Research article: small-scale spatial variation of δ13C and δ15N isotopes in Antarctic carbon sources and consumers

Regional food web studies that fail to account for small-scale isotopic variability can lead to a mismatch between an organism’s inferred and true trophic position. Misinterpretation of trophic status may result, substantially limiting spatial and temporal comparability of food web studies. We sampled several carbon sources and consumers in a nested design to assess the variability of food web members across small spatial scales (100 s of m to several km) in regions around the Windmill Islands and Vestfold Hills in East Antarctica. For carbon sources, δ¹³C in sea ice POM was particularly variable between locations (km apart) and between sites (100 s of m apart) with replicate samples varying by up to 16‰. Macroalgae δ¹³C was less variable (replicate samples ranging up to 6.9‰ for the red alga Iridaea cordata), yet still differed between locations. Sediment POM and pelagic POM were the least variable, displaying minimal differences between locations or sites for δ¹³C and δ¹⁵N. Three out of eight consumers were significantly different between locations for δ¹³C, and five out of eight for δ¹⁵N, with the fish Trematomus bernacchii the most variable for both δ¹³C and δ¹⁵N. At smaller scales, the amphipod Paramorea walkeri showed significant variation between sites in δ¹³C but not in δ¹⁵N. We attribute small-scale variability to the dynamic physical environment for carbon sources in coastal systems and a close coupling of diet to habitat for consumers. We highlight the need to account for small-scale spatial variation in sampling designs for regional food web studies.     

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.     

Species-specific differences in temporal and spatial variation in δ13C of plant carbon pools and dark-respired CO2 under changing environmental conditions

Stable carbon isotope signatures are often used as tracers for environmentally driven changes in photosynthetic δ¹³C discrimination. However, carbon isotope signatures downstream from carboxylation by Rubisco are altered within metabolic pathways, transport and respiratory processes, leading to differences in δ¹³C between carbon pools along the plant axis and in respired CO₂. Little is known about the within-plant variation in δ¹³C under different environmental conditions or between species. We analyzed spatial, diurnal, and environmental variations in δ¹³C of water soluble organic matter (δ¹³C_WSOM) of leaves, phloem and roots, as well as dark-respired δ¹³CO₂ (δ¹³C_res) in leaves and roots. We selected distinct light environments (forest understory and an open area), seasons (Mediterranean spring and summer drought) and three functionally distinct understory species (two native shrubs—Halimium halimifolium and Rosmarinus officinalis—and a woody invader—Acacia longifolia). Spatial patterns in δ¹³C_WSOM along the plant vertical axis and between respired δ¹³CO₂ and its putative substrate were clearly species specific and the most δ¹³C-enriched and depleted values were found in δ¹³C of leaf dark-respired CO₂ and phloem sugars, ~?15 and ~?33 ‰, respectively. Comparisons between study sites and seasons revealed that spatial and diurnal patterns were influenced by environmental conditions. Within a species, phloem δ¹³C_WSOM and δ¹³C_res varied by up to 4 ‰ between seasons and sites. Thus, careful characterization of the magnitude and environmental dependence of apparent post-carboxylation fractionation is needed when using δ¹³C signatures to trace changes in photosynthetic discrimination.     

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.     

The environmental water of the middle Eocene Arctic: Evidence from δD, δ18O and δ13C within specific compounds

The extensive vertical exposure (> 150 m) of terrestrial sediments on Axel Heiberg Island, which contain thick fossiliferous lignites, presents an exceptional opportunity to follow the establishment and re-establishment of Arctic Metasequoia forests during the middle Eocene. We compared δD values in n-alkanes of chain length 23, 25, 27 and 29 with δ¹⁸O values in phenylglucosazone (P-G) derived from α-cellulose; we also analyzed %-abundance of ferns, gymnosperms and angiosperms using pollen and spores isolated from each lignite. Our results showed that forest composition was altered upon uplift, as gymnosperms became more abundant within the relatively well-drained upland sediments. This was also reflected in the small (1‰), but significant, increase in the δ¹³C value of TOM from lowland to upland environments. However, neither the δD values of n-alkanes nor the δ¹⁸O in P-G were statistically different in the upland sediments, as compared to the lowland sediments; from this we inferred that the oxygen isotope signature of environmental water available to the forests for plant growth was relatively uniform throughout the time of the fossil forests. The δD value of environmental water implied by both n-alkanes and P-G ranged from ? 168 to ? 131% and was considerably enriched compared to all environmental water samples available from the modern Arctic region (< ? 180%). In addition to indicating a warmer Eocene Arctic, subject to meteoric transport patterns different from today's, these results argue against the presence of an Eocene polar ice cap.     

