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Carbon and nitrogen stable isotope ratios (δ13C and δ15N) have been used to study the structure of food webs. However, few studies have examined how a terrestrial food web can be depicted by this technique. We measured δ13C and δ15N in various consumers of four trophic groups (detritivores, herbivores, omnivores and predators), including vertebrates and invertebrates (14 orders, ≥24 families), as well as canopy and understorey leaves in a tropical rain forest in Malaysia. We found that δ13C and δ15N of the consumers differed significantly among the trophic groups. The predators had significantly higher δ13C than the herbivores, and were similar in δ13C to the detritivores, suggesting that most predators examined depend largely on below-ground food webs. δ15N was higher in predators than detritivores by about 3‰. The comparison of δ13C in plant materials and herbivores suggests that most herbivores are dependent on C fixed in the canopy layers. The vertebrates had significantly higher δ15N and δ13C than the invertebrates of the same trophic group, likely reflecting differences in the physiological processes and/or feeding habits. This study indicates that stable isotope techniques can help better understanding of the terrestrial food webs in terms of both trophic level and the linkage of above- and below-ground systems.

Distinct patterns can be discerned in the extensive ritual tooth ablation found among the human skeletal remains of the Late–Final Jomon period (ca. 3200–2800 cal BP) in Japan. Based on comparative observations of sex and grave patterns in the skeletal remains, two major patterns in ritual tooth ablation, termed type 4I and type 2C, have been assigned to locals and immigrants, respectively. In order to test this hypothesis, strontium (Sr) isotope ( 87 Sr/ 86 Sr) analyses were performed on human skeletal remains from the Yoshigo shell mound in Aichi Prefecture, central Japan. Plants in the surrounding area were also examined to illustrate the geographic 87 Sr/ 86 Sr distribution. The Sr isotopic variation in human tooth enamel ( 87 Sr/ 86 Sr = 0.70868–0.71028) was greater than that in human bones ( 87 Sr/ 86 Sr = 0.70871–0.70943). Individuals with higher Sr isotope ratios in their tooth enamel than seawater Sr values of 0.7092 can be identified as immigrants (36% of population). The presence of these isotopically identified immigrants between both type 2C and type 4I individuals does not support the previous hypothesis. The intra-population 87 Sr/ 86 Sr distribution of tooth enamel of type 2C individuals showed a significantly higher mean ratio than that of type 4I individuals, suggesting a higher proportion of immigrants among the former.

In the prolonged absence of catastrophic disturbance, ecosystem retrogression occurs, and this involves increased nutrient limitation, and reduced aboveground and belowground ecosystem processes rates. Little is known about how the nitrogen and carbon stable isotope ratios (delta(15)N and delta(13)C) of plants, soils and consumer organisms respond to retrogression in boreal forests. We investigated a 5000 year chronosequence of forested islands in the boreal zone of northern Sweden, for which the time since lightning-induced wildfire increases with decreasing island size, leading to ecosystem retrogression. For this system, tissue delta(15)N of three abundant plant species (Betula pubescens, Vaccinium myrtillus and Pleurozium schreberi) and humus all increased as retrogression proceeded. This is probably due to enhanced ecosystem inputs of N by biological fixation, and greater dependency of the plants on organic N during retrogression. The delta(13)C of B. pubescens and plant-derived humus also increased during retrogression, probably through nutrient limitation increasing plant physiological stress. Unlike the plants, delta(15)N of invertebrates (lycosid spiders and ants) did not increase during retrogression, probably because of their partial dependence on aquatic-derived prey that had a variable delta(15)N signature. The delta(13)C of the invertebrates increased as retrogression proceeded and converged towards that of an aquatic prey source (chironomid flies), suggesting increased dependence on aquatic-derived prey during retrogression. These results show that measurement of delta(15)N and delta(13)C of plants, soils, and consumers across the same environmental gradient can provide insights into environmental factors that drive both the aboveground and belowground subsystems, as well as the linkages between them.

