Trophic level delineation and resource partitioning in a South African afromontane forest bird community using carbon and nitrogen stable isotopes

Southern African forests are naturally fragmented yet hold a disproportionately high number of bird species. Carbon and nitrogen stable isotopes were measured in feathers from birds captured at Woodbush (n = 27 species), a large afromontane forest in the eastern escarpment of Limpopo province, South Africa. The δ¹³C signatures of a range of forest plants were measured to categorise the food base. Most plants sampled, including two of five grass species, had δ¹³C signatures typical of a C₃ photosynthetic pathway (?29.5 ± 1.9‰). Three grass species had a C₄ signature (?12.0 ± 0.6‰). Most bird species had δ¹³C values representing a predominantly C₃‐based diet (?24.8‰ to ?20.7‰). δ¹⁵N values were as expected, with higher levels of enrichment associated with a greater proportion of dietary animal matter. The cohesive isotopic niche defining most species (n = 22), where the ranges for δ¹³C and δ¹⁵N were 2.4‰ and 3.4‰, respectively, highlight the difficulties in understanding diets of birds in a predominantly C₃‐based ecosystem using carbon and nitrogen stable isotopes. However, variation in isotopic values between and within species provides insight into possible niche width and the use of resources by different birds within a forest environment.     

Little Tracker

Little Tracker . Directed by Johan Heynes; camera, Fanie van der Merwe, Jacques Barnard, Thomas van der Merwe; sound, Laura van der Merwe; Fevertree Productions; 2000; 53 mins., color; narration in English, dialogue in Afrikaans. Distributed by ProVision Distribution Network, PO Box 49232, Hercules, 0030, South Africa. Tel: +27-12-3762608; Fax: +27-12-3762119; E-mail: provision@intekom.co.za     

Within trophic level shifts in collagen–carbonate stable carbon isotope spacing are propagated by diet and digestive physiology in large mammal herbivores

Stable carbon isotope analyses of vertebrate hard tissues such as bones, teeth, and tusks provide information about animal diets in ecological, archeological, and paleontological contexts. There is debate about how carbon isotope compositions of collagen and apatite carbonate differ in terms of their relationship to diet, and to each other. We evaluated relationships between δ¹³C_collagen and δ¹³C_carbonate among free‐ranging southern African mammals to test predictions about the influences of dietary and physiological differences between species. Whereas the slopes of δ¹³C_collagen–δ¹³C_carbonate relationships among carnivores are ≤1, herbivore δ¹³C_collagen increases with increasing dietary δ¹³C at a slower rate than does δ¹³C_carbonate, resulting in regression slopes >1. This outcome is consistent with predictions that herbivore δ¹³C_collagen is biased against low protein diet components (¹³C‐enriched C₄ grasses in these environments), and δ¹³C_carbonate is ¹³C‐enriched due to release of ¹³C‐depleted methane as a by‐product of microbial fermentation in the digestive tract. As methane emission is constrained by plant secondary metabolites in browse, the latter effect becomes more pronounced with higher levels of C₄ grass in the diet. Increases in δ¹³C_carbonate are also larger in ruminants than nonruminants. Accordingly, we show that Δ¹³C_collagen‐_carbonate spacing is not constant within herbivores, but increases by up to 5 ‰ across species with different diets and physiologies. Such large variation, often assumed to be negligible within trophic levels, clearly cannot be ignored in carbon isotope‐based diet reconstructions.     

