Vertical canopy gradients in δ13C correspond with leaf nitrogen content in a mixed-species conifer forest

Stable carbon isotope composition varies markedly between sun and shade leaves, with sun leaves being invariably more enriched (i.e., they contain more¹³C). Several hypotheses have emerged to explain this pattern, but controversy remains as to which mechanism is most general. We measured vertical gradients in stable carbon isotope composition (δ¹³C) in more than 200 trees of nine conifer species growing in mixed-species forests in the Northern Rocky Mountains, USA. For all species except western larch, δ¹³C decreased from top to bottom of the canopy. We found that δ¹³C was strongly correlated with nitrogen per unit leaf area (N _area), which is a measure of photosynthetic capacity. Usually weaker correlations were found between δ¹³C and leaf mass per area, nitrogen per unit leaf mass, height from the ground, or depth in the canopy, and these correlations were more variable between trees than for N _area. Gradients of δ¹³C (per meter canopy depth) were steeper in small trees than in tall trees, indicating that a recent explanation of δ¹³C gradients in terms of drought stress of upper canopy leaves is unlikely to apply in our study area. The strong relationship between N _area and δ¹³C here reported is consistent with the general finding that leaves or species with higher photosynthetic capacity tend to maintain lower CO₂ concentrations inside leaves. We conclude that photosynthetic capacity is a strong determinant of δ¹³C in vertical canopy profiles, and must be accounted for when interpreting δ¹³C values in conifer forests.     

Foliar δ13C within a temperate deciduous forest: spatial,temporal, and species sources of variation

Summary Foliar ¹³C-abundance (¹³C) was analyzed in the dominant trees of a temperate deciduous forest in east Tennessee (Walker Branch Watershed) to investigate the variation in foliar ¹³C as a function of time (within-year and between years), space (canopy height, watershed topography and habitat) and species (deciduous and coniferous taxa). Various hypotheses were tested by analyzing (i) samples collected from the field during the growing season and (ii) foliar tissues maintained in an archived collection. The ¹³C-value for leaves from the tops of trees was 2 to 3%. more positive than for leaves sampled at lower heights in the canopy. Quercus prinus leaves sampled just prior to autumn leaf fall had significantly more negative ¹³C-values than those sampled during midsummer. On the more xeric ridges, needles of Pinus spp. had more positive ¹³C-values than leaves from deciduous species. Foliar ¹³C-values differed significantly as a function of topography. Deciduous leaves from xeric sites (ridges and slopes) had more positive ¹³C-values than those from mesic (riparian and cove) environments. On the more xeric sites, foliar ¹³C was significantly more positive in 1988 (a dry year) relative to that in 1989 (a year with above-normal precipitation). In contrast, leaf ¹³C in trees from mesic valley bottoms did not differ significantly among years with disparate precipitation. Patterns in foliar ¹³C indicated a higher ratio of net CO₂ assimilation to transpiration (A/E) for trees in more xeric versus mesic habitats, and for trees in xeric habitats during years of drought versus years of normal precipitation. However, A/E (units of mmol CO₂ fixed/mol H₂O transpired) calculated on the basis of ¹³C-values for leaves from the more xeric sites was higher in a wet year (6.6±1.2) versus a dry year (3.4±0.4). This difference was attributed to higher transpiration (and therefore lower A/E) in the year with lower relative humidity and higher average daily temperature. The calculated A/E values for the forest in 1988–89, based on ¹³C, were within ±55% of estimates made over a 17 day period at this site in 1984 using micrometeorological methods.     

