Forest growth along a rainfall gradient in Hawaii: Acacia koa stand structure,productivity, foliar nutrients,and water- and nutrient-use efficiencies

We tested whether variation in growth of native koa (Acacia koa) forest along a rainfall gradient was attributable to differences in leaf area index (LAI) or to differences in physiological performance per unit of leaf area. Koa stands were studied on western Kauai prior to Hurricane Iniki, and ranged from 500 to 1130 m elevation and from 850 to 1800 mm annual precipitation. Koa stands along the gradient had basal area ranging from 8 to 42 m²/ha, LAI ranging from 1.4 to 5.4, and wood increment ranging from 0.7 to 7.1 tonnes/ha/year. N, P, and K contents by weight of sun leaves (phyllodes) were negatively correlated with specific leaf mass (SLM, g m^-2) across sites; on a leaf area basis, N increased whereas P and K decreased with SLM. LAI, aboveground woody biomass increment, and production per unit leaf area (E) increased as phyllode ¹³C became more negative. The ¹³C data suggested that intrinsic water-use efficiency (ratio of assimilation to conductance) increased as water availability decreased. In five of the six sites, phyllode P contents increased as LAI increased, but biomass increment and E were not correlated with phyllode nutrient contents, suggesting that productivity was limited more by water than by nutrient availability. Because vapor pressure deficits increased with decreasing elevation, actual water-use efficiency (ratio of assimilation to transpiration) was lower at drier, low-elevation sites. There was a trade-off between intrinsic water-use efficiency and production per unit of canopy N or P across the gradient. In summary, koa responds to water limitation both by reducing stand LAI and by adjusting gas exchange, which results in increased intrinsic water-use efficiency but decreased E.     

Influence of stand structure on carbon-13 of vegetation, soils, and canopy air within deciduous and evergreen forests in Utah, United States

Carbon isotope ratios (δ¹³C) were studied in evergreen and deciduous forest ecosystems in semi-arid Utah (Pinus contorta, Populus tremuloides, Acer negundo and Acer grandidentatum). Measurements were taken in four to five stands of each forest ecosystem differing in overstory leaf area index (LAI) during two consecutive growing seasons. The δ¹³C_leaf (and carbon isotope discrimination) of understory vegetation in the evergreen stands (LAI 1.5–2.2) did not differ among canopies with increasing LAI, whereas understory in the deciduous stands (LAI 1.5–4.5) exhibited strongly decreasing δ¹³C_leaf values (increasing carbon isotope discrimination) with increasing LAI. The δ¹³C values of needles and leaves at the top of the canopy were relatively constant over the entire LAI range, indicating no change in intrinsic water-use efficiency with overstory LAI. In all canopies, δ¹³C_leaf decreased with decreasing height above the forest floor, primarily due to physiological changes affecting c _i/c _a (> 60%) and to a minor extent due to δ¹³C of canopy air (< 40%). This intra-canopy depletion of δ¹³C_leaf was lowest in the open stand (1‰) and greatest in the denser stands (4.5‰). Although overstory δ¹³C_leaf did not change with canopy LAI, δ¹³C of soil organic carbon increased with increasing LAI in Pinus contorta and Populus tremuloides ecosystems. In addition, δ¹³C of decomposing organic carbon became increasingly enriched over time (by 1.7–2.9‰) for all deciduous and evergreen dry temperate forests. The δ¹³C_canopy of CO₂ in canopy air varied temporally and spatially in all forest stands. Vertical canopy gradients of δ¹³C_canopy, and [CO₂]_canopy were larger in the deciduous Populus tremuloides than in the evergreen Pinu contorta stands of similar LAI. In a very wet and cool year, ecosystem discrimination (Δ_e) was similar for both deciduous Populus tremulodies (18.0 ± 0.7‰) and evergreen Pinus contorta (18.3 ± 0.9‰) stands. Gradients of δ¹³C_canopy and [CO₂]_canopy were larger in denser Acer spp. stands than those in the open stand. However, ¹³C enrichment above and photosynthetic draw-down of [CO₂]_canopy below tropospheric baseline values were larger in the open than in the dense stands, due to the presence of a vigorous understory vegetation. Seasonal patterns of the relationship δ¹³C_canopy versus 1/[CO₂]_canopy were strongly influenced by precipitation and air temperature during the growing season. Estimates of Δ_e for Acer spp. did not show a significant effect of stand structure, and averaged 16.8 ± 0.5‰ in 1933 and 17.4 ± 0.7‰ in 1994. However, Δ_e varied seasonally with small fluctuations for the open stand (2‰), but more pronounced changes for the dense stand (5‰). Received: 15 April 1996 / Accepted: 19 October 1996     