Many epiphytic ferns in tropical forest canopies have domatia that provide habitat for arboreal arthropods such as ants. Two species of fern that were predominantly occupied by Crematogaster difformis ants were collected from the canopy of a forest in Borneo. In the fern domatia, two cockroach species, Pseudoanaplectinia yumotoi and Blatta sp., were found to live with C. difformis. Pseudoanaplectinia yumotoi were found in large numbers and accounted for approximately 20% of the total arthropod population, while Blatta sp. comprised only 1% of the arthropods living in the domatia. Behavioural experiments were conducted to evaluate the response of C. difformis workers to newly introduced cockroaches of each species and to allospecific ant workers. Crematogaster difformis workers were highly aggressive and quickly attacked allospecific workers. Both cockroach species could fend off ant attacks; however, dead Blatta sp. were often attacked by the ants while dead P. yumotoi were infrequently bothered.

Deterioration of aquatic ecosystems resulting from enhanced anthropogenic N loading has become an issue of increasing concern worldwide, and methods are needed to trace sources of N in rivers. Because nitrate from sewage is enriched in 15N relative to nitrate from natural soils, delta(15)N values of stream nitrate (delta(15)Nnitrate) should be an appropriate index of anthropogenic N loading to rivers, as should the delta(15)N values of riparian plants (delta(15)Nplant) because they are consumers of nitrate. We determined the delta(15)N values of stream nitrate and six species of riparian macrophytes in 31 rivers in the Lake Biwa Basin in Japan. We then tested the correlation between these values and various land-use parameters, including the percentage of land used for residential and agricultural purposes as well as for natural areas. These delta(15)N values were significantly positively correlated with land use (%) that had a high N load (i.e., residential or agricultural use) and significantly negatively correlated with forest (%). These findings indicate that delta(15)N values of stream nitrate and riparian plants might be good indicators of anthropogenic inputs of nitrogen.

We measured stable nitrogen (N) and carbon (C) isotope ratios, lignin-derived phenols, and fossil pigments in sediments of known ages to elucidate the historical changes in the ecosystem status of Lake Biwa, Japan, over the last 100 years. Stable N isotope ratios and algal pigments in the sediments increased rapidly from the early 1960s to the 1980s, and then remained relatively constant, indicating that eutrophication occurred in the early 1960s but ceased in the 1980s. Stable C isotope ratios of the sediment increased from the 1960s, but decreased after the 1980s to the present. This decrease in stable C isotope ratios after the 1980s could not be explained by annual changes in either terrestrial input or algal production. However, when the C isotope ratios were corrected for the Suess effect, the shift to more negative isotopic value in atmospheric CO 2 by fossil fuel burning, the isotopic value showed a trend, which is consistent with the other biomarkers and the monitoring data. The trend was also mirrored by the relative abundance of lignin-derived phenols, a unique organic tracer of material that originated from terrestrial plants, which decreased in the early 1960s and recovered to some degree in the 1980s. We detected no notable difference in the composition of lignin phenols, suggesting that the terrestrial plant composition did not change markedly. However, we found that lignin accumulation rate increased around the 1980s. These results suggest that although eutrophication has stabilized since the 1980s, allochthonous organic matter input has changed in Lake Biwa over the past 25 years.

We examined how nitrogen-stable isotopic signatures of food web components (basal resources, primary and lower consumers, and omnivores) in rivers change with increasing levels of human population density (HPD) in their watersheds. Samples were collected from 22 rivers flowing in the Lake Biwa basin, Japan. Among three potential resources at the base of food webs (epilithon, benthic and suspended particulate organic matter), the mean isotopic values (δ15N) of the epilithon (4.5–7.8%) were consistently higher than those of other items (1.9–4.2%) and displayed the most pronounced elevation (by 3.3%) with increasing HPD. The mean δ15N values of the individual taxa of lower consumers (bivalve, snail and caddisfly) tended to increase with increasing HPD, although the pattern and the extent of the elevation were highly variable among the taxa. These results suggest a taxon-specific feature in the N source (or sources) of lower consumers. Our data suggested that human activities (e.g. nutrient loading) potentially induce changes in the N baselines of river food webs. The major N source of bivalves appeared to be shifted from suspended particulate organic matter to other items with increasing HPD. Trophic levels of goby fish (Rhinogobius sp. OR) and shrimp (Palaemon paucidens), being estimated to be at 2.4–3.8 and 2.1–3.4, respectively, did not differ significantly among rivers with different HPD levels.