Is there coordination of leaf and fine root traits at local scales? A test in temperate forest swamps

Examining the coordination of leaf and fine root traits not only aids a better understanding of plant ecological strategies from a whole‐plant perspective, but also helps improve the prediction of belowground properties from aboveground traits. The relationships between leaf and fine root traits have been extensively explored at global and regional scales, but remain unclear at local scales. Here, we measured six pairs of analogous leaf and fine root traits related to resource economy and organ size for coexisting dominant and subordinate vascular plants at three successional stages of temperate forest swamps in Lingfeng National Nature Reserve in the Greater Hinggan Mountains, NE China. Leaf and fine root traits related to resource acquisition (e.g., specific leaf area [SLA], leaf N, leaf P, root water content, and root P) decreased with succession. Overall, we found strong linear relationships between leaf dry matter content (LDMC) and root water content, and between leaf and root C, N, and P concentrations, but only weak correlations were observed between leaf area and root diameter, and between SLA and specific root length (SRL). The strong relationships between LDMC and root water content and between leaf and root C, N, and P held at the early and late stages, but disappeared at the middle stage. Besides, C and P of leaves were significantly correlated with those of roots for woody plants, while strong linkages existed between LDMC and root water content and between leaf N and root N for herbaceous species. These results provided evidence for the existence of strong coordination between leaf and root traits at the local scale. Meanwhile, the leaf–root trait relationships could be modulated by successional stage and growth form, indicating the complexity of coordination of aboveground and belowground traits at the local scale.     

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.     

Characteristics of βA chromosome haplotypes in Japanese

DNA polymorphism patterns linked to the ^A-globin gene were analyzed in healthy Japanese using four different restriction endonucleases. The chromosomes with the ^A-globin gene were mapped through an evaluation of the presence of seven different restriction sites (HincII 5 to ; HindIII in ^G and ^A; HincII in, and 3 to, ₁; AvaII in ; Bam-HI 3 to ). Among 36 chromosomes analyzed, 20 chromosomes had a haplotype of [+–––––+]. Among 55 individuals examined, 7 possessed a homozygous haplotye of [+–––––+]. All Japanese with the ^AT-globin gene had a subhaplotype of [–++–+] 5 to the -globin gene. Their major haplotypes were [–++–+–+] and [–++–++–]. It was expected that the presence of the ^AT-globin gene in Japanese may be deduced from subhaplotypes 5 to the -globin gene.     

Fleshy fruit characteristics in a temperate deciduous forest of Japan: how unique are they?

This study investigated the fleshy fruit characteristics of 28 woody species in a Japanese temperate forest where large sedentary seed-dispersing mammals are present. We tested whether the findings in previous studies in temperate forests of Europe and North America are universal or not. Results have suggested that fruits of all species were eaten both by birds and mammals except for four species with larger fruits, which were eaten only by mammals. A gradient was found from a syndrome characterized by small, oily, and large-seeded fruits to a syndrome characterized by large, succulent, non-oily, and small-seeded fruits. The sizes and colors of the fruits were not conspicuously different from previous findings in Europe and North America. On the other hand, nitrogen and lipids in the fleshy part did not show seasonally increasing trends, or even seasonally decreasing trends in terms of dry weight. This result, suggesting the absence of community-level adaptation of fruit traits to migratory bird dispersers, contrasted with findings in Europe and North America. Large sedentary arboreal or tree-climbing mammals may have a greater effect on the evolution of fruit-disperser relations than opportunistic migratory birds.     

Variations of root traits in three Xizang grassland communities along a precipitation gradient北大核心CSCD

Aims Root functional traits and their variations mediate coexistence and adaptive strategy of plant species. Yet, strong environmental constraints may induce convergence of root traits among different plant species. To study the variations of root traits and clarify the diverse adaptive strategies across plant species, we sampled three alpine grasslands along a precipitation gradient in the Xizang Plateau. Methods In three grassland communities along a precipitation gradient: Nagqu, Baingoin and Nyima from east to west of Xizang Plateau, we collected 22 coexisting plant species and measured three key root traits: 1st-order root diameter, 1st-order lateral root length and root branch intensity. Important findings The main results showed that: (1) the root of plants in the alpine grassland was generally thin, and the interspecific variation was also small (22.76%); (2) the root diameter of 86% plant species was in the range from 0.073 mm to 0.094 mm. Compared with the thick-root species, thin-root species had a higher root branching intensity, but shorter lateral root length. In addition, at community-level, plants mainly increased root diameter and lateral root length, but reduced root branching intensity to adapt to the decreasing precipitation; while at species-level, the plant species exhibited diverse adaptive strategies along the precipitation gradient. © Chinese Journal of Plant Ecology.     