Soil-surface carbon dioxide efflux and microbial biomass in relation to tree density 13 years after a stand replacing fire in a lodgepole pine ecosystem

The effects of fire on soil‐surface carbon dioxide (CO₂) efflux, F_S, and microbial biomass carbon, C_mic, were studied in a wildland setting by examining 13‐year‐old postfire stands of lodgepole pine differing in tree density (< 500 to > 500 000 trees ha^?1) in Yellowstone National Park (YNP). In addition, young stands were compared to mature lodgepole pine stands (～110‐year‐old) in order to estimate ecosystem recovery 13 years after a stand replacing fire. Growing season F_S increased with tree density in young stands (1.0 µmol CO₂ m^?2 s^?1 in low‐density stands, 1.8 µmol CO₂ m^?2 s^?1 in moderate‐density stands and 2.1 µmol CO₂ m^?2 s^?1 in high‐density stands) and with stand age (2.7 µmol CO₂ m^?2 s^?1 in mature stands). Microbial biomass carbon in young stands did not differ with tree density and ranged from 0.2 to 0.5 mg C g^?1 dry soil over the growing season; C_mic was significantly greater in mature stands (0.5–0.8 mg C g^?1 dry soil). Soil‐surface CO₂ efflux in young stands was correlated with biotic variables (above‐ground, below‐ground and microbial biomass), but not with abiotic variables (litter and mineral soil C and N content, bulk density and soil texture). Microbial biomass carbon was correlated with below‐ground plant biomass and not with soil carbon and nitrogen, indicating that plant activity controls not only root respiration, but C_mic pools and overall F_S rates as well. These findings support recent studies that have demonstrated the prevailing importance of plants in controlling rates of F_S and suggest that decomposition of older, recalcitrant soil C pools in this ecosystem is relatively unimportant 13 years after a stand replacing fire. Our results also indicate that realistic predictions and modeling of terrestrial C cycling must account for the variability in tree density and stand age that exists across the landscape as a result of natural disturbances.     

Interpretation of soil δ13C as an indicator of vegetation change in African savannas

Question: The relationship between carbon‐13 in soil organic matter and C₃ and C₄ plant abundance is complicated because of differential productivity, litter fall and decomposition. As a result, applying a mass balance equation to δ¹³C data from soils cannot be used to infer past C₃ and C₄ plant abundance; only the proportion of carbon derived from C₃ and C₄ plants can be estimated. In this paper, we compare δ¹³C of surface soil samples with vegetation data, in order to establish whether the ratio of C₃:C₄ plants (rather than the proportion of carbon from C₃ and C₄ plants) can be inferred from soil δ¹³C. Location: The Tsavo National Park, in southeastern Kenya. Methods: We compare vegetation data with δ¹³C of organic matter in surface soil samples and derive regression equations relating the δ¹³C of soil organic matter to C₃:C₄ plant abundance. We use these equations to interpret δ¹³C data from soil profiles in terms of changes in inferred C₃:C₄ plant ratio. We compare our method of interpretation with that derived from a mass balance approach. Results: There was a statistically significant, linear relationship between the δ¹³C of organic matter in surface soil samples and the natural logarithm of the ratio of C₃:C₄ plants in the 100m² surrounding the soil sample. Conclusions: We suggest that interpretation of δ¹³C data from organic matter in soil profiles can be improved by comparing vegetation surveys with δ¹³C of organic matter in surface soil samples. Our results suggest that past C₃ plant abundance might be under‐estimated if a mass balance approach is used.     

The evolution of ecological specialization in southern African ungulates: competition‐ or physical environmental turnover?