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.     

Variations in leaf β13C along a vertical profile of irradiance in a temperate Japanese forest

The vertical profile of stable carbon isotope ratios (δ¹³C) of leaves was analyzed for 13 tree species in a cool-temperate deciduous forest in Japan. The vertical distribution of long-term averaged δ¹³C in atmospheric CO₂ (δ_a) was estimated from δ¹³C of dry matter from NADP-malic enzyme type C₄ plant (Zea mays L. var. saccharata Sturt.) grown at a tower in the forest for 32␣days, assuming constant Δ value (3.3‰) in Z. mays against height. The δ_a value obtained from δ¹³C in Z.␣mays was lowest at the forest floor (−9.30 ± 0.03‰), increased with height, and was almost constant above 10␣m (−7.14 ± 0.14‰). Then leaf Δ values for the tree species were calculated from tree leaf δ¹³ C andδ_a. Mean leaf Δ values for the three tall deciduous species (Fraxinus mandshurica, Ulmus davidiana, and Alnus hirsuta) were significantly different among three height levels in the forest: 23.1 ± 0.7‰ at the forest floor (understory), 21.4 ± 0.5‰ in lower canopy, and 20.5 ± 0.3‰ in upper canopy. The true difference in tree leaf Δ among the forest height levels might be even greater, because Δ in Z. mays probably increased with shading by up to ∼‰. The difference in tree leaf Δ among the forest height levels would be mainly due to decreasing intercellular CO₂ (C _i) with the increase in irradiance. Potential assimilation rate for the three tree species probably increased with height, since leaf nitrogen content on an area basis for these species also increased with height. However, the increase in stomatal conductance for these tree species would fail to meet the increase in potential assimilation rate, which might lead to increasing the degree of stomatal limitation in photosynthesis with height. Received: 30 September 1995 / Accepted: 25 October 1996     

Stipa tenacissima Does not Affect the Foliar δ13C and δ15N of Introduced Shrub Seedlings in a Mediterranean Semi-arid Steppe

Recent studies have shown that the tussock grass Stipa tenacissima L. facilitates the establishment of late-successional shrubs, in what constitutes the first documented case of facilitation of woody plants by grasses. With the aim of increasing our knowledge of this interaction, in the present study we investigated the effects of S. tenacissima on the foliar δ13C, δ15N, nitrogen concentration, and carbon: nitrogen ratio of introduced seedlings of Pistacia lentiscus L., Quercus coccifera L., and Medicago arborea L. in a semi-arid Mediterranean steppe. Six months after planting, the values of δ13C ranged between -26.9‰ and -29.6‰,whereas those of δ15N ranged between -1.9‰ and 2.7‰. The foliar C: N ratio ranged between 10.7 and 53.5,and the nitrogen concentration ranged between 1.0% and 4.4%. We found no significant effect of the microsite provided by S. tenacissima on these variables in any of the species evaluated. The values of δ13C were negatively correlated with predawn water potentials in M. arborea and were positively correlated with relative growth rate in Q. coccifera. The values of δ15N were positively correlated with the biomass allocation to roots in the latter species. The present results suggest that the modification of environmental conditions in the are surrounding S. tenacissima was not strong enough to modify the foliar isotopic and nitrogen concentration of shrubs during the early stages after planting.     