Presence of N-fixing neighbors increases leaf N and δ13C in Castilleja applegatei,a root hemiparasite

Plant Ecology - Parasitic plants are known for their high transpiration rates and low water use efficiency (WUE), which the N-parasitism hypothesis posits is driven by N limitation. Thus,...     

Phylogeography of Liquidambar styraciflua (Altingiaceae) in Mesoamerica: survivors of a Neogene widespread temperate forest (or cloud forest) in North America?

We investigate the genetic variation between populations of the American sweetgum (Liquidambar styraciflua), a tree species with a disjunct distribution between northeastern Texas and Mexico, by analyzing sequences of two chloroplast DNA plastid regions in Mesoamerica. Our results revealed phylogeographical structure, with private haplotypes distributed in unique environmental space at either side of the Trans‐Mexican Volcanic Belt, and a split in the absence of gene flow dating back ca. 4.2–1.4 million years ago (MYA). Species distribution modeling results fit a model of refugia along the Gulf and Atlantic coasts but the present ranges of US and Mesoamerican populations persisted disjunct during glacial/interglacial cycles. Divergence between the US and Mesoamerican (ca. 8.4–2.8 MYA) populations of L. styraciflua and asymmetrical gene flow patterns support the hypothesis of a long‐distance dispersal during the Pliocene, with fragmentation since the most recent glacial advance (120,000 years BP) according to coalescent simulations and high effective migration rates from Mesoamerica to the USA and close to zero in the opposite direction. Our findings implicate the Trans‐Mexican Volcanic Belt as a porous barrier driving genetic divergence of L. styraciflua, corresponding with environmental niche differences, during the Pliocene to Quaternary volcanic arc episode 3.6 MYA, and a Mesoamerican origin of populations in the USA.     

Inverse gradients in leaf wax δD and δ13C values along grass blades of Miscanthus sinensis: Implications for leaf wax reproduction and plant physiology

Compound specific hydrogen and carbon isotopic ratios of higher plant leaf waxes have been extensively used in paleoclimate and paleoenvironmental reconstructions. However, studies so far have focused on the comparison of leaf wax isotopic differences in bulk leaf samples between different plant species. We sampled three different varieties of tall grasses (Miscanthus sinensis) in six segments from base to tip and determined hydrogen and carbon isotopic ratios of leaf waxes, as well as hydrogen and oxygen isotopic ratios of leaf water samples. We found an increasing, base-to-tip hydrogen isotopic gradient along the grass blades that can probably be attributed to active leaf wax regeneration over the growth season. Carbon isotopic ratios, on the other hand, show opposite trends to hydrogen isotopic ratios along the grass blades, which may reflect different photosynthetic efficiencies at different blade locales.     

Variations in the foliar δ13C and C3/C4 species richness in the Japanese flora of Poaceae among climates and habitat types under human activity

For 383 Poaceae species harvested over the Japanese islands and stored as herbarium specimens along several decades, we determined C₃ and C₄ types of photosynthesis from leaf stable carbon isotope ratio (δ¹³C). Then, we sought the relationships between C₄ species richness and climatic factors or habitat types. Except for the two Panicum species (P. lanuginosum and P. scoparium) having the possibility of C₃–C₄ intermediate, 227 and 154 species were classified into C₃ and C₄. The C₄ species richness increased from northern to southern islands in Japan, positively correlated with mean annual air temperature. Greater C₄ species richness in the seashore habitats, and smaller C₄ species richness in the shaded, wet and highland habitats would be related to the photosynthetic responses to local environmental factors such as irradiance level and temperature regime. No difference of leaf δ-value of C₃ Poaceae was obtained between the habitats with different soil water availability, suggesting the less importance of soil water availability on leaf water-use efficiency in C₃ Poaceae species in Japan having humid climate. Additionally, possible effects of human activity around the harvested time or site on leaf δ-value were estimated, because the habitat includes the sites with high human activity. Leaf δ-value was decreased with sampling year, and it was higher in the densely inhabited district for both C₃ and C₄. They are probably due to a historical decrease in the atmospheric δ-value via increasing human activity, and high gas emission at the districts of high human density.     

