Shoot growth phenology of co-existing evergreen and deciduous species in an oak forest

Shoot growth phenology was compared for the saplings of evergreen and deciduous woody species sharing the same microsite. Growth initiation occurred earlier in evergreens (among co-stratal species) while deciduous species completed their growth earlier. Shoot growth rate was significantly greater (P<0.01) for deciduous trees than evergreen trees. The amount of shoot elongations and shoot diameter was also significantly greater (P<0.01) for deciduous trees than evergreens. On the other hand, among shrubs the amount of shoot elongation and shoot diameter was greater for evergreens but the rate of elongation and diameter was more or less similar for both. The duration of shoot elongation and shoot diameter was significantly longer in evergreens than the deciduous species. Leaf packing (number of leaves per shoot) was significantly more dense in evergreen trees (P<0.01) than in deciduous tree species. Leaf packing was more dense in evergreen than deciduous shrubs but the difference was not significant. Leaf area (per individual leaf) at full expansion was significantly greater (P<0.01) in deciduous species. Leaf dry mass and specific leaf mass in the initial stage was significantly greater for evergreen species than for deciduous species. The number of buds/10 cm of shoot was higher in evergreens. However, the per cent mortality was also higher in them.     

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.     

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.     

Leaf and bud demography and shoot growth in evergreen and deciduous trees of central Himalaya, India

Leaf and bud demography and shoot growth were studied in 10 evergreen (ES) and 15 deciduous (DS) tree species occurring between 600 and 2200 m elevation in the central Himalayan mountains in India. Results were analyzed to help explain why ES prevail in the vegetation of this region, even though the number of ES is no greater than for DS. Although each species had its own pattern with regard to leaf and bud demography and seasonality of shoot extension and radial growth, it was possible to group the species on the basis of shoot growth phenology. In most species, leaves emerged during March-April, at the onset of warm and dry summer season. The ES recruit leaves in shoots more rapidly than the DS. Across all species, peak number of leaves per shoot (5.8–20.7), peak leaf area per shoot (116.2–1559.2 cm²), peak number of vegetative buds per shoot (1.9–14.5), bud survival per shoot (23–84%), shoot extension growth (6.4–40.8 cm) and shoot extension period (13–30 weeks) varied considerably. The peak leaf area per shoot (587.7 vs. 246.7 cm²) and shoot extension growth (19.3 vs. 11.2 cm) were significantly greater for DS than for ES, and these two functional groups of species were clearly separable with regard to shoot growth characteristics.Results indicate that rapid recruitment of leaf crop in the shoots, longer leaf life-span, and access to ground water due to deep roots were some of the advantages, the ES had over the DS, that may have likely enable them to maintain growth for a longer period in this region of warm winters and longer winter day length as compared to temperate climates. In the shallow rooted DS, shoot growth seems to be much affected by a seasonal drought in winter and they are likely to be affected more in the event of failure of monsoon rains in this region.     

Seasonal changes in stem diameter and leaf development in a tropical montane forest

Abstract. Seasonal patterns of stem diameter changes in evergreen and deciduous species of a tropical montane forest in the Central Himalayas (300–2250 m a.s.l.) were investigated in relation to leaf development. Ca. 75 % of the annual rainfall in this region occurs in a short period, from mid-June to mid-September and the remaining months are dry. It was assumed that changes in stem diameter are correlated with changes in water stress. Each evergreen species could be characterized by leaf longevity of about one year; each species showed pronounced summer leaf drop and simultaneous new leaf formation. Winter stem shrinkage was more pronounced in deciduous species than in evergreen ones. The deciduous species also showed a greater proportional loss of leaf mass (before abscission) than the evergreen species. Winter leaf fall in deciduous species was related to the pronounced stem shrinkage. The leaf fall enabled these species to control further water loss. Being more resistant to desiccation, the evergreen species retained their leaves throughout the winter but showed gradual loss of leaf mass, presumably in order to control water loss. In all species, leaf expansion was completed before the onset of the rainy season, when water stress was high. This strategy has definite advantages in a climate with a monsoon pattern of rainfall. Evergreen species, showing pronounced leaf drop in summer, have advantages over deciduous species; hence their preponderance in the region.     