The vascular epiphytes of a lowland forest in Panama—species composition and spatial structure

We conducted a comprehensive census of the vascular epiphytes in a lowland forest in Panama by means of a canopy crane. In 0.4 ha of ca. 40-m tall forest, 103 species of vascular epiphytes with 13,099 individuals were found. The orchids were the most important family both in species and individual numbers, accounting alone for >40% of all species and >50% of all individuals. There was a clear vertical segregation of species with a concentration at intermediate heights: more than 50% of all individuals were found between 15 m and 25 m above ground. Tree species identity, tree size and the position of a tree in the forest (“space”) all influenced species composition. However, none of the two environmental variables nor space alone explained more than 10% of the total variation in epiphyte assemblages in several canonical correspondence analyses. By far the largest proportion of the observed variation remained unexplained and is arguably due to mere chance. In the future, our results will be used as a baseline data-set for the direct observation of the long-term dynamics in a diverse epiphyte community.     

Foliar δ13C Within Temperate Deciduous Forest: Its Spatial Change and Interspecies Variation

The stable carbon isotope ratios (δ13C) of dominant tree and shrub species in typical temperate broad-leaved forests in Dongling Mountain, northwestern part of Beijing, China, were studied. The results indicated that the values of foliar 8 13C differed among the species tested, in the following order of Prunus armeniaca var. ansu Maxim. > Fraxinus rhychophylla Hce. ≈ Virex negundo var. heterophylla (Franch.) ReM. ≈ Quercus liaotungensis Koiz. ≈ Syringa pekinensis Rupr. > Juglans mandshurica Maxim. The δ13C values were also affected by the maturity of leaves. The newly-developed leaves had higher δ13C values than the mature ones, especially in Juglans mandshurica and Quercus liaotungensis. The conditions of habitats, particularly soil depth and moisture content, also had pronounced effect on foliar δ13C. The foliar δ13C values of plants grown in dry habitats were higher than those in moist ones. They were also affected by the position of leaves in the canopy: leaves at upper canopy showed higher δ13C values than the lower ones. It is suggested that stable isotope technique may be effective for the study of ecophysiology of plants in the warm temperate regions of China.     

Links between plant species’ spatial and temporal responses to a warming climate

To generate realistic projections of species’ responses to climate change, we need to understand the factors that limit their ability to respond. Although climatic niche conservatism, the maintenance of a species’s climatic niche over time, is a critical assumption in niche-based species distribution models, little is known about how universal it is and how it operates. In particular, few studies have tested the role of climatic niche conservatism via phenological changes in explaining the reported wide variance in the extent of range shifts among species. Using historical records of the phenology and spatial distribution of British plants under a warming climate, we revealed that: (i) perennial species, as well as those with weaker or lagged phenological responses to temperature, experienced a greater increase in temperature during flowering (i.e. failed to maintain climatic niche via phenological changes); (ii) species that failed to maintain climatic niche via phenological changes showed greater northward range shifts; and (iii) there was a complementary relationship between the levels of climatic niche conservatism via phenological changes and range shifts. These results indicate that even species with high climatic niche conservatism might not show range shifts as instead they track warming temperatures during flowering by advancing their phenology.     

Short-term variations in δ<Superscript>13</Superscript>C of ecosystem respiration reveals link between assimilation and respiration in a deciduous forest

We present a comprehensive dataset of hourly, daily, and monthly measurements of carbon isotope measurements of CO₂ in canopy air from a temperate deciduous forest with the aim to identify the relevance of short-term variations in the isotopic signature of ecosystem respiration (¹³C_R) and to understand its underlying physiological processes. We show that during daytime low vertical mixing inside the canopy can lead to decoupling of the air in the lower and upper canopy layer resulting in large spatial variation of ¹³C in CO₂ of canopy air. Intercept of Keeling Plots also showed large temporal variation (3.8) over the course of the day demonstrating that intercepts can differ between day and night and suggesting that choosing the right time for sampling is essential to capture the isotopic signature of ecosystem respiration (¹³C_R). ¹³C_R as obtained from night-time measurements showed large variation of up to 2.65 on a day-to-day basis, which was similar to the observed variation of ¹³C_R over the seasonal cycle (3.08). This highlights the importance of short-term physiological processes within ecosystems for the isotopic composition of CO₂ in the atmosphere, not reflected by bulk plant and soil organic samples. At daily and monthly time scales, ¹³C_R increased with increasing ratio of vapour pressure deficit to photosynthetically active radiation, measured 4–5 days before. This suggests that ecosystem respiration was isotopically linked to assimilation. Furthermore, assimilates recently fixed in the canopy seem to form a labile carbon pool with a short mean residence time that is respired back to the atmosphere after 4–5 days.     