Fire modulates the effects of introduced ungulates on plant–insect interactions in a Patagonian temperate forest

Disturbances like biological invasions and fire may affect in unexpected ways plant-animal interactions. In northwestern Patagonia, introduced ungulates (cattle, horses and deers) are widespread and very common occupying more than 50% of forests and shrublands, widely affecting these habitats. In addition, fire play a major role in creating landscape patterns in this region. We evaluated whether fire modify the impacts of introduced ungulates on plant-animal interactions. In a mature forest (unburnt) and in an early post-fire area (burnt) we used structural equation modeling (SEM) to analyzed the impacts of introduced ungulates on insect herbivory, pollination and pre-dispersal seed predation on Berberis darwinii, one of the most common understory shrub of temperate forests. We found that the effects of cattle on pollination and fruit set depended on the habitat condition (i.e. unburnt or burnt). Introduced ungulates in unburnt forest decreased fruit set through a reduction on pollinator visits. Conversely, introduced ungulates in burnt forest increased pollinator visits and flower production without affecting fruit set. On the other hand, damage patterns (herbivory and fruit/seed predation) were unaffected by cattle in both forests types. Either, low browsing pressure or induction of plant defences may explain our results. This study illustrates how modifications on biotic and abiotic conditions produced by fire may affect in complex ways the effect of introduced ungulates on plant-animal interactions.     

Litter in a first–order stream of a temperate deciduous forest (Margaraça Forest, central Portugal)

To evaluate the importance and fate of organic matter inputs in forested streams, we determined the litterfall inputs and the benthic coarse particulate organic matter (CPOM) in one headwater stream flowing through a mixed deciduous forest, during one year. Both vertical traps and the stream bottom were sampled monthly. The material collected was sorted into four main categories: leaves, fruits and flowers, twigs and debris. Litter production was 715 g m−2 y−1 and seasonal, with 73% of the annual total during October–December (autumn). Leaves comprised the largest litter component. Benthic organic matter was 1880 g m−2 y−1, and was also seasonal. Highest accumulation was attained in spring, and twigs and branches comprised the major component. This revised version was published online in July 2006 with corrections to the Cover Date.     

Variation of δ<Superscript>13</Superscript>C of wood and foliage with canopy height differs between evergreen and deciduous species in a temperate forest

We investigated post-photosynthetic fractionation and carbon transfer mechanisms of different plant functional types growing under the same climatic conditions in North-eastern China. The variations in δ¹³C of trunk and branches were compared with leaf δ¹³C at different canopy heights of Pinus koraiensis (evergreen coniferous species), Larix gmelinii (deciduous coniferous species) and Quercus mongolica (deciduous broad-leaved species). Results showed that δ¹³C of leaves increased (became more enriched) with increasing canopy height for both coniferous species (P. koraiensis, L. gmelinii) but not for Q. mongolica (a deciduous broad-leaved species). δ¹³C of both trunk and branches also increased with sampling height for the evergreen conifer P. koraiensis but did not significantly vary for either of the deciduous species (L. gmelinii or Q. mongolica), except a significant increase in branch δ¹³C for L. gmelinii. Similarly, δ¹³C of trunk and branches were strongly correlated with leaf δ¹³C only in the evergreen conifer, P. koraiensis. ¹³C was consistently more enriched in trunk, branches, and roots compared to leaves in all three species. Our findings suggest that, even under the same climatic conditions, different plant functional types may exhibit different carbon transfer mechanisms. This is contrary to the previous hypothesis that different carbon transfer mechanisms operate in forests of different climatic zones, especially in tropical and temperate forests. Particularly, the differences occur predominantly between evergreen and deciduous trees rather than between coniferous and broad-leaved trees. The significant difference in δ¹³C between leaves and wood tissues confirms a previous post-photosynthetic isotope fractionation in temperate forests.     