Woody plant phenology in the West Africa savanna

In the savanna of West Africa the seasonality of rainfall, with a drought period of at least four months, strongly influences the vegetation. Rainfall is a very critical abiotic variable and therefore plant species must be well adapted to survive in this habitat.
In our research, phenological patterns of 120 woody plant species have been examined based on the presence of green leaves. According to the patterns found, these species can be classified in phenological groups, which represent different strategies for survival. Two extreme strategies are found to resist drought: (1) by using the waterstorage in the deeper soil layers and river beds and by restricting drought-damage through scleromorphic features, and (2) by avoiding the drought through foliage shedding in the dry period.
The first strategy is represented by the riparian and upland evergreens , and the semi-evergreens . The evergreens bear leaves the whole year, gradually replacing old leaves by new ones. The riparian evergreens are strictly bound to riverbeds and grow in or immediately adjacent to them. The semi-evergreens shed their leaves and start sprouting during a short period (one-two weeks) once a year. Because the evergreens and the semi-evergreens are in leaf in the dry period they have to protect themselves to drought damage by scleromorphic features.
Contrary to these species are the deciduous species which are bare for at least some months per year. When the dry season starts their leaves dry out and are subsequently shed. They start sprouting before or at the beginning of the first rains. Although much less in number, some deciduous trees also have scleromorphic features to resist drought-damage. The strategy of sprouting just before the rainy season begins indicates that certain water resources remain available to these deep-rooting woody plants throughout the year, providing them with a fully operating photosynthetic apparatus when favourable conditions arrive.     

Seasonal patterns of leaf production in co‐occurring trees with contrasting leaf phenology: time and quantitative divergences

Neotropical savannas (‘cerrados’) of Central Brazil are characterized by the coexistence of a large diversity of tree species with divergent phenological behaviors, which reflect a great diversity in growth strategies. In the present study time behavior and quantitative aspects of shoot growth, shoot mortality, and leaf longevity and production were analyzed in 12 woody species of contrasting leaf phenology, adopting a functional group approach where 12 species were categorized into three functional groups: evergreen, decidous and brevideciduous, according to their leaf phenology. Shoot growth and leaf production were seasonal for the three functional groups, differing in their time of occurrence, but being concentrated during the last months of the dry season. Shoot growth differed between evergreens and deciduous, as well leaf production. Evergreens had higher rates of shoot growth, produced a higher number of leaves and had longer leaf longevity (around 500 days against 300 days in deciduous and brevideciduous). Leaf longevity was associated with patterns of leaf production when accounting for all phenological groups studied. It was possible to identify different patterns of aerial growth in savanna phenological groups, providing evidence of great functional variability amongst the groups studied.     

Relative growth rate in phylogenetically related deciduous and evergreen woody species

Relative growth rate (RGR) and other growth parameters were studied in eight pairs of closely related deciduous and evergreen species (within the same genus or family). The main objective of this study was to test the association between leaf turnover rate and RGR, specific leaf area (SLA, leaf area/leaf dry weight) and other growth variables. Plants were grown for 6 months in a greenhouse under favourable water and nutrient conditions. Variation in RGR among the 16 woody species was due mainly to differences in morphological parameters such as leaf area ratio (LAR, whole plant area/whole plant dry weight) and SLA). However, temporal variation in RGR within species was due mainly to variation in net assimilation rate. When phylogeny was not taken into account, analyses showed that deciduous species grew faster than evergreens. In contrast, when phylogeny was taken into account, the data analysis showed that a faster RGR is not consistently associated with the deciduous habit (in five pairs it was, but in the other three it was not). The faster growth of the deciduous trees (in the five positive contrasts) could be explained by their higher LAR and higher SLA relative to evergreens. The lack of differences in RGR between deciduous and evergreens (in three pairs) was due to the higher leaf mass ratio (LMR, leaf dry biomass/total dry biomass) for the evergreens, which offset the higher SLA of the deciduous species, resulting in a similar LAR in both functional groups (LAR=LMR2SLA). Deciduous species had consistently higher SLA than evergreens. We suggest that SLA, more than RGR, could be an important parameter in determining adaptive advantages of deciduous and evergreen species.     

