of co-occurring Mediterranean oaks that differ in leaf lifespan

Freezing sensitivity of leaves and xylem was examined in four co-occurring Mediterranean oaks (Quercus spp.) grown in a common garden to determine whether freezing responses of leaves and xylem were coordinated and could be predicted by leaf lifespan. Freezing-induced embolism and loss of photosynthetic function were measured after overnight exposure to a range of subzero temperatures in both summer and winter. Both measures were found to be dependent on minimum freezing temperature and were correlated with leaf lifespan and vessel diameter. The dependence of xylem embolism on minimum freezing temperature may result from the decline in water potential with ice temperature that influences the redistribution of water during freezing and leads to an increase in xylem tension. Winter acclimatization had a relatively small effect on the vulnerability to freezing-induced embolism, although leaf photosynthetic function showed a strong acclimatization response, particularly in the two evergreen species. Quercus ilex, the species with the longest leaf lifespan and narrowest vessel diameters, showed the highest freezing tolerance. This helps explain its ability to inhabit a broad range throughout the Mediterranean region. By contrast, the inability of the deciduous oaks to maintain photosynthetic and vascular function throughout the winter indicates a competitive disadvantage that may prevent them from expanding their ranges.     

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.     

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.     

Stem xylem features in three Quercus (Fagaceae) species along a climatic gradient in NE Spain

 Stem xylem features in two evergreen Quercus species (Q. coccifera and Q. ilex) and a deciduous one (Q. faginea) were analysed along an Atlantic-Mediterranean climatic gradient in which rainfall and winter cold experience strong variation. Mean maximum vessel diameter, vessel density, vessel element length, xylem transverse sectional area, Huber value (xylem transverse sectional area per leaf area unit), theoretical leaf specific conductivity (estimated hydraulic conductance per leaf area unit) and total leaf area were determined in 3-year-old branches. Q. faginea presented the widest vessels and the highest theoretical leaf specific conductivity while Q. coccifera showed the lowest total leaf area and the highest Huber value. Studied features did not exhibit significant correlations with mean minimum January temperature in any species but did show significant relationships with rainfall. In Q. coccifera, mean maximum vessel diameter, vessel element length and theoretical leaf specific conductivity increased with higher rainfall while vessel density decreased. Mean maximum vessel diameter and total leaf area in Q. ilex increased with precipitation whereas variables of Q. faginea did not show any significant trend. Results suggest that aridity, rather than minimum winter temperature, controls stem xylem responses in the studied evergreen species. Q. faginea traits did not show any response to precipitation, probably because this species develops deep roots, which in turn makes edaphic and topographic factors more important in the control of soil water availability. In response to aridity Q. coccifera only exhibits adjustment at a xylem level by reducing its water transport capacity through a reduction of vessel diameter without changing the amount of xylem tissue or foliage, whereas Q. ilex adjusts its water transport capacity in parallel to the foliage area. Received: 13 January 1997 / Accepted: 8 April 1997     

Aboveground nitrogen and phosphorus use by five plantation-grown trees with different leaf longevities

Aboveground nitrogen (N) and phosphorus (P) requirement, retranslocation and use efficiency were determined for 28-year-old red oak (Quercus rubra L.), European larch (Larix decidua Miller), white pine (Pinus strobes L.), red pine (Pinus resinosa Ait.) and Norway spruce (Picea abies (L) Karst.) plantations on a similar soil in southwestern Wisconsin. Annual aboveground N and P requirements (kg/ha/yr) totaled 126 and 13 for red oak, 86 and 9 for European larch, 80 and 9 for white pine, 38 and 6 for red pine, and 81 and 13 for Norway spruce, respectively. Nitrogen and P retranslocation from current foliage ranged from 81 and 72%, respectively, for European larch, whereas red pine retranslocated the smallest amount of N (13%) and Norway spruce retranslocated the smallest amount of P (18%). In three evergreen species, uptake accounted for 72 to 74% of annual N requirement whereas for two deciduous species retranslocation accounted for 76 to 77% of the annual N requirement. Nitrogen and P use (ANPP/uptake) was more efficient in deciduous species than evergreen species. The results from this common garden experiment demonstrate that differences in N and P cycling among species may result from intrinsic characteristics (e.g. leaf longevity) rather than environmental conditions.     