Abundance of climbing plants in a southern temperate rain forest: host tree characteristics or light availability?

Question: In a southern temperate rain forest, we addressed three questions: (1) Does the abundance of climbing plants increase with light availability? (2) Do host tree species differ in their susceptibility to vine infestation? (3) How does the relationship between host tree trunk diameter and relative abundance of vines vary with their climbing mechanism? Location: Two sites in the temperate evergreen rain forest of southern Chile: Puyehue (40°39′S, 72°09′W; 350 m a.s.l.) and Pastahue (42°22′S, 73°49′W; 285 m a.s.l.). Methods: We sampled vines in 60 25‐m² plots, with 20 plots in each of three light environments: mature forest, forest edges and canopy gaps. In each plot, for every tree ≥1.50‐m tall of any diameter we counted and identified all climbing plant individuals at a height of 1.30 m. We also counted, measured (trunk diameter at 1.30 m) and identified all these trees, and determined prevalence of vine infestation for each tree species. Results: Light availability in forest plots did not affect vine abundance when the number and size of host trees was taken into account. Overall, vine abundance increased with host tree trunk diameter. Tree species did not differ in the prevalence of vine infestation. The relative abundance of stem twiners and adhesive climbers decreased and increased with trunk diameter, respectively. The densities of stem twiners and adhesive climbers were negatively correlated across the forest. Conclusion: We provide further evidence that the pattern of vine abundance is independent of light availability in southern temperate rain forests, in contrast to results commonly reported for tropical rain forests. We also show that support suitability across the forest varies with the mechanism by which vines climb, probably due in part to biomechanical constraints and in part to vine interspecific competition, a virtually unexplored ecological factor.     

What factors potentially influence the ability of phylogenetic distance to predict trait dispersion in a temperate forest?

Although phylogenetic‐based approaches have been frequently used to infer ecological processes, they have been increasingly criticized in recent years. To date, the factors that affect phylogenetic signals and further the ability of phylogenetic distance to predict trait dispersion have been assumed but not empirically tested. Therefore, we investigate which factors potentially influence the ability of phylogenetic distance to predict trait dispersion. We quantified the phylogenetic and trait dispersions across size classes and spatial scales in a 9‐ha old‐growth temperate forest dynamics plot in northeastern China. Phylogenetic signals at the community level were generally lower than those at the species pool level, and phylogenetically clustered communities showed lower phylogenetic signals than did overdispersed communities. This pattern might explain the other three findings of our study. First, phylogenetically overdispersed communities performed better at predicting trait dispersion than did clustered communities. Second, the mean pairwise distance (MPD)‐based metric exhibited a stronger correlation with trait dispersion than did the mean nearest taxon distance (MNTD)‐based metric. Finally, the MNTD‐based metric showed that the prediction accuracy for trait dispersion decreased with increasing spatial scales, whereas its effects were weak on the MPD‐based metric. In addition, phylogeny could not determine the dispersions of all functional axes but was able to predict certain traits depending on whether they were evolutionarily conserved. These results were conserved when we removed the effects of space and environment. Our findings highlighted that using phylogenetic distance as a proxy of trait similarity might work in a temperate forest depending on the species in local communities sampled from total pool as well as the traits measured. Utilizing these rules, we should rethink the conclusions of previous studies that were based on phylogenetic‐based approaches.     