Leaf Mass per Area (LMA) and Its Relationship with Leaf Structure and Anatomy in 34 Mediterranean Woody Species along a Water Availability Gradient

Leaf mass per area (LMA) is a morphological trait widely used as a good indicator of plant functioning (i.e. photosynthetic and respiratory rates, chemical composition, resistance to herbivory, etc.). The LMA can be broken down into the leaf density (LD) and leaf volume to area ratio (LVA or thickness), which in turn are determined by anatomical tissues and chemical composition. The aim of this study is to understand the anatomical and chemical characteristics related to LMA variation in species growing in the field along a water availability gradient. We determined LMA and its components (LD, LVA and anatomical tissues) for 34 Mediterranean (20 evergreen and 14 deciduous) woody species. Variation in LMA was due to variation in both LD and LVA. For both deciduous and evergreen species LVA variation was strongly and positively related with mesophyll volume per area (VA or thickness), but for evergreen species positive relationships of LVA with the VA of epidermis, vascular plus sclerenchyma tissues and air spaces were found as well. The leaf carbon concentration was positively related with mesophyll VA in deciduous species, and with VA of vascular plus sclerenchymatic tissues in evergreens. Species occurring at the sites with lower water availability were generally characterised by a high LMA and LD.     

Construction and maintenance costs of mediterranean-climate evergreen and deciduous leaves I. Growth and CO2 exchange analysis

Summary Gas exchange and leaf growth analysis were used in conjunction to determine leaf-construction and maintenance costs in three co-occurring shrubs of the california chaparral, one evergreen, Heteromeles arbutifolia, and two drougth deciduous species, Lepechinia calycina, and Diplacus aurantiacus. The construction costs per unit of leaf weight were similar among the three species and very close to values reported for other evergreens but considerably higher than leaf construction costs for other deciduous or herbaceous plants. Maintenance costs per unit of leaf weight were significantly greater in one deciduous species, L. calycina, than in the evergreen. Maintenance costs for all species were in the range reported for herbaceous species and considerably above those reported for other evergreens.     

Leaf functional traits of Neotropical savanna trees in relation to seasonal water deficit

The seasonal savannas (cerrados) of Central Brazil are characterized by a large diversity of evergreen and deciduous trees, which do not show a clear differentiation in terms of active rooting depth. Irrespective of the depth of the root system, expansion of new foliage in deciduous species occurs at the end of the dry season. In this study, we examined a suite of leaf traits related to C assimilation, water and nutrients (N, P) in five deciduous and six evergreen trees that were among the dominant families of cerrado vegetation. Maximum CO₂ assimilation on a mass basis (A_mass) was significantly correlated with leaf N and P, and specific leaf area (SLA; leaf area per unit of leaf mass). The highest leaf concentrations of both nutrients were measured in the newly mature leaves of deciduous species at the end of the dry period. The differences in terms of leaf N and P between evergreen and deciduous species decreased during the wet season. Deciduous species also invested less in the production of non-photosynthetic leaf tissues and produced leaves with higher SLA and maintained higher water use efficiency. Thus, deciduous species compensated for their shorter leaf payback period by maintaining higher potential payback capacity (higher values of A_mass) and lower leaf construction costs (higher SLA). Their short leafless period and the capacity to flush by the end of the dry season may also contribute to offset the longer payback period of evergreen species, although it may involve the higher cost of maintaining a deep-root system or a tight control of plant water balance in the shallow-rooted ones.     

Differential impact of liana colonization on the leaf functional traits of co-occurring deciduous and evergreen trees in a tropical dry scrub forest