Interspecific variability of δ13C among trees in rainforests of French Guiana: functional groups and canopy integration

The interspecific variability of sunlit leaf carbon isotope composition (δ¹³C), an indicator of leaf intrinsic water-use efficiency (WUE, CO₂ assimilation rate/leaf conductance for water vapour), was investigated in canopy trees of three lowland rainforest stands in French Guiana, differing in floristic composition and in soil drainage characteristics, but subjected to similar climatic conditions. We sampled leaves with a rifle from 406 trees in total, representing 102 species. Eighteen species were common to the three stands. Mean species δ¹³C varied over a 6.0‰ range within each stand, corresponding to WUE varying over about a threefold range. Species occurring in at least two stands displayed remarkably stable δ¹³C values, suggesting a close genetic control of species δ¹³C. Marked differences in species δ¹³C values were found with respect to: (1) the leaf phenology pattern (average δ¹³C=–29.7‰ and –31.0‰ in deciduous-leaved and evergreen-leaved species, respectively), and (2) different types of shade tolerance defined by features reflecting the plasticity of growth dynamics with respect to contrasting light conditions. Heliophilic species exhibited more negative δ¹³C values (average δ¹³C=–30.5‰) (i.e. lower WUE) than hemitolerant species (–29.3‰). However, tolerant species (–31.4‰) displayed even more negative δ¹³C values than heliophilic ones. We could not provide a straightforward ecophysiological interpretation of this result. The negative relationship found between species δ¹³C and midday leaf water potential (Ψ_wm) suggests that low δ¹³C is associated with high whole tree leaf specific hydraulic conductance. Canopy carbon isotope discrimination (Δ _A ) calculated from the basal area-weighed integral of the species δ¹³C values was similar in the three stands (average Δ _A =23.1‰), despite differences in stand species composition and soil drainage type, reflecting the similar proportions of the three different shade-tolerance types among stands. Received: 30 November 1999 / Accepted: 23 March 2000     

Relationships of stable carbon isotopes, plant water potential and growth: an approach to asses water use efficiency and growth strategies of dry land agroforestry species

The relationships between annual wood stable carbon isotope composition (δ¹³C), dry season midday plant water potential, and annual growth rate were investigated to asses the ability of agroforestry species to adapt to climate changes. 6–8 stem disks from four co-occurring species (Acacia senegal, A. seyal, A. tortilis and Balanites aegyptiaca) were collected for radial growth measurements using tree-ring analysis spanning 1930–2003. Annual δ¹³C was measured on three tree disks per species for the period 1970–2002. Midday plant water potential was measured during the dry season. Annual radial growth and midday plant water potential ranged from 0.27 to 9.12 mm and −1.0 to −5.0 MPa, respectively, with statistically significant differences. After correcting annual wood δ¹³C for atmospheric changes in δ¹³C, carbon isotopic composition ranged from −22.22 to −26.58‰. Relationships between δ¹³C, radial growth and plant water potentials revealed the interaction of water availability, stomatal conductance, δ¹³C values and growth. Two contrasting water use strategies and competitive advantages can be distinguished. Species with lower mean δ¹³C values (A. senegal and A. seyal) show high plant water potential and, hence, better growth during moist years. Thus, they indicate low water use efficiency (WUE) and opportunistic water use strategy. On the other hand, species with lower water potentials (A. tortilis and B. aegyptiaca) showed relative better growth performance and less increase in δ¹³C in drought years, reflecting their high WUE and conservative water use strategy. These results suggest that δ¹³C in tree rings can be useful in estimating historic changes in plant WUE and hence in screening drought tolerant species in the face of expected climate changes, as well as for assessing the functional diversity and risk reduction in mixed vegetation.     