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。     

Aspects of tissue water relations and seasonal changes of leaf water potential components of evergreen and deciduous species coexisting in tropical dry forests

Summary This study compared the tissue water relations and seasonal changes in leaf water potential components of an evergreen tree,Morisonia americana, and two evergreen shrubs,Capparis verrucosa andC. aristiquetae, with two deciduous trees,Humboltiella arborea andLonchocarpus dipteroneurus, and the deciduous vineMansoa verrucifera. All these species coexist in a tropical dry forest in Venezuela. Leaves of the evergreen species are sclerophyllous, while those of the deciduous species are mesophytic. Leaf area to leaf weight ratios of fully mature leaves were about 75 and 170 cm² g^–1 in evergreen and deciduous species, respectively. Seasonal fluctuations of leaf water content per unit of dry weight, water potential, and turgor pressure were smaller in evergreen than in deciduous species. The analysis of tissue water relations using pressurevolume curves showed that evergreen species could develop a higher leaf turgor and lose turgor at lower leaf water potentials than deciduous species. This was related to a lower osmotic potential at full turgor in evergreen (-3.0 MPa)_than in deciduous (-2.0 MPa) species, rather than to the elastic properties of leaf tissue. The volumetric modulus of elasticity was 14 MPa in evergreen compared with 7–10 MPa in deciduous species. Thus, leaf characteristics are important in determining the drought resistance of evergreen species of this tropical dry forest.     

Hydraulic architecture of deciduous and evergreen dry rainforest tree species from north-eastern Australia

Hydraulic conductivity and xylem anatomy were examined in stems of two evergreen species, Alphitonia excelsa (Fenzal) Benth. and Austromyrtus bidwillii (Benth.) Burret., and two drought-deciduous species, Brachychiton australis (Schott and Endl.) A. Terracc. and Cochlospermum gillivraei Benth., from a seasonally dry rainforest in north Queensland, Australia. The deciduous species possessed hydraulic architecture typical of drought-sensitive plants, i.e. low wood density, wider xylem vessels, higher maximal rates of sapwood specific hydraulic conductivity (K_s) and high vulnerability to drought-induced embolism. In contrast, the evergreen species had lower rates of K_h and leaf specific conductivity (K_L) but were less susceptible to embolism. The evergreen species experienced leaf water potentials <–4.0 MPa during the dry season, while the deciduous species shed their leaves before leaf water potentials declined below –2.0 MPa. Thus, the hydraulic architecture of the evergreens allows them to withstand the greater xylem pressure gradients required to maintain water transport to the canopy during the dry season. Our results are consistent with observations made in neotropical dry forests and demonstrate that drought-deciduous species with low wood density and high water storage capacity are likely to be more hydraulically efficient, but more vulnerable to embolism, than coexisting evergreens.     

亚热带常绿落叶阔叶混交林植物功能性状的种间和种内变异

不同物种间的功能性状差异是自然生态系统中物种共存的基础, 而物种内个体间的性状变异对物种的共存和分布同样具有重要作用。本文以湖北星斗山自然保护区亚热带常绿落叶阔叶混交林内28种主要树种(通过物种多度排序获得, 其中常绿和落叶树种各14种)为研究对象, 探讨不同叶习性树种的4种功能性状(比叶面积、叶干物质含量、叶面积和比茎密度)在种间和种内的差异程度。结果表明: (1)常绿和落叶树种在4种功能性状上均存在显著差异, 常绿树种的比叶面积和叶面积显著低于落叶树种, 但叶干物质含量和比茎密度则显著高于落叶树种; (2)比叶面积的变化主要来源于叶习性(57.49%), 叶面积变化主要来源于种间(66.80%)和种内变异(27.52%), 叶干物质含量的变化主要来源于种间(38.12%)和种内(33.88%)变异, 但比茎密度的变化主要来源于种内变异(51.50%), 其次为种间变异(32.52%); (3)常绿和落叶树种种间水平的性状相关性可能掩盖各功能性状之间的相关性。种内变异能够显著影响群落间的植物功能性状差异, 但不同功能性状的种内变异程度存在差异。     

A Cost-Benefit Analysis of Leaves of Eight Australian Savanna Tree Species of Differing Leaf Life-Span