Using long‐term diet reconstructions spanning the past one million years, we contrast hypotheses that biotic interactions versus physical environmental changes are primary drivers of evolutionary turnover in mammals. We use stable carbon (δ¹³C) and oxygen (δ¹⁸O) isotope ratios in tooth enamel carbonate to trace herbivore niche shifts through the Late Quaternary Land Mammal Ages (LMAs) of grassland savannas in the South African interior (Cornelian‐1.0 to 0.6 Ma; Florisian‐500 to 10 ka; and Holocene/modern). Data reveal niche separation amongst closely related coeval taxa, and dispersals through time into empty niche spaces following extinctions. This suggests a primary role of competitive exclusion and niche displacement for speciation and extinctions in these early grassland environments. However, niche changes through time show a similar trend in many taxa, entailing increased δ¹³C (elevated C₄ grass consumption) from the Cornelian to the Florisian, and from the Florisian to the Holocene/modern, and elevated δ¹⁸O in Holocene/modern taxa that reflect global aridification around the terminal Pleistocene. Commonality in isotopic trends implies universal environmental forcing of ecological, and ultimately macroevolutionary, turnover. Yet some taxa shift from a mixed C₃/C₄ diet in the Florisian to a near‐pure C₃ diet today. Indeed, we find that while δ¹³C data are normally distributed for Cornelian fossils, non‐normal distributions characterize more recent time intervals. Such distributions are in line with the bimodal distribution of δ¹³C and diet in contemporary African ungulates. Thus, while environmental forcing did not, by necessity, lead to increases in C₄ intake, the results show changes from mixed to more specialized diets. We propose that this niche specialization was a function of long‐term exposure to C₄ grasslands, consistent with predictions that relatively high metabolic demands of C₄ grazing in subtropical environments forced the differentiation of herbivores into one of two highly specialized feeding niches, i.e. C₃ browsing or C₄ grazing.     

The carbon origin of arbuscular mycorrhizal fungi estimated from δ13C values of individual spores

 The origin of carbon in the spores of arbuscular mycorrhizal (AM) fungi was quantified based on their obligate symbiosis with C₃ and C₄ plants showing clearly different δ¹³C values. The δ¹³C values of individual spores of the AM fungus Gigaspora margarita were analyzed. In monoculture pots of a C₃ or a C₄ plant species, spore δ¹³C values were ca. 3.5‰ lower than those of host roots. In coculture pots of a C₃ and a C₄ plant species, spore δ¹³C values varied between those of the roots of C₃ and C₄ plants, and increased linearly from the C₃ to the proximity of the C₄ plant (P<0.01). This reflects the higher δ¹³C values in C₄ plants than in C₃ plants. Thus the carbon origin of G. margarita spores changed with growth state and combination of host plants. In the presence of fresh plant residue instead of living host plants, spore δ¹³C values did not vary with distance from the residue. This finding supports the current view that AM fungi are obligate symbionts. Accepted: 12 February 1999     

Effect of host shoot clipping on carbon and nitrogen sources for arbuscular mycorrhizal fungi

The effect of clipping of the host-plant shoot on the sources of carbon and nitrogen for the arbuscular mycorrhizal (AM) fungus Gigaspora margarita was determined by measuring ¹³C in spores and hyphae in cocultures of C₃ and C₄ plants and by differential ¹⁵N labeling. C₃ and C₄ plants, which have different δ¹³C values, were grown in the same container separated by a series of hyphal compartments. The C₃ and C₄ plants were applied with ¹⁴N- and ¹⁵N-urea, respectively. After clipping of the C₃ shoots, spore δ ¹³C gradually approached that of the C₄ roots. Hyphal δ ¹³C paralleled that of spores. Spore % ¹⁵N was similar to that of mineral N in the C₄ plant compartment. Thus C in G. margarita coming from the clipped plants decreased with time. This demonstrates that C in AM fungi comes from living plants, whilst the N in spores comes mostly from the soil. Accepted: 28 November 2000     

Carbon isotopic abundances in Mesozoic and Cenozoic fossil plants: Palaeoecological implications