Temporal variation of δ<Superscript>13</Superscript>C of larch leaves from a montane boreal forest in Mongolia

This paper reports the temporal variation (2002–2004) in foliar δ¹³C values, which are indicative of long-term integrated photosynthetic and water use characteristics, of Siberian larch (Larix sibirica Ledeb.) trees in a montane forest at Mongonmorit, NE Mongolia. At the stand, the δ¹³C value for understory shaded leaves was more negative by 2‰ on average than that for sunlit leaves sampled concurrently from open and sun-exposed environments in a forest gap. The δ¹³C value of both sunlit and shaded leaves showed pronounced intra- but relatively small inter-seasonal variations. The δ¹³C value was more positive for juvenile than mature leaves. We conjecture that juvenile leaves may derive carbon reserves in woody tissues (e.g., stems). Regardless of leaf habitats, the δ¹³C value was also affected by insect herbivores occurred in mid summer of 2003, being more negative in newly emerging leaves from the twigs after defoliation than in non-defoliated mature leaves. This pattern seems to contrast with that for the juvenile leaves in the early growing season. We surmise that the newly emerging leaves used stored organic carbon that was depleted due to fractionation during remobilization and translocation for leaf regrowth. There was also intra- and inter-seasonal variation in the foliar N concentrations and C:N ratios. A good positive (negative) correlation between the foliar δ¹³C values and N concentrations (C:N ratios) was also observed for both sunlit and shaded leaves, suggesting that the relationship between water and nitrogen use is a crucial factor affecting the plant carbon–water relationship in this mid latitude forest with a cold semiarid climate. Our isotopic data demonstrate that the larches in NE Mongolia exhibits relatively higher water use efficiency with a distinct within-season variability.     

The forest–alpine ecotone: a multi-scale approach to spatial and temporal dynamics of treeline change at Niwot Ridge

Background: Current understanding of treeline or forest-alpine ecotone (FAE) dynamics does not fully explain past and present FAE patterns and their underlying processes, nor allow prediction of their response to climate change.

Aims: We address the overarching hypothesis that the FAE is a mosaic of distinct landscape units of vegetation and landforms that result in differential responses to climate change. We focus on climate-related, landscape and vegetation characteristics, but also consider the effect of landscape heterogeneity on biogeochemistry and overall resilience of the FAE to climate change.

Results: There are three distinct FAE land units at Niwot Ridge, generated by different interactions of climate with vegetation, landforms and topography. Within these FAEs, a process of self-organisation takes place from organism to patch to landscape scales, and is modulated by positive and negative feedback loops along an elevation gradient. The underlying controls cannot be attributed solely to temperature, but to a combination of interactions along a physical/biotic gradient.

Conclusions: FAE dynamics result from interactions among mechanisms and processes at the microsite, patch and landscape scales: (1) tree persistence; (2) forest patch establishment; (3) drivers of patch forest configurations and (4) resilience, increasing along a gradient of biotic control.     

Foliar chemistry and tree ring δ13C of Pinus densiflora in relation to tree growth along a soil pH gradient

Background and aims

Soil acidification is known to be one of the constraints of tree growth; however, it is unclear how it affects tree growth at photosynthesis level (i.e., through affecting stomatal conductance vs. carboxylation rate) during the growth of trees. This paper studied the effects of soil acidification on Pinus densiflora foliar chemistry and tree ring C isotope ratio (¹³C/¹²C, expressed as δ¹³C) and their relationship with tree growth.

Methods

Tree growth (diameter, annual growth ring area, and root biomass), soil chemistry (pH, mineral N, and exchangeable Ca and Al), foliage chemistry (N, Ca/Al, and δ¹³C), and tree ring δ¹³C in P. densiflora stands along a soil pH gradient (from 4.38 to 4.83, n?=?9) in southern Korea were investigated.