Cost-benefit analysis of foliar construction and maintenance costs and of carbon assimilation of leaves of differing life-span were conducted using two evergreen, three semi-deciduous, and three deciduous tree species of savannas of north Australia. Rates of radiant-energy-saturated CO₂ assimilation (P _max) and dark respiration were measured and leaves were analysed for total nitrogen, fat, and ash concentrations, and for heat of combustion. Specific leaf area, and leaf N and ash contents were significantly lower in longer-lived leaves (evergreen) than shorter-lived leaves (deciduous) species. Leaves of evergreen species also had significantly higher heat of combustion and lower crude fat content than leaves of deciduous species. On a leaf area basis, P _max was highest in leaves of evergreen species, but on a leaf dry mass basis it was highest in leaves of deciduous species. P _max and total Kieldahl N content were linearly correlated across all eight species, and foliar N content was higher in leaves of deciduous than evergreen species. Leaf construction cost was significantly higher and maintenance costs were lower for leaves of evergreen than deciduous species. Maintenance and construction costs were linearly related to each other across all species. Leaves of evergreen species had a higher cost-benefit ratio compared to leaves of deciduous species but with longer lived leaves, the payback interval was longer in evergreen than deciduous species. These results support the hypotheses that longer lived leaves are more expensive to construct than short-lived leaves, and that a higher investment of N into short-lived leaves occurs which supports a higher P _max over a shorter payback interval. This revised version was published online in June 2006 with corrections to the Cover Date.     

Litter decomposition of woody species in shrublands of NW Patagonia: how much do functional groups and microsite conditions influence decomposition?

The study examined the effects of leaf traits, soil microsite, and microclimate characteristics on litter decomposition of the dominant species in two functional groups (FG), deciduous and evergreen, in shrublands in NW Patagonia, Argentina. Leaf traits considered were nutrient concentration (C, N, P, C/N, and N/P) and physical characteristics (area, strength, specific leaf area, and dry matter content). Soil microsite characteristics measured were pH, C, N, P, C/N and water retention capacity, while soil microclimate characteristics recorded were soil and air, temperature and moisture, and solar radiation. Five evergreen and five deciduous woody shrub species were selected. During 1 year, litter and microsite properties were measured below canopy: (i) senescent leaf chemical and physical properties, and the quantity as well as field decomposition of litter and (ii) soil chemistry, and soil and air physical properties. The factors controlling litter decomposition were different for each FG. In deciduous species, C/N ratio had a negative effect on decomposition. In evergreen species, decomposition was affected negatively by leaf carbon and dry matter content. Litter decomposition depended exclusively on the inherent senescent leaves traits. The common decomposition pattern between species of both FG could be attributed to similar leaf traits and the correlation between variables that control decomposition in both groups. Plant nutrient inputs associated with the litter decomposition process did not explain the soil nutrient content. These results suggest that other organic matter sources (roots, branches, and fruits) are more important than leaves on soil fertility.     

Seedling growth strategies and seed size effects in fourteen oak species native to different soil moisture habitats

 Seedling growth and morphology are thought to reflect evolutionary responses to habitat or influences of seed size. To test these hypotheses, we selected fourteen species of North American oaks differing in soil moisture habitat preference and seed size. Seedlings were grown for 1 – 2 years with abundant soil water and moderate soil nutrition in pots placed outdoors and in a common garden. Oak species native to xeric environments produced the smallest seedlings. Oaks from hydric soils had more shoot weight per unit of root weight and more height per unit of total plant weight than did mesic or xeric oaks. Essentially no differences in leaf area per unit of total plant weight were detected. Species with thinner and larger individual leaves tended to produce larger seedlings. Within species, seed size was generally unrelated to seedling growth, although results may have been complicated by uncontrolled genotypic variability. However, when species were compared, those with larger mean seed size produced larger seedlings. Root/shoot allometry, height growth and leaf thickness in the tested species may reflect evolutionary responses to soil moisture and flooding. Although seed size influenced seedling growth, no clear relationship between seed size and soil moisture habitat was found. Received: 26 March 1995 / Accepted: 30 November 1995     

Similar variation in carbon storage between deciduous and evergreen treeline species across elevational gradients

Background and Aims

The most plausible explanation for treeline formation so far is provided by the growth limitation hypothesis (GLH), which proposes that carbon sinks are more restricted by low temperatures than by carbon sources. Evidence supporting the GLH has been strong in evergreen, but less and weaker in deciduous treeline species. Here a test is made of the GLH in deciduous–evergreen mixed species forests across elevational gradients, with the hypothesis that deciduous treeline species show a different carbon storage trend from that shown by evergreen species across elevations.