We analyzed stable carbon and nitrogen isotope ratios in a sample of human and nonhuman mammal bones excavated from the Yoshigo and Inariyama shell mounds of the Late-Final Jomon pe- riods in Aichi Prefecture, Japan, with a view to determining individual dietary differences. To inves- tigate possible reasons for the dietary variations identified, we related isotope ratios to sex and tooth ablation patterns. At both sites, large intra-site variations in δ 13 C and δ 15 N values were found, com- pared with other Jomon populations previously studied, suggesting higher than usual levels of dietary variability, and at both sites there was a positive correlation between δ 13 C and δ 15 N values. The diet of the Jomon people at both these sites had two main protein sources: marine (marine finfish and shell- fish) and terrestrial (C3 plants and terrestrial mammals) protein. The intra-site variability is probably explained by consumption of these resources in different proportions. Analysis of the Yoshigo shell mound data indicated that sex is one of the factors determining dietary difference. It was also found that individual differences in diet in Yoshigo males are greater than in females. This pattern was re- peated in the Inariyama shell mound data. Dietary differences were found to be related to ritual tooth ablation characteristics, particularly in males. At Inariyama, type 4I ritual tooth ablation was associated with comparatively greater dependency on terrestrial resources, while type 2C tooth ablation was as- sociated with greater dependency on marine resources. This may indicate that type 4I males engaged predominantly in hunting, and type 2C males in fishing, as a means of food acquisition. These results are possibly the earliest evidence of occupational differentiation in the Jomon people.

We investigated the effects of natural environments and human activity on Lake Biwa, central Japan. We determined the concentrations of 19 elements and the compositions of stable S and Sr isotopes in the main tributaries of the lake and compared them with the corresponding values obtained from the lake water during the circulation period. Results of a principal component analysis indicated that the components dissolved in the lower reaches of the tributaries can be divided into group 1 (HCO(3), SO(4), NO(3), Ca, Mg, Sr) and group 2 components (Cl, Br, Na, K, Ba, Rb, Cs). The concentrations of group 1 components were high in the rivers of the southern area, which is urbanized and densely populated, and the eastern area, which consists of plains where agriculture predominates, compared with the rivers of the northern and western areas, which are mostly mountainous and sparsely populated. The concentrations of group 2 components tended to be high in the river water of industrial areas. The delta(34)S values of SO(4) in the river water converged to 0+/-2 per thousand as the SO(4) concentration increased and, excluding the areas where limestone is extensively distributed, as the HCO(3) concentration increased. In particular, both the delta(34)S values (0+/-2 per thousand) and the (87)Sr/(86)Sr ratios (0.7117+/-0.0005) fell within narrow ranges in the small and medium rivers of the eastern plain area, where rice is widely grown. These values agreed respectively with the delta(34)S values of the fertilizers used in the Lake Biwa basin and the soil-exchangeable (87)Sr/(86)Sr in the eastern plain. The characteristics of water quality in the small and medium rivers of the eastern area can be explained by a model in which sulfuric, nitric, and bicarbonic acids generated by the decomposition of agricultural fertilizer and paddy rice selectively leached out alkaline-earth elements adsorbed on the soil and sediments of the plain or dissolved calcium carbonate enriched with Mg and Sr. Compared with tributary waters, the lake water was depleted in NO(3), owing to denitrification, and in Mn, owing to mineralization, which occur under the redox condition of bottom sediments. Excluding NO(3) and Mn, the compositions of both the dissolved elements and the Sr and S isotopes in the water of Lake Biwa can be approximately reproduced by simple mixing of the tributary water, indicating that these components provide effective indices for evaluating the relationship between the waters of the lake and its tributaries.