Seasonal variations in leaf δ<Superscript>13</Superscript>C and nitrogen associated with foliage turnover and carbon gain for a wet subalpine fir forest in the Gongga Mountains,eastern Tibetan Plateau

It is still unclear to what extent variations in foliar δ¹³C and nitrogen can be used to detect seasonal changes in canopy productivity. We hypothesize that in a wet and cloudy fir forest, seasonally higher litterfall and lower leaf area index (LAI) are correlated with higher mass-based leaf nitrogen (N _mass) and net primary productivity (NPP), while foliar δ¹³C may change with specific leaf area (SLA), area-based leaf nitrogen (N _area), and/or starch concentration. In order to test our hypotheses, stand-level litterfall and the means of δ¹³C, N _mass, N _area, SLA, and starch concentration of canopy needles for a wet and cloudy Abies fabri forest in the Gongga Mountains were monthly measured during the growing season. Seasonal estimates of LAI were obtained from our previous work. A conceptual model was used to predict seasonal NPP of the fir forest. Seasonal mean δ¹³C and N _mass and climatic variables were used as inputs. The δ¹³C across 1–7-year-old needles increased from May to September associated with decreasing SLA and increasing N _area. There were no significant differences in seasonal starch concentration. With increasing litterfall and decreasing LAI, seasonal mean N _mass increased, while the δ¹³C varied little. The simulated NPP increased with increasing litterfall and related traits of N _mass and N _area. Our data generally supported the hypotheses. The results also suggest that in the forest with relatively moist and cloudy environment, the largest fraction of annual carbon gain may occur in the early part of the growing season when higher litterfall results in higher N _mass of canopy leaves.     

Water deficit and plant competition effects on growth and water-use efficiency of white clover ( Trifolium repens,L.) and ryegrass (Lolium perenne,L.)

The combined effects of soil water deficit and above and below ground interspecific plant competition on the growth, water-use efficiency (WUE), and measured carbon isotopic composition (δ¹³C) values of white clover and ryegrass were studied. White clover and ryegrass were grown in specially designed crates 1) individually; 2) in shoot competition; or 3) in shoot + root competition and either well-watered or at a moderate or severe soil water deficit. The effects of shoot + root competition on shoot dry matter growth were substantial and benefited both white clover and ryegrass when well-watered or at a moderate soil water deficit, while severely reducing white clover shoot dry matter growth at severe soil water deficit. Plant competition did not affect the WUE of white clover or ryegrass. As soil water deficit increased, the WUE of white clover did not change whereas the WUE of ryegrass increased and was greater than that of white clover. This was attributed to the lower leaf water conductance of ryegrass which conserved water and maintained growth longer compared to white clover. A stronger correlation existed between soil water deficit and measured δ¹³C values for ryegrass at each plant competition level (P<0.001) than existed for white clover (individual: P<0.01; shoot + root: P<0.001; shoot: P<0.10). Unlike white clover, the relationship between measured δ¹³C values and shoot dry matter growth indicated that C assimilation for ryegrass was dependent on type of plant competition. That WUE remained constant for white clover while measured δ¹³C values increased as soil water deficit increased, suggests that the role below ground respiration rate played in determining δ¹³C values increased. The WUE of white clover appears to be independent of the nature of the competition between plants and the soil water deficit level at which it is grown, whereas for ryegrass, the addition of root competition to shoot competition should lead to increases in its WUE. This revised version was published online in June 2006 with corrections to the Cover Date.     

Carbon and Nitrogen Dynamics of Boreal Jack Pine Stands With and Without a Green Alder Understory

We compared the species composition, structure and selected components of the carbon (C) and nitrogen (N) budgets of similar-aged, mature boreal jack pine (Pinus banksiana Lamb.) forests with and without green alder [Alnus crispa (Ait.) Pursh.] in two different boreal environments. The C and N content of the overstory biomass components (for example, stem, branch, and foliage), total vegetation, forest floor, and mineral soil were greater (P= 0.05 to P= 0.10) for jack pine with alder (JPA) stands than for jack pine without alder (JP) stands at both study areas. Jack pine foliage N isotopic discrimination (δ¹⁵N) and annual litterfall N content were significantly greater (P < 0.05) in the JPA than the JP stands at both study areas, suggesting that alder was fixing N and that N availability was greater in the JPA than the JP stands. The greater leaf area index (LAI) and overstory C accumulation in the JPA than the JP stands (P < 0.05) is likely because of the greater N availability in the JPA stands, but the effect of soil texture discontinuity on water availability in the JPA stands can not be dismissed. Percent ground cover by feathermoss varied among the jack pine communities and was positively correlated with overstory LAI (r ²= 0.83, P< 0.05). One index of N-use efficiency (NUE), defined as aboveground net primary productivity (ANPP) per litterfall N, was significantly greater (P < 0.05) for the JP than the JPA stands, but a second index of NUE, ANPP/N uptake, did not differ between the two jack pine communities. Jack pine trees growing without alder produced more organic matter per unit of N, but percent N retranslocation from senescing foliage and N mean residence time in the overstory did not differ between the JPA and the JP stands. A conceptual model is presented that illustrates the potential influence of alder on the species composition, structure, and function of boreal jack pine forests. Received 6 January 1998; accepted 15 April 1998.     