The present study was carried out to analyze the leaf functional traits of co-occurring evergreen and deciduous tree species in a tropical dry scrub forest. This study also intended to check whether the species with contrasting leaf habits differ in their leaf trait plasticity, responding to the canopy infestation by lianas. A total of 11 leaf functional traits were studied for eight tree species with contrasting leaf habits (evergreen and deciduous) and liana-colonization status (with or without liana). In the liana-free environment (L^–), evergreen trees had significantly higher leaf tissue density (LTD) and total chlorophyll (CHL_t) than the deciduous species. Whereas the deciduous trees had higher specific leaf area (SLA) and mass-based leaf nitrogen concentration (N_mass). The leaf trait-pair relationship in the present study agreed with the well-established global trait-pair relationships (leaf thickness (LT) vs. SLA, N_mass vs. LT, SLA vs. N_mass, and LDMC vs. SLA). There was a significant difference between L⁺ and L^– individuals in leaf area (LA), petiole length (PL), SLA, LDMC, and CHL_t in the deciduous species. On the other hand, evergreen species showed marked differences across LT, SLA, LTD, N_mass, and chlorophyll components between L⁺ and L^– individuals of the same species. The results revealed the differential impact of liana colonization on the host trees with contrasting leaf habits. The deciduous species with the acquisitive strategy can have a competitive advantage over evergreen species in the exposed environments (L^–), whereas evergreen species with shade-tolerant properties were better acclimated to the shaded environments (L⁺). Therefore, liana colonization can significantly impact the C-fixation strategies of the host trees by altering their light environment and further, the magnitude of such impact may vary among species of different leaf habits. The result also indicated the patterns of convergence and divergence in some of the leaf functional traits between evergreen and deciduous species explaining the patterns of species co-existence.

     

Vegetative phenology of alpine plants at Tateyama Murodo-daira in central Japan

The vegetative phenology of 29 alpine species, including herbaceous and woody summergreens and evergreens, was investigated. Summergreen species initiated and completed leaf growth earlier than evergreen species. The green period of leaves in summergreen plants was determined largely by the time of growth initiation. Early initiation of growth in summergreen plants contributes to the increase in photosynthetic carbon gain. Early cessation of growth in summergreens is advantageous for the growth in the following year because it leads to an increase in stored photosynthates. The growth period of leaves and stems in alpine plants correlated with the time of growth initiation more strongly than with the time of growth cessation, indicating the importance of early growth initiation for the increase in plant growth. The growth period of leaves was positively correlated with the sum of leaf lengths and the number of leaves. Herbs with a long growth period of more than 50 days had perennial shoot axes not terminated by inflorescences, suggesting a relationship between the growth period and shoot habit. Two summergreen species were completely dead by mid-September, before the air temperature decreased below 0°C. The remaining summergreen species died immediately after the air temperature decreased to −1.4°C in late September.     

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.     

Nutrient conservation strategies in native Andean‐Patagonian forests

Abstract. Nutrient conservation in vegetation affects rates of litter decomposition and soil nutrient availability. Although resorption has been traditionally considered one of the most important plant strategies to conserve nutrients in temperate forests, long leaf life‐span and low nutrient requirements have been postulated as better indicators. We aimed at identifying nutrient conservation strategies within characteristic functional groups of NW Patagonian forests on Andisols. We analysed C‐, N‐, P‐, K‐ and lignin‐concentrations in mature and senescent leaves of ten native woody species within the functional groups: broad‐leaved deciduous species, broad‐leaved evergreens and conifers. We also examined mycorrhizal associations in all species. Nutrient concentration in mature leaves and N‐ resorption were higher in broad‐leaved deciduous species than in the other two functional groups. Conifers had low mature leaf nutrient concentrations, low N‐resorption and high lignin/N ratios in senescent leaves. P‐ and K‐resorptions did not differ among functional groups. Broad‐leaved evergreens exhibited a species‐dependent response. Nitrogen in mature leaves was positively correlated with both N resorption and soil N‐fertility. Despite the high P‐retention capacity of Andisols, N appeared to be the more limiting nutrient, with most species being proficient in resorbing N but not P. The presence of endomycorrhizae in all conifers and the broad‐leaved evergreen Maytenus boaria, ectomycorrhizae in all Nothofagus species (four deciduous, one evergreen), and cluster roots in the broad‐leaved evergreen Lomatia hirsuta, would be possibly explaining why P is less limiting than N in these forests.     

Global meta-analysis shows that relationships of leaf mass per area with species shade tolerance depend on leaf habit and ontogeny

It was predicted that relationships of leaf mass per area (LMA) with juvenile shade tolerance will depend on leaf habit, and on whether species are compared at a common age as young seedlings, or at a common size as saplings. A meta-analysis of 47 comparative studies (372 species) was used to test predictions, and the effect of light environment on this relationship. The LMA of evergreens was positively correlated with shade tolerance, irrespective of ontogeny or light environment. The LMA of young seedlings (相似文献   

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.     

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 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.     

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.