Carbon isotopic abundances have been measured for more than one hundred samples of fossil plants ranging in age from middle Triassic to late Tertiary. Most of the plant fossils were identified at the specific or generic level and were selected as representing a variety of continental environments, including xeric and humid habitats. Material analysed included numerous fragments of flowers, seeds, fruits, leaves and wood, as well as a single amorphous lignite sample. The analyses performed for the plant fragments indicate relatively constant isotopic compositions during this time interval, with plant δ¹³C values ranging between -28 and -20%. These values are within the range for living terrestrial plants with C₃, photosynthesis, although values more positive than -23% are rare in C₃ plants and typically found in plants growing under environmental stress. Lower δ¹³C values might have been expected owing to the much higher CO₂, levels of the Cretaceous atmosphere that have been inferred from marine carbonates. No fossils with values indicating C₄, photosynthesis were discovered. Fossil plants from inferred mesic environments showed δ¹³C values ranging between -26.7 and -24.1%. Highest δ¹³C values in angiosperms (up to -20.1%) were measured for Late Cretaceous combretaceous flowers from Portugal. Some cheirolepidiaceous conifers from the Early Cretaceous also showed high δ¹³C values. Values measured for Pseudofrenelopsis varians and Glenrosa taxensis were -21.9%, and values of gymnosperm wood, probably of cheirolepidiaceous affinity, were -19.0%. These high values are in accordance with inferred ecological conditions for the fossil plants. They may suggest a tendency for C₄,-like photosynthesis, although the data are equivocal. Higher values (-17.3%) clearly falling outside the C₃, range were, however, obtained from a single lignite fragment of Late Cretaceous (Maastrichtian) age. The nature of this plant fragment is unknown, but the result suggests that C₄-like photosynthesis was present at least in some latest Cretaceous vegetation. A hadrosaurian dinosaur with well-preserved collagen-like organic matter from the same deposit showed δ¹³C values around-16%, which also suggests the presence of CAM or even C₄ plants in the latest Cretaceous. □Carbon isotopic abundances, δ¹³C values, dinosaurs, plants, photosynthetic pathways, Mesozoic.     

The intramolecular ¹³C-distribution in ethanol reveals the influence of the CO₂ -fixation pathway and environmental conditions on the site-specific ¹³C variation in glucose

Efforts to understand the cause of ¹²C versus ¹³C isotope fractionation in plants during photosynthesis and post‐photosynthetic metabolism are frustrated by the lack of data on the intramolecular ¹³C‐distribution in metabolites and its variation with environmental conditions. We have exploited isotopic carbon‐13 nuclear magnetic resonance (¹³C NMR) spectrometry to measure the positional isotope composition (δ¹³C_i, ‰) in ethanol samples from different origins: European wines, liquors and sugars from C₃, C₄ and crassulacean acid metabolism (CAM) plants. In C₃‐ethanol samples, the methylene group was always ¹³C‐enriched (～2‰) relative to the methyl group. In wines, this pattern was correlated with both air temperature and δ¹⁸O of wine water, indicating that water vapour deficit may be a critical defining factor. Furthermore, in C₄‐ethanol, the reverse relationship was observed (methylene‐C relatively ¹³C‐depleted), supporting the concept that photorespiration is the key metabolic process leading to the ¹³C distribution in C₃‐ethanol. By contrast, in CAM‐ethanol, the isotopic pattern was similar to but stronger than C₃‐ethanol, with a relative ¹³C‐enrichment in the methylene‐C of up to 13‰. Plausible causes of this ¹³C‐pattern are briefly discussed. As the intramolecular δ¹³C_i‐values in ethanol reflect that in source glucose, our data point out the crucial impact on the ratio of metabolic pathways sustaining glucose synthesis.     

δ<Superscript>13</Superscript>C values,photosynthetic pathways,and plant functional types for some plants from saline meadows,Northeastern China

Based on stable carbon isotope ratio (δ¹³C) measurements, photosynthetic pathway types were determined for 61 species in 54 genera and 24 families of flowering plants from the saline meadows of Northeastern China. Of these total vascular plants, 18 species in 17 genera from 6 families were found to have C₄ photosynthesis; 43 species in 38 genera from 20 families had C₃ photosynthesis. Six dicotyledonous species exhibited C₄ pathway, 12 monocotyledonous species were found with C₄ photosynthesis. The dicotyledonous C₄ species had relative greater mean δ¹³C value and less total carbon content than both monocotyledonous C₄ and C₃ species. Most dicotyledonous C₄ species were annual forbs and halophytes. Some C₄ species had been previously documented, but their δ¹³C values varied remarkably from those of the present study. Even though there are some fluctuations for the δ¹³C values of some C₄ species, δ¹³C value was still more reliable for C₃ and C₄ identification than the use of the enzyme ratio method and of low CO₂ compensation concentration.     