Inter-annual and seasonal variability of radial growth, wood density and carbon isotope ratios in tree rings of beech (Fagus sylvatica) growing in Germany and Italy

We investigated the variability of tree-ring width, wood density and ¹³C/¹²C in beech tree rings (Fagus sylvatica L.), and analyzed the influence of climatic variables and carbohydrate storage on these parameters. Wood cores were taken from dominant beech trees in three stands in Germany and Italy. We used densitometry to obtain density profiles of tree rings and laser-ablation-combustion-GC-IRMS to estimate carbon isotope composition (δ ¹³C) of wood. The sensitivity of ring width, wood density and δ ¹³C to climatic variables differed; with tree-ring width responding to environmental conditions (temperature or precipitation) during the first half of a growing season and maximum density correlated with temperatures in the second part of a growing season (July–September). δ ¹³C variations indicate re-allocation and storage processes and effects of drought during the main growing season. About 20% of inter-annual variation of tree-ring width was explained by the tree-ring width of the previous year. This was confirmed by δ ¹³C of wood which showed a contribution of stored carbohydrates to growth in spring and a storage effect that competes with growth in autumn. Only mid-season δ ¹³C of wood was related to concurrent assimilation and climate. The comparison of seasonal changes in tree-ring maximum wood density and isotope composition revealed that an increasing seasonal water deficit changes the relationship between density and ¹³C composition from a negative relation in years with optimal moisture to a positive relationship in years with strong water deficit. The climate signal, however, is over-ridden by effects of stand density and crown structure (e.g., by forest management). There was an unexpected high variability in mid season δ ¹³C values of wood between individual trees (−31 to −24‰) which was attributed to competition between dominant trees as indicated by crown area, and microclimatological variations within the canopy. Maximum wood density showed less variation (930–990 g cm⁻³). The relationship between seasonal changes in tree-ring structure and ¹³C composition can be used to study carbon storage and re-allocation, which is important for improving models of tree-ring growth and carbon isotope fractionation. About 20–30% of the tree-ring is affected by storage processes. The effects of storage on tree-ring width and the effects of forest structure put an additional uncertainty on using tree rings of broad leaved trees for climate reconstruction.     

Variation in woody plant δ13C along a topoedaphic gradient in a subtropical savanna parkland

δ¹³C values of C₃ plants are indicators of plant carbon–water relations that integrate plant responses to environmental conditions. However, few studies have quantified spatial variation in plant δ¹³C at the landscape scale. We determined variation in leaf δ¹³C, leaf nitrogen per leaf area (N_area), and specific leaf area (SLA) in April and August 2005 for all individuals of three common woody species within a 308 × 12-m belt transect spanning an upland–lowland topoedaphic gradient in a subtropical savanna in southern Texas. Clay content, available soil moisture, and soil total N were all negatively correlated with elevation. The δ¹³C values of Prosopis glandulosa (deciduous N₂-fixing tree legume), Condalia hookeri (evergreen shrub), and Zanthoxylum fagara (evergreen shrub) leaves increased 1–4‰ with decreasing elevation, with the δ¹³C value of P. glandulosa leaves being 1–3‰ higher than those of the two shrub species. Contrary to theory and results from previous studies, δ¹³C values were highest where soil water was most available, suggesting that some other variable was overriding or interacting with water availability. Leaf N_area was positively correlated with leaf δ¹³C of all species (p < 0.01) and appeared to exert the strongest control over δ¹³C along this topoedaphic gradient. Since leaf N_area is positively related to photosynthetic capacity, plants with high leaf N_area are likely to have low p _I/p _a ratios and therefore higher δ¹³C values, assuming stomatal conductance is constant. Specific leaf area was not correlated significantly with leaf δ¹³C. Following a progressive growing season drought in July/August, leaf δ¹³C decreased. The lower δ¹³C in August may reflect the accumulation of ¹³C-depleted epicuticular leaf wax. We suggest control of leaf δ¹³C along this topoedaphic gradient is mediated by leaf N_area rather than by stomatal conductance limitations associated with water availability.     