Results

Overall, trees with relatively poor growth under more acidic soil conditions (low pH and Ca/Al) had lower values of foliar N concentration and δ¹³C and tree ring δ¹³C, suggesting that restricted N uptake under more acidic soil conditions caused N limitation for photosynthesis, leading to poor tree growth. In addition, relationships between mean annual area increment and carbon isotope discrimination of tree rings at five-yr intervals from 1968 to 2007 revealed that the impact of soil acidification on tree growth became severer during the last 15 yrs as negative correlations between them became significant after 1993.

Conclusions

Reduced N uptake under acidic soil conditions resulted in lower radial growth of P. densiflora via non-stomatal limitation of photosynthesis.     

Can spatial variation and inter-annual variation in precipitation explain the seed density and species richness of the germinable soil seed bank in a tropical dry forest in north-eastern Brazil?

The interaction between microhabitat and inter-annual variation in precipitation has an important role on the dynamics of the seed bank and can play a crucial role in survival and maintenance of plant populations in semi-arid environments. We hypothesized that the type of microhabitat and the inter-annual variability in precipitation can explain the richness and density of the seed bank in a semi-arid region in Brazil. The study was conducted in an area of tropical dry forest with shrub-tree physiognomy, locally called caatinga. We collected 35 soil samples in three distinct microhabitats, at the end of rainy and dry seasons, respectively, over three years, totalling 630 samples. The seed bank (richness and seed density) were determined by seedling emergence method. Over the three years, 79 species emerged from the seed bank, 64, 45 and 42 in riparian, non-riparian and rocky microhabitats, respectively. We recorded differences in species richness and average density between microhabitats and between years, with significant statistical interaction between them. Inter-annual precipitation explained 48% and 5% of the variation in richness and seed density, respectively. Spatial variation explained 7% of the species richness and 31% of the density. Our results show that the interaction between spatial variation and precipitation has an important role on the spatial and temporal heterogeneity of the richness and density of seed banks in dry environments.     

Does spatial distribution of tree size account for spatial variation in soil respiration in a tropical forest?

We explored the relationship between soil processes, estimated through soil respiration (R _soil ), and the spatial variation in forest structure, assessed through the distribution of tree size, in order to understand the determinism of spatial variations in R _soil in a tropical forest. The influence of tree size was examined using an index (I _c ) calculated for each tree as a function of (1) the trunk cross section area and (2) the distance from the measurement point. We investigated the relationships between I _c and litterfall, root mass and R _soil , respectively. Strong significant relationships were found between I _c and both litterfall and root mass. R _soil showed a large range of variations over the 1-ha experimental plot, from 1.5 to 12.6 g_C m^?2 d^?1. The best relationship between I _c and R _soil only explained 17% of the spatial variation in R _soil . These results support the assumption that local spatial patterns in litter production and root mass depend on tree distribution in tropical forests. Our study also emphasizes the modest contribution of tree size distribution–which is mainly influenced by the presence of the biggest trees (among the large range size of the inventoried trees greater than 10 cm diameter at 1.30 m above ground level or at 0.5 m above the buttresses)–in explaining spatial variations in R _soil .     

Do disturbance and productivity influence evenness of seedling,sapling and adult tree species across a semi‐deciduous tropical forest landscape?

Disturbance and productivity may influence and alter community structure by affecting the partitioning of resources among species. Here, we examined how evenness in the relative abundance of growth stages (seedlings, saplings and adults) of woody species is related to measures of productivity (i.e. total diameter breast high (dbh) and tree volume) and aspects of human disturbance (i.e. number of tree stumps, area covered by charcoal making holes and trail length) in a bamboo‐deciduous forest, northeastern of Thailand. Our results using stepwise multiple regressions showed that productivity (total dbh) explained a significant part of the variation in evenness only at the adult stage where evenness decreased with productivity. We found a decreased evenness of saplings with the area covered by charcoal making holes. Evenness of seedlings was not related to productivity or human disturbance variables and other environmental variables contribute more to explain the variation in evenness at this stage. Evenness was correlated consistently between the growth stages but the strength of relationships diminishes across the stages. Our results suggest that high productivity and human disturbance may facilitate competitive dominant species, affecting evenness in woody communities.