Fractionation of the stable isotopes of inorganic carbon by seagrasses

The δ¹³C values for seagrasses collected along the Texas Gulf Coast range from −10.9 to −11.4‰. These values are similar to the δ¹³C values of terrestrial C₄ plants, but seagrasses lack bundle sheath cells which are important in determining the δ¹³C values of C₄ plants. This work attempts to explain the reason the δ¹³C values of seagrasses resemble the δ¹³C values of C₄ plants.     

Estimation of food composition of Hodotermes mossambicus (Isoptera: Hodotermitidae) based on observations and stable carbon isotope ratios

Abstract The diet of the harvester termite Hodotermes mossambicus was investigated at two sites with distinct dietary components: C₄ grasses (δ¹³C isotope values, ?13.8‰ to ?14.0‰) and C₃ plants (δ¹³C isotope values, ?25.6‰ to ?27.1‰). By comparing observations of food items carried into the colony by the termites and carbon isotope ratios of whole termites (that determined assimilated carbon), the relative proportion of the C₃ and C₄ plant food components of the termite diet was estimated. There was agreement between the observational data and stable carbon isotopic data, with grass representing approximately 93% of the diet of H. mossambicus at two study sites (urban and rural) on the South African highveld. However, when correcting for mass of food items, that is, C₃ and C₄, carried by termites, the proportion of grass (C₄) in the diet may be underestimated.     

The Signs of Maize? A Reconsideration of What δ13C Values Say about Palaeodiet in the Andean Region

Palaeodietary isotope studies have long assumed C₄ signals in South American archaeological populations to be due to the consumption of maize (Zea mays), which in turn, underlie interpretations important social processes. We presents δ¹³C data from wild plants (n?=?89) from the south coast of Peru, which may have been significant in the diets of humans and animals in the past. A combination of these with previously published results from domesticates of the Andean region (n?=?144) brings the proportion of C₄ species likely to have contributed to the human dietary isotopic signal, whether directly or indirectly, to almost one third. This undermines the widespread assumption that maize is the only plant to contribute a C₄ signal to diets. By considering both direct and indirect routes whereby C₄ plants may have contributed to the human isotopic signal we show the need for a reassessment of how palaeodietary studies are interpreted in the Andes, and perhaps elsewhere in the Americas.     

中国区域478种C3植物叶片碳稳定性同位素组成与水分利用效率

目前, 在中国区域关于植物碳稳定性同位素组成(δ¹³C)已经有了很多的研究, 同时, δ ¹³C作为植物水分利用效率(WUE)的替代指标, 得到了越来越广泛的应用。而这些研究多集中在站点或小的区域尺度, 那么在整个中国区域尺度上, δ¹³C能否作为植物WUE的替代指标值得探讨。该文通过对文献资料的收集整理, 研究了中国区域187个采样点478种C₃植物叶片的δ¹³C, 统计分析结果表明δ¹³C的变化范围为-33.50‰- -22.00‰, 均值为-(27.10 ± 1.70)‰。在乔木、灌木和草本3种不同的生活型间, 叶片δ¹³C的差异达到极显著水平, 其中以草本的δ¹³C最高, 乔木最低, 这与在站点或小的区域尺度上的研究结果不同。对不同系统发育类型的植物而言, 种子植物的δ¹³C极显著地大于蕨类植物; 祼子植物与被子植物间的差异未达到显著水平; 单子叶植物极显著地大于双子叶植物。叶片δ¹³C值随经度的变化没有明显的规律, 但是随纬度的增加, δ ¹³C极显著地升高。随年均温度和年均降雨量的降低, 叶片δ¹³C值极显著升高。年均降雨量与δ¹³C间的这种极显著的负相关关系, 与WUE和降水量间的关系一致, 这表明在大的区域尺度上, δ¹³C可以作为植物WUE的指示指标。     