Forest Canopy Properties and Variation in Aboveground Net Primary Production over Upper Great Lakes Landscapes

Relationships among aboveground net primary production (ANPP) and forest canopy properties were investigated in secondary successional forests of similar age and disturbance history in northern Lower Michigan, USA. Aboveground biomass, ANPP, canopy leaf area index (LAI), and several canopy nitrogen (N) measures were estimated from 12 stands representing major landform-level ecosystems and vegetation associations. Stand single-date and growing season average normalized difference vegetation indices (NDVI) were derived from Landsat TM. ANPP correlated most strongly with total canopy N content (r ² = 0.81, P < 0.001), followed by LAI (r ² = 0.73, P < 0.001) and area-based canopy-average leaf N concentration (r ² = 0.37, P < 0.05). No significant relationship was detected between ANPP and mass-based canopy-average leaf N concentration. Stand ANPP correlated positively with both total canopy N content (r ² = 0.62, P < 0.05) and mass-based leaf N concentration (r ² = 0.53, P < 0.05) of commonly dominant Populus spp. Relatively higher ANPP, total canopy N content and LAI corresponded to simultaneous presence of shade-intolerant P. grandidentata with shade-tolerant species. Both forms of NDVI were significantly related to ANPP, and more strongly to total canopy N content and LAI; relationships were stronger for seasonally averaged (r ² ≥ 0.75, P < 0.001) than for single-date NDVI (r ² ≥ 0.52, P < 0.01). Results indicate that on the transitioning study landscapes, ANPP was more closely related to canopy N content than to LAI, seasonally averaged NDVI was a more reliable predictor of ANPP and canopy properties than the single-date index, whereas measured canopy characteristics varied significantly between major landform-level ecosystems. The ongoing decline of P. grandidentata is likely to alter aboveground carbon and pools and fluxes in the course of succession.     

Stable isotope composition of <Emphasis Type="Italic">Chara rudis</Emphasis> incrustation in Lake Jasne,Poland

Stable isotope composition (δ¹³C and δ¹⁸O) was analysed in mineral incrustation of Chara rudis and surrounding waters. This macroalga forms dense and extensive charophyte meadows and may significantly contribute to the calcium carbonate precipitation and deposition of marl lake sediments. The study aimed to find out if charophyte calcium carbonate was precipitated in an isotopic equilibrium with lake water and if the precipitation was related to the environmental conditions. Two apical internodes of 10 individuals of C. rudis were collected monthly between June and late October 2008 at three permanent study sites (1.0 m, 1.5 m and 2.0 m deep) in a small (15.1 ha) and shallow (mean depth: 4.3 m) mid-forest lake with extensively developed charophyte meadows (Lake Jasne, mid-Western Poland). Basic physical–chemical analyses were performed at each study site, and water samples for further laboratory determinations, including stable isotope analyses, were collected from the above searched C. rudis stands and, simultaneously, at three comparative sites in the macrophyte-free pelagial. The difference in δ¹³C between incrustation and water from above C. rudis exceeded 2‰ V-PDB at each site. In the case of δ¹⁸O, it exceeded 2‰ between July and September. Accordingly, it is postulated that calcium carbonate was not precipitated in an isotopic equilibrium with lake water. Incrustation was enriched in heavier carbon isotope, ¹³C, and water was enriched in ¹⁸O. δ¹³C of incrustation and DIC were positively correlated, whereas negative relation was found between δ¹⁸O of incrustation and water. Several dependencies were found with water chemistry above the plants. The content of mineral incrustation in Chara dry weight had negative influence on the δ¹⁸O but not on the δ¹³C. Community depth, structure and PVI had no effect. No significant differences appeared between isotope composition in the pelagic zone and Chara stands.     