Effect of soil moisture on leaf ecophysiology of <Emphasis Type="Italic">Parasenecio yatabei</Emphasis>, a summer-green herb in a cool–temperate forest understory in Japan

Leaf physiological and gas-exchange traits of a summer-green herbaceous perennial, Parasenecio yatabei, growing along a stream were examined in relation to leaf age. In its vegetative phase, the aerial part of this plant consists of only one leaf and provides an ideal system for the study of leaf longevity. Volumetric soil water content (SWC) decreased with increasing distance from the stream, whereas relative light intensity was nearly constant. The light-saturated net CO₂ assimilation rate (A _sat) and leaf stomatal conductance (gs) were approximately 1.5-fold and 1.4-fold higher, respectively, in the lower slope near the mountain stream than in the upper slope far from the mountain stream. The lifespan of aerial parts of vegetative plants significantly increased with decreasing SWC. The leaf mass-based nitrogen content of the leaves (N _mass) was almost constant (ca. 2.2%); however, the maximum carboxylation rate by ribulose-1,5-biphosphate carboxylase/oxygenase (rubisco) (V _cmax) and photosynthetic nitrogen use efficiency (PNUE, A _sat/N _area) decreased more slowly in the upper slope than in the lower slope. The higher leaf photosynthetic activity of P. yatabei plants growing lower on the slope leads to a decrease in V _cmax and PNUE in the early growing season, and to a shorter leaf lifespan.     

What is the optimal number of leaves when measuring leaf area of tree species in a forest community?

植物形态性状叶面积简单易测, 能够反映植物对环境的适应与响应, 指示生态系统的功能与过程。在野外测定叶面积时, 叶片取样数量往往采用约定俗成的10-20片, 但到底采集多少叶片才是最优和最具代表性, 却少有探究。该研究以浙江金华山常绿落叶阔叶混交林的优势树种木荷(Schima superba)与枫香树(Liquidambar formosana)为研究对象, 通过对5个胸径等级植株和每个植株6个方位开展大批量叶片取样(>2 500个), 分析两个树种的叶面积变异特征, 探讨叶片取样数量为多少才能最代表该森林类型的叶片大小性状规律。结果表明, 常绿乔木木荷平均叶面积与变幅均小于落叶乔木枫香树。木荷叶面积与胸径无显著相关性, 而枫香树叶面积与胸径有较显著相关性, 但两个树种均在中胸径等级(15-20 cm)差异不显著; 两个树种的叶面积与采样方位无显著相关性, 但在东、西和底部的差异不显著。因此, 综合考虑代表性与野外可操作性, 叶片采集首选中胸径成树的底部叶片。随机抽样统计可知, 树木叶面积测定的最适叶片采集数量因物种而异, 木荷的最适叶片采集数量为40, 而枫香树最少为170片。因此, 在叶面积测定时, 叶片采集的数量应该不能只局限在10-20片, 在人力、物力和时间等条件允许的情况下, 应该尽可能多地测定较多叶片的叶面积。     

The impact of forest management on litter-dwelling invertebrates: a subtropical–temperate contrast

Land use intensification in forests is a main driver of global biodiversity loss. Although historical state of land use differs between subtropical and temperate zones, gradients of land-use intensities similarly range from unmanaged to very intensively managed forests. Irrespective of similar land use forces in both climate zones, comparative studies on land use effects are still rare. Such studies are, however, promising in discovering more general impacts and geographical specifics of land use intensification. We studied litter-dwelling invertebrates along a gradient of increasing land use intensity in subtropical forests in Southern Brazil and temperate forests in Central Europe using similar sampling designs. Effects of land use intensity on the entire community were analyzed on the level of orders and feeding guilds. In both climate zones a similar number of individuals were caught when standardizes to 100 pitfall trap days, but taxa richness was higher in the subtropics. Moreover, community composition differed between both climate zones. In both regions, land use intensity did not affect taxa richness, but invertebrate abundance was affected in opposite ways; while increasing land use intensity resulted in a decrease of invertebrate abundance in the subtropics, an increase was observed in the temperate zone and this was mostly consistent regarding different feeding guilds. Management practices should take into account that the effect of land use intensity on biodiversity can differ drastically among climatic regions.