Tracing the Origin of Dietary Protein in Tropical Dry Forest Birds1

Fundamental to our understanding of the ecology of animal communities in the tropics is knowledge of the effect of seasonal changes in the abundance of food sources in consumer diets. We determined stable‐isotope composition (¹³C/¹²C and ¹⁵N/¹⁴N) in whole blood of 14 resident avian species in a tropical dry forest to quantify the origin of their assimilated protein. We used a probabilistic approach (IsoSource) to estimate the relative contribution of C₃ plants, CAM‐C₄ plants, C₃ insects, and CAM‐C₄ insects during the dry and rainy seasons. IsoSource iteratively creates each possible combination of source contribution and produces a distribution of all feasible combinations that adequately predict the observed isotopic signature of the consumer. Granivore–frugivores and granivore–frugivore–insectivores were modeled as predominantly dependent upon plants whereas insectivorous birds were modeled to derive protein almost exclusively from insects. Between these extremes there were several species using mixed diets such as insectivore–frugivores or insectivore–granivores. In most species, virtually all assimilated food was of C₃ origin with the exception of Ruddy Ground‐Doves (Columbina talpacoti) in which CAM or C₄ plants contributed significantly. Seasonal changes in relative food source contribution were followed in eight species of birds. Of these species, White‐tipped Doves (Leptotila verreauxi), Grayish Saltators (Saltator coerulescens), and Social Flycatchers (Myiozetetes similis) increased their use of insects in the rainy season, in contrast to Great Kiskadees (Pitangus sulphuratus), which decreased their use of insects. Our study suggests that that diverse strategies are used by various avian species to obtain dietary proteins within seasonal habitats.     

Evidence of shift in C4 species range in central Argentina during the late Holocene

Aim

Millennial-scale biogeographic changes are well understood in many parts of the world, but little is known about long-term vegetation dynamics in subtropical regions. Here we investigate shifts in C₃/C₄ plant abundance occurred in central Argentina during the past few millennia

Methods

We determined present day soil organic matter ??¹³C signatures of grasslands, shrublands and woodlands, containing different mixtures of C₃ and C₄ plants. We measured past changes in the relative cover of C₃/C₄ plants by comparing ??¹³C values in soil profiles with present day ??¹³C signatures. We analyzed ¹⁴C activity in soil depths that showed major changes in vegetation.

Results

Present day relative cover of C₃/C₄ plants determines whole ecosystem ??¹³C signatures integrated as litter and superficial soil organic matter (R²?=?0.78; p?<?0.01). Deeper soils show a consistent shift in ??¹³C, indicating a continuous replacement of C₄ by C₃ plants since 3,870 (±210) YBP. During this period, the relative abundance of C₃ plants increased 32% (average across sites) with significant changes being observed in all studied ecosystems.

Conclusions

Our results show that C₄ species were more abundant in the past, but C₃ species became dominant during the late Holocene. We identified increases in the relative C₃/C₄ cover in grasslands, shrublands and woodlands, suggesting a physiological basis for changes in vegetation. The replacement of C₄ by C₃ plants coincided with changes in climate towards colder and wetter conditions and could represent a climatically driven shift in the C₄ species optimum range.     