Aboveground Net Primary Production and Leaf-Area Index in Early Postfire Vegetation in Yellowstone National Park

Aboveground net primary production (ANPP) and leaf-area index (LAI) of lodgepole pine (Pinus contorta var. latifolia Engelm. ex Wats.) saplings and aboveground productivity of herbaceous vegetation components were determined 9 years after the 1988 fires in Yellowstone National Park (YNP). Measurements were made in four sites representing a wide range of early postfire vegetation present in YNP, including high-density lodgepole pine, low-density lodgepole pine, and two nonforest stands. LAI of the pine saplings and total ANPP (trees plus herbs) generally increased with increasing sapling density, from 0.002 m² m^{− 2} and 0.25 Mg ha^{− 1} year^{− 1} in the infertile nonforest stand (100 pine saplings ha^{− 1}) to 1.8 m² m^{− 2} and 4.01 Mg ha^{− 1} year^{− 1} in the high-density pine stand (62,800 saplings ha^{− 1}). Aboveground herbaceous productivity was not strongly correlated with sapling density, but appeared to be influenced by soil fertility. In the high-density pine stand, tree ANPP and LAI were within the lower range of values reported for similar mature coniferous forests. This finding suggests that at least some ecosystem processes (related to ANPP and LAI) may have nearly recovered after only 9 years of postfire succession, in at least some of the young forests developing after the 1988 Yellowstone fires. Received 7 April 1998; accepted 1 December 1998.     

Intra-annual variability of anatomical structure and δ<Superscript>13</Superscript>C values within tree rings of spruce and pine in alpine,temperate and boreal Europe

Tree-ring width, wood density, anatomical structure and ¹³C/¹²C ratios expressed as δ¹³C-values of whole wood of Picea abies were investigated for trees growing in closed canopy forest stands. Samples were collected from the alpine Renon site in North Italy, the lowland Hainich site in Central Germany and the boreal Flakaliden site in North Sweden. In addition, Pinus cembra was studied at the alpine site and Pinus sylvestris at the boreal site. The density profiles of tree rings were measured using the DENDRO-2003 densitometer, δ¹³C was measured using high-resolution laser-ablation-combustion-gas chromatography-infra-red mass spectrometry and anatomical characteristics of tree rings (tracheid diameter, cell-wall thickness, cell-wall area and cell-lumen area) were measured using an image analyzer. Based on long-term statistics, climatic variables, such as temperature, precipitation, solar radiation and vapor pressure deficit, explained <20% of the variation in tree-ring width and wood density over consecutive years, while 29–58% of the variation in tree-ring width were explained by autocorrelation between tree rings. An intensive study of tree rings between 1999 and 2003 revealed that tree ring width and δ¹³C-values of whole wood were significantly correlated with length of the growing season, net radiation and vapor pressure deficit. The δ¹³C-values were not correlated with precipitation or temperature. A highly significant correlation was also found between δ¹³C of the early wood of one year and the late wood of the previous year, indicating a carry-over effect of the growing conditions of the previous season on current wood production. This latter effect may explain the high autocorrelation of long-term tree-ring statistics. The pattern, however, was complex, showing stepwise decreases as well as stepwise increases in the δ¹³C between late wood and early wood. The results are interpreted in the context of the biochemistry of wood formation and its linkage to storage products. It is clear that the relations between δ¹³C and tree-ring width and climate are multi-factorial in seasonal climates.     

Carbon isotope composition,water-use efficiency and biomass productivity of Eucalyptus microtheca populations under different water supplies

Variation in carbon isotope composition (δ¹³C), water-use efficiency (WUE) and biomass productivity were compared among three populations of Eucalyptus microtheca F. Muell. in a greenhouse. Seedlings were maintained under one well-watered (Control, keeping the soil at field capacity) and two different water deficit conditions (Drought stress I, keeping the same soil water content; Drought stress II, keeping the same soil water supply). In each treatment, significant population differences in δ¹³C, WUE, and dry matter accumulation and allocation were detected. A negative correlation between WUE and biomass productivity was detected under control and drought stress I, but a positive correlation under drought stress II. The results suggested that there were different water-use strategies among the populations, the southeastern population with lower WUE may employ a prodigal water-use strategy, whereas the northwestern and central populations with higher WUE may employ a conservative water-use strategy. This knowledge may be useful as criteria for genotype selection within a breeding program for this species. This revised version was published online in June 2006 with corrections to the Cover Date.     