We measured the natural stable carbon (C) and nitrogen (N) isotope ratio patterns of collembola and the organic substrates of their habitats and potential food sources in a warm temperate coniferous forest. Based on previous studies, we classified collembola into successional classes along litter decomposition gradients: early colonizers, late colonizers, and dominants-throughout. The stable C and N isotope ratios of late colonizers exceeded those of early colonizers, and those of the dominants-throughout were intermediate between early and late colonizers, which is consistent with previous studies on two macro-invertebrates, earthworms and termites. The C and N isotopic signature differences in collembola may reflect food resource partitioning along decomposition gradients.

The degradation of tropical forests is progressing rapidly and its ecological effects on wild animals are a global concern. We evaluated the hypothesis that small mammals in highly degraded forest occupy higher trophic levels than those in somewhat degraded forests, as indicated by diets high in consumers such as insects, in a tropical rain forest in Sarawak, Malaysia. After correcting for differences in the δ15N values for primary production among the study sites, the δ15Ncorrected values for rats and mice (Muridae) differed significantly among forest types. Rats and mice in more degraded forest had higher δ15Ncorrected values than those in less degraded or primary forest; in contrast, treeshrews (Tupaiidae) and squirrels (Sciuridae) showed no significant differences in the δ15Ncorrected values among forest types. We found significant positive correlations between canopy openness and the δ15Ncorrected values for one species of squirrel and two species of rats. This hypothesis was supported for small mammals that have normal dietary preferences for plants, i.e., omnivorous rodents, but not for those that normally prefer insects, i.e., treeshrews. The δ15N values for omnivorous mammals may be useful as an indicator of changes in food-web structure in response to forest disturbance.

Summary 1We propose that diets of consumers in a food web have various ages, where age is defined as the time elapsed since carbon (C) in the diet was fixed from atmospheric CO2 by primary producers. To examine the diet ages for primary consumers in a detrital food web, we measured the radiocarbon (14C) content of termites collected in Thailand in 1998 and 2004. Diet ages were estimated by comparing the 14C content of samples with records of atmospheric 14CO2, which doubled in the early 1960s as a result of nuclear weapons tests and decreased after the nuclear test ban treaty. For comparison, we measured the 14C content of bees as primary consumers in a grazing web at the same study site. Stable carbon and nitrogen (N) isotope ratios were also analysed. 2The 14C contents of the same species of termites decreased during the sampling interval, indicating that they used organic matter produced after the peak in atmospheric 14CO2. The diet ages were estimated to be 12–18, 7–13 and 5–9 years for the wood-feeder (Microcerotermes crassus), the soil-feeders (Dicuspiditermes makhamensis and Termes comis) and the fungus-grower (Macrotermes carbonarius), respectively. One colony of soil-feeder (T. comis), which nested in a fallen tree trunk, had exceptionally low 14C content, and its diet age was estimated to be around 50 years. The two bee species had lower 14C contents compared with the termites, and their diet ages were estimated to be 0 (Apis florea) and 2–4 years (Trigona sp.). 3Stable C and N isotope ratios of termites showed similar patterns as previously reported, and no clear difference was observed between 1998 and 2004. Although the bees and the fungus-growing termite had similar stable C and N isotope ratios, their diet ages differed. 4Our study suggests that radiocarbon can be used to estimate the diet ages of consumers in terrestrial food webs. Diet age should provide new insight into the trophic positions of organisms in grazing and detrital food webs and the interactions between these two webs.