Reliability of δ13C and δ15N in faeces for reconstructing savanna herbivore diet

We tested the reliability of herbivore faecal δ¹³C and δ¹⁵N values for reconstructing diet through review of an extensive database derived from a 3-year study of ungulates in South Africa's Kruger National Park. Faeces are a useful material for stable isotope studies of diet because they record dietary turnover at very short time scales, and because sampling is non-invasive. However, the validity of faecal isotope proxies may be questioned because they represent only undigested food remains. Results from Kruger Park confirm that free-ranging browsers have faecal δ¹³C consistent with C₃ feeding, grazer faeces are C₄, and mixed-feeder faeces intermediate. Although the respective ranges do not overlap, there is significant variation in faecal δ¹³C of browsers and grazers (～2.0–4.0‰) across space and through time. We demonstrate that most (～70%) of this variation can be ascribed to corresponding patterns of variation in the δ¹³C of C₃ and C₄ plants, respectively, re-enforcing the fidelity of faecal isotope proxies for diet but highlighting a need for mixing models that control for variations in plant δ¹³C in order to achieve accurate diet reconstructions. Predictions for the effects of climate (rainfall) and ecophysiology on ¹⁵N-abundance variations in mammals do not persist in faeces. Rather, faecal δ¹⁵N tracks changes in plant δ¹⁵N, with further fractionation occurring primarily due to variations in dietary protein (reflected by %N). Controlling for these effects, we show that a dual-isotope multiple source mixing model (Isosource) can extend diet reconstructions for African savanna herbivores beyond simplified C₃/C₄ distinctions, although further understanding of variations in mammal δ¹⁵N are needed for greater confidence in this approach.     

Tracking dietary habits of cave arthropods associated with deposits of hematophagous bat guano: A study from a neotropical savanna

Carbon and nitrogen stable isotopes were used to track major dietary variations in arthropods found in a cave located in a neotropical savanna in Central Brazil. We determined the δ¹³C and δ¹⁵N for cave crickets, cockroaches, spiders, guano of a hematophagous bat, and leaf litter found on the ground near the cave entrance. The δ¹³C and δ¹⁵N for the cricket and cockroaches showed that bat guano was not the only food item for these arthropods. They had intermediate δ¹³C and δ¹⁵N between bat guano and leaf litter, which means that they consumed additional food resources other than guano in their diet, independent of distribution of guano deposits in the cave. The spiders, predators with great mobility, seemed to not have a preferential diet item since their isotope signals indicated they hunt both crickets and cockroaches. The δ¹³C of the bat guano (?15.4‰) indicated that the diet of these hematophagous bats relied on animals fed with C₄ plants. The conversion of native savanna vegetation of which the debris is mostly from C₃ plants, to pasture, based on C₄ African grass, a remarkably common land use transformation in the region, is indirectly influencing the diet of hematophagous bats, which basically relied on exotic fauna. The C and N stable isotope ratios showed that the arthropods inside the cave do not rely solely on bat guano for their diet, but interact directly with the external environment, through litter debris, indicating a significant exchange of energy and matter between the cave environment and the surrounding area.     

Carbon isotopes and water use efficiency: sense and sensitivity

We revisit the relationship between plant water use efficiency and carbon isotope signatures (δ¹³C) of plant material. Based on the definitions of intrinsic, instantaneous and integrated water use efficiency, we discuss the implications for interpreting δ¹³C data from leaf to landscape levels, and across diurnal to decadal timescales. Previous studies have often applied a simplified, linear relationship between δ¹³C, ratios of intercellular to ambient CO₂ mole fraction (C _i/C _a), and water use efficiency. In contrast, photosynthetic ¹³C discrimination (Δ) is sensitive to the ratio of the chloroplast to ambient CO₂ mole fraction, C _c/C _a (rather than C _i/C _a) and, consequently, to mesophyll conductance. Because mesophyll conductance may differ between species and over time, it is not possible to determine C _c/C _a from the same gas exchange measurements as C _i/C _a. On the other hand, water use efficiency at the leaf level depends on evaporative demand, which does not directly affect Δ. Water use efficiency and Δ can thus vary independently, making it difficult to obtain trends in water use efficiency from δ¹³C data. As an alternative approach, we offer a model available at to explore how water use efficiency and ¹³C discrimination are related across leaf and canopy scales. The model provides a tool to investigate whether trends in Δ indicate changes in leaf functional traits and/or environmental conditions during leaf growth, and how they are associated with trends in plant water use efficiency. The model can be used, for example, to examine whether trends in δ¹³C signatures obtained from tree rings imply changes in tree water use efficiency in response to atmospheric CO₂ increase. This is crucial for predicting how plants may respond to future climate change. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.