��13C signature of tree rings and radial increment of Fagus sylvatica trees as dependent on tree neighborhood and climate

We conducted dendroecological analyses in 80-year-long tree ring chronologies to detect neighborhood effects (competition intensity, species identity) on the δ¹³C signature of tree rings and radial stem increment of Fagus sylvatica trees growing either in monospecific or mixed patches of a temperate forest. We hypothesized that tree ring δ¹³C is a more sensitive indicator of neighborhood effects and the impact of climate variability on growth than is ring width. We found a closer correlation of summer precipitation to δ¹³C than to ring width. While the ring width showed a decline over the test period (1926–2005), the mean curve of δ¹³C increased until the mid of the 1970s, remained high until about 1990, and markedly decreased thereafter. Possible explanations related to ontogeny and environmental change (‘age effect’ due to canopy closure; elevated atmospheric SO₂ concentrations in the 1960s–1980s) are discussed. Beech target trees surrounded by many allospecific trees had a significantly lower mean δ¹³C in the period 1926–1975 than beech with predominantly or exclusively conspecific neighborhood, possibly indicating a more favorable water supply of beech in diverse stands. Contrary to expectation, trees subject to more intense competition by neighboring trees (measured by Hegyi’s competition index) had lower δ¹³C values in their tree rings, which is thought to reflect denser canopies being linked to increased shading. We conclude that tree ring δ¹³C time series represent combined archives of climate variability, stand history and neighborhood effects on tree physiology and growth that may add valuable information to that obtained from conventional tree ring analysis.     

Photosynthesis,water-use efficiency and δ<Superscript>13</Superscript>C of five cowpea genotypes grown in mixed culture and at different densities with sorghum

A field experiment involving two planting densities (83,333 and 166,666 plants per ha), two cropping systems (monoculture and mixed culture) and five cowpea [Vigna unguiculata L. (Walp.)] genotypes was conducted at Nietvoorbij (33°54S, 18°14E), Stellenbosch, South Africa, to select cowpea material with superior growth and water-use efficiency (WUE). The results showed significantly higher photosynthetic rates, stomatal conductance and transpiration in leaves of plants at low density and in monoculture due to greater chlorophyll (Chl) levels relative to those at high density and in mixed culture. As a result, C concentration in leaves and the amount of C, P, K, Ca, Mg, Fe, Cu, Zn, Mn, and B accumulated in shoots at low density and under monoculture were also much higher. Even though no marked differences in photosynthetic rates were found between and among the five cowpea genotypes, leaf C concentration and shoot C, P, K, Ca, Mg, Fe, Cu, Zn, Mn, and B contents differed considerably, with Sanzie exhibiting the highest C concentration and C, P, K, Ca, Mg, Fe, Cu, Zn, Mn, and B contents in shoots, followed by Bensogla and Omondaw, while ITH98-46 and TVu1509 had the lowest shoot concentration and contents of C, P, K, Ca, Mg, Fe, Cu, Zn, Mn, and B. WUE (calculated as photosynthate produced per unit water molecule transpired) was significantly greater in plants at low density and monoculture relative to those at high density and in mixed culture. Isotopic analysis revealed significant differences in δ¹³C values of sorghum [Sorghum bicolor L. (Moench.)] and cowpea, with higher δ¹³C values being obtained for plants at low density and in monoculture relative to those at high density or in mixed culture. The five cowpea genotypes also showed significant differences in δ¹³C values, with Sanzie exhibiting the most negative value (i.e. low WUE) and ITH98-46, the least negative δ¹³C value (i.e. high WUE). Whether measured isotopically or from gas-exchange studies, sorghum (a C₄ species) exhibited much higher WUE relative to cowpea (a C₃ species). Both correlation and regression analyses revealed a positive relationship between WUE from gas-exchange studies and δ¹³C values from isotopic analysis of cowpea and sorghum shoots.     

Variation in leaf area index and stand leaf mass of European beech across gradients of soil acidity and precipitation

Leaf area index (LAI, the one-sided foliage area per unit ground surface area) is a key determinant of plant productivity which has a large influence on water and energy exchange between vegetation and the atmosphere. The variation in forest LAI across landscapes and environmental gradients and its causes are not sufficiently understood. We measured the LAI of European beech (Fagus sylvatica) by litter trapping in 23 closed, mature stands across gradients of rainfall and soil acidity or fertility. With a mean LAI of 7.4 m² m⁻² (minimum: 5.6, maximum: 9.5 m² m⁻²), beech stands maintained a comparably high leaf area index with relatively small variation along steep environmental gradients. Contrary to expectation, decreasing water availability (rainfall gradient from 1030 to 520 mm yr⁻¹) or increasing soil acidity (pH 3–7) had no significant effect on LAI. Stand leaf mass (M _l) increased slightly with soil fertility (C/N ratio, base saturation). We regressed parameters of site water availability (rainfall), soil fertility or acidity (pH, base saturation, C/N ratio, exchangeable Mg and Al content), and stand structure (stand age and stem density) against LAI and M _l in order to detect environmental controls of stand leaf area. Stand age was the most influential factor for both LAI and M _l (negative relationship). Stem density and the base saturation of the soil affected M _l significantly, but had a weak influence on LAI. We conclude that the leaf area index of beech is mainly under control of age-related physiological factors, whereas the influence of soil chemistry and rainfall is comparably low.     