Methods

Tree growth and concentrations of non-structural carbohydrates (NSCs) in foliage, branch sapwood and stem sapwood tissues were measured at four elevations in six deciduous–evergreen treeline ecotones (including treeline) in the southern Andes of Chile (40°S, Nothofagus pumilio and Nothofagus betuloides; 46°S, Nothofagus pumilio and Pinus sylvestris) and in the Swiss Alps (46°N, Larix decidua and Pinus cembra).

Key Results

Tree growth (basal area increment) decreased with elevation for all species. Regardless of foliar habit, NSCs did not deplete across elevations, indicating no shortage of carbon storage in any of the investigated tissues. Rather, NSCs increased significantly with elevation in leaves (P < 0·001) and branch sapwood (P = 0·012) tissues. Deciduous species showed significantly higher NSCs than evergreens for all tissues; on average, the former had 11 % (leaves), 158 % (branch) and 103 % (sapwood) significantly (P < 0·001) higher NSCs than the latter. Finally, deciduous species had higher NSC (particularly starch) increases with elevation than evergreens for stem sapwood, but the opposite was true for leaves and branch sapwood.

Conclusions

Considering the observed decrease in tree growth and increase in NSCs with elevation, it is concluded that both deciduous and evergreen treeline species are sink limited when faced with decreasing temperatures. Despite the overall higher requirements of deciduous tree species for carbon storage, no indication was found of carbon limitation in deciduous species in the alpine treeline ecotone.     

Relative growth rate variation of evergreen and deciduous savanna tree species is driven by different traits

Background and Aims

Plant relative growth rate (RGR) depends on biomass allocation to leaves (leaf mass fraction, LMF), efficient construction of leaf surface area (specific leaf area, SLA) and biomass growth per unit leaf area (net assimilation rate, NAR). Functional groups of species may differ in any of these traits, potentially resulting in (1) differences in mean RGR of groups, and (2) differences in the traits driving RGR variation within each group. We tested these predictions by comparing deciduous and evergreen savanna trees.

Methods

RGR, changes to biomass allocation and leaf morphology, and root non-structural carbohydrate reserves were evaluated for juveniles of 51 savanna species (34 deciduous, 17 evergreen) grown in a common garden experiment. It was anticipated that drivers of RGR would differ between leaf habit groups because deciduous species have to allocate carbohydrates to storage in roots to be able to flush leaves again, which directly compromises their LMF, whereas evergreen species are not subject to this constraint.

Key Results

Evergreen species had greater LMF and RGR than deciduous species. Among deciduous species LMF explained 27 % of RGR variation (SLA 34 % and NAR 29 %), whereas among evergreen species LMF explained between 2 and 17 % of RGR variation (SLA 32–35 % and NAR 38–62 %). RGR and LMF were (negatively) related to carbohydrate storage only among deciduous species.

Conclusions

Trade-offs between investment in carbohydrate reserves and growth occurred only among deciduous species, leading to differences in relative contribution made by the underlying components of RGR between the leaf habit groups. The results suggest that differences in drivers of RGR occur among savanna species because these have different selected strategies for coping with fire disturbance in savannas. It is expected that variation in the drivers of RGR will be found in other functional types that respond differently to particular disturbances.     

Different responses to shade of evergreen and deciduous oak seedlings and the effect of acorn size

An evergreen oak species, Cyclobalanopsis multinervis, and a deciduous oak species, Quercus aliena var. acuteserrata were grown from acorns under two light levels (full sunlight and shade at about 18 % of full sunlight, simulating the light intensities in forest clearings and gaps, respectively) for one growing season. Three hypotheses were tested: (i) the deciduous species grows faster than the evergreen species in forest gaps and clearings; (ii) the deciduous species responds more strongly in terms of growth and morphology to variation in light climate than the evergreen species; and (iii) seedling size is positively correlated to acorn size. The results showed: (i) at both light levels, the deciduous seedlings gained significantly more growth in biomass and height than the evergreen seedlings; (ii) both species produced significantly more biomass in full sunlight than in shade, without showing any significant difference in height between treatments. Increase in light intensity improved the growth of the deciduous seedlings more strongly; (iii) at a similar age, the deciduous seedlings showed a greater response in leaf morphology and biomass allocation to variation in light levels, but when compared at a similar size, biomass allocation patterns did not differ significantly between species; (iv) bigger acorns tended to produce larger seedlings, larger leaf sizes and more leaf area, between and within species. These differences demonstrate that the deciduous species is gap-dependent and has the advantage over the evergreen species in forest gaps and clearings.     

Periodicity of leaf growth and leaf dry mass changes in the evergreen and deciduous species of Southern Assam,India

The study described patterns of leaf dry mass change, leaf mass per area (LMA), relative growth rate and leaf life span (LL) for 14 evergreen and 7 deciduous species of a tropical forest of Southern Assam, India. Leaf expansion in both the groups was, in general, completed before June (i.e. well before the onset of monsoon rains). Although leaf dry mass during leaf initiation phase was significantly higher (P < 0.01) in evergreen species than in deciduous species, at the time of full leaf expansion, average leaf dry mass relative to the peak leaf dry mass, realised by the evergreen species was lower (66 %) than for deciduous species (76 %). Leaf dry mass increase in both groups continued after leaf full expansion. Evergreen species had a longer leaf dry mass steady phase than deciduous species (2–6 vs 2–3 months). Average LMA of mature leaves for evergreen species (77.43 g m^?2) was significantly greater than that of deciduous species (48.43 g m^?2). LL ranged from 165 days in Gmelina arborea (deciduous) to 509 days in Dipterocarpus turbinatus (evergreen). LMA was correlated positively with LL, indicating that evergreen species with higher leaf construction cost retain leaves for longer period to pay back. The average leaf dry mass loss before leaf shedding was greater (P < 0.01) for deciduous species (30.29 %) than for evergreen species (18.31 %). Although the cost of leaf construction in deciduous species was lower than for evergreen species, they replace leaves at a faster rate. Deciduous species perhaps compensate the cost involved in faster leaf replacement through higher reabsorption of dry mass during senescence, which they remobilise to initiate growth in the following spring when soil resources remain limiting.     

Global quantification of contrasting leaf life span strategies for deciduous and evergreen species in response to environmental conditions

Aim Species with deciduous and evergreen leaf habits typically differ in leaf life span (LLS). Yet quantification of the response of LLS, within each habit, to key environmental conditions is surprisingly lacking. The aim of this study is to quantify LLS strategies of the two leaf habits under varying temperature, moisture and nutrient conditions, using a global database. We hypothesize that deciduous LLS reflects the length of the growing season, avoiding unfavourable conditions regardless of the cause. Evergreen species adjust to unfavourable periods and amortize lower net carbon gains over several growing seasons, with increasing LLS associated with increasingly short favourable versus unfavourable season lengths. Location Global. Methods Data on LLS and environmental variables were compiled from global datasets for 189 deciduous and 506 evergreen species across 83 study locations. Individual and combined effects of measures of seasonality of temperature, water and nutrient availability on length of the growing season and on LLS were quantified using linear mixed models. The best models for predicting LLS were obtained using Akaike's information criterion (AIC) and ΔAIC. Results The LLS of deciduous and evergreen species showed opposite responses to changes in environmental conditions. Under unfavourable conditions, deciduous LLS decreases while evergreen LLS increases. A measure of temperature alone was the best predictor of the growing season. The LLS of deciduous species was independent of environmental conditions after expressing LLS in relation to the number of growing seasons. Evergreen species, on the other hand, adjusted to unfavourable conditions by increasing LLS up to four growing seasons. Contrary to expectations, variation in LLS was best explained solely by temperature, instead of by combined measures of temperature, moisture and nutrient availability. Shifts in the photosynthesis to respiration balance might provide a physiological explanation. Main conclusions Temperature, and not drought or nutrient availability, is the primary driver of contrasting responses of LLS for different leaf habit types.