Altitude trends in conifer leaf morphology and stable carbon isotope composition

The natural ratio of stable carbon isotopes (δ¹³C) was compared to leaf structural and chemical characteristics in evergreen conifers in the north-central Rockies, United States. We sought a general model that would explain variation in δ¹³C across altitudinal gradients. Because variation in δ¹³C is attributed to the shifts between supply and demand for carbon dioxide within the leaf, we measured structural and chemical variables related to supply and demand. We measured stomatal density, which is related to CO₂ supply to the chloroplasts, and leaf nitrogen content, which is related to CO₂ demand. Leaf mass per area was measured as an intermediate between supply and demand. Models were tested on four evergreen conifers: Pseudotsuga menziesii, Abies lasiocarpa, Picea engelmannii, and Pinus contorta, which were sampled across 1800 m of altitude. We found significant variation among species in the rate of δ¹³C increase with altitude, ranging from 0.91‰ km^–1 for A. lasiocarpa to 2.68‰ km^–1 for Pinus contorta. Leaf structure and chemistry also varied with altitude: stomatal density decreased, leaf mass per area increased, but leaf nitrogen content (per unit area) was constant. The regressions on altitude were particularly robust in Pinus contorta. Variables were derived to describe the balance between supply and demand; these variables were stomata per gram of nitrogen and stomata per gram of leaf mass. Both derived variables should be positively related to internal CO₂ supply and thus negatively related to δ¹³C. As expected, both derived variables were negatively correlated with δ¹³C. In fact, the regression on stomatal density per gram was the best fit in the study (r ²=0.72, P<0.0001); however, the relationships were species specific. The only general relationship observed was between δ¹³C and LMA: δ¹³C (‰)=–32.972+ 0.0173×LMA (r ²=0.45, P<0.0001). We conclude that species specificity of the isotopic shift indicates that evergreen conifers demonstrate varying degrees of functional plasticity across environmental gradients, while the observed convergence of δ¹³C with LMA suggests that internal resistance may be the key to understanding inter-specific isotopic variation across altitude. Received: 1 June 1999 / Accepted: 2 November 1999     

Mechanism for the small-scale movement of carbon among estuarine habitats: organic matter transfer not crab movement

In theory, carbon is highly mobile in aquatic systems. Recent evidence from carbon stable isotopes of crabs (Parasesarma erythrodactyla and Australoplax tridentata), however, shows that in subtropical Australian waters, measurable carbon movement between adjacent mangrove and saltmarsh habitats is limited to no more than a few metres. We tested whether the pattern in crab δ¹³C values across mangrove and saltmarsh habitats was explained by crab movement, or the movement of particulate organic matter. We estimated crab movement in a mark–recapture program using an array of pitfall traps on 13 transects (a total of 65 traps) covering an area of 600 m² across the interface of these two habitats. Over a 19-day period, the majority of crabs (91% for P. erythrodactyla, 93% for A. tridentata) moved <2 m from the place of initial capture. Crab movement cannot, therefore, explain the patterns in δ¹³C values of crabs. δ¹³C values of detritus collected at 2-m intervals across the same habitat interface fitted a sigmoidal curve of a similar form to that fitting the δ¹³C values of crabs. δ¹³C values of detritus were 2–4‰ more depleted in saltmarsh (−18.5±0.6‰), and 4–7‰ more depleted in mangroves (−25.9±0.1‰) than δ¹³C values of crabs recorded previously in each habitat. Assimilation by crabs of very small detrital fragments or microphytobenthos, more enriched in ¹³C, may explain the disparity in δ¹³C values. Nevertheless, the pattern in δ¹³C values of detritus suggests that crabs obtain their carbon from up to several metres away, but without themselves foraging more then a metre or so from their burrow. Such detailed measurements of carbon movement in estuaries provide a spatially explicit understanding of the functioning of food webs in saltmarsh and mangrove habitats.