しろかき期において強制落水を行ったときに水田から流出する懸濁物 (SS), 窒素, リンの量を圃場 (30×100m) における流出実験から見積もった。強制落水における総流出量は, 水位の減少の小さい圃場 (8mm, 水田A) ではSSが33kg, 全窒素が85g, 全リンが46g, 水位の減少が大きい圃場 (72mm, 水田B) ではSSが110kg, 全窒素が589g, 全リンが146gであった。水深が深い圃場からのリンの流出が特に大きいことが明らかになった。さらに, 強制落水による排水のN/P(モル比)は, それぞれ4.1 (水田A), 8.8 (水田B) であった。植物プランクトンのN/Pが16であることを考えると, 強制落水は特にリンを高い効率で流出させることがわかった。琵琶湖へのリンの供給は富栄養化の促進の点から望ましくなく, 水田の粗放的水管理の改善が今後の琵琶湖の水質を考える上で重要であることが指摘された。

To study the deterioration of the water quality in Lake Biwa, Japan, over the last 40 years, we measured the concentrations and isotopic ratios of sulfur and strontium of water in 41 inflowing rivers and one discharging river. The concentrations of SO4 and Sr of inflowing rivers at downstream sites were generally high in the southern urban area and in the eastern area, where a large agricultural plain is situated, but low in the northern and western areas, whose watersheds are mountainous and with low population density. SO4 and Sr concentrations are also lower at upstream sites, which are closer to mountainous areas. Thus, the inflowing river receives large amounts of SO4 and Sr as it flows across the plain, where human activity levels are high. The delta34S or 87Sr/86Sr values of most eastern rivers at downstream sites are lower than those of water in Lake Biwa, and values become more uniform as the proportion of the plain area in the watershed increases. River water in other areas has higher values of delta34S or 87Sr/86Sr than the lake water. This result indicates that the decadal decrease of delta34S and 87Sr/86Sr in the lake water has been caused mainly by the increased flux of SO4 and Sr from rivers in the eastern plain. We assume that in the plain, sulfur, nitrogen, and organic compounds induced by human activities generate sulfuric, nitric, and organic acids in the water, which accelerate the extraction of Sr from bedrocks, leading to the generation of Sr in the river water in the area.

We examined the crown architecture of two canopy species, Fagus crenata Blume, which is highly shade tolerant, and Quercus crispula Blume, which is less shade tolerant, in upper and lower positions on a slope in a cool-temperate forest in Japan. The purpose of this study was to investigate the relative effects of the competitive position for the light environment and developmental constraints on the crown architecture of two species that have different degrees of shade tolerance. In both species, the maximum attainable height was smaller on the upper slope than on the lower slope. The light environment brightened with increasing tree height. Trees in similar competitive position and thus similar light environments were shorter on the upper slope than on the lower slope. When we considered tree height relative to the maximum attainable height, the difference between the upper and lower slopes in the relationship of competitive position to tree height disappeared. Thus, relative tree height was a good index of the competitive position for the light environment when the trees of both slope positions were pooled. The relationships of crown depth and width to relative tree height differed between trees on the upper and lower slopes, while the relationships of crown depth and width to actual tree height were similar. This suggests that developmental constraints are more important than the competitive position in determining the crown architecture of the two species. However, variation in crown architecture was greater in Q. crispula than in F. crenata , probably because of the difference in shade tolerance between the two species.

Summary 1To investigate whether the role of symbiotic fungi in lignin degradation and food provision differs among fungus-growing termites (four species of Macrotermes, three species of Odontotermes, Hypotermes makhamensis, Ancistrotermes pakistanicus and Pseudacanthotermes militaris), the chemical composition of fungus combs of different ages and the chitinolytic activity in the gut of termites were analysed. In addition, the carbon stable isotope ratios (δ13C) in old combs, worker termite tissues and fungal nodules (aggregated conidia) were compared. 2In Macrotermes spp., the carbohydrate : lignin ratio of the combs increased with increased comb age, but it decreased or remained the same in Odontotermes spp., H. makhamensis, A. pakistanicus and P. militaris. In contrast, the chitin : carbohydrate ratio and ash content increased more over time in the combs of Odontotermes spp., H. makhamensis, A. pakistanicus and P. militaris than in the combs of Macrotermes spp. 3Chitinolytic activity in the gut of workers was higher in O. takensis, H. makhamensis and A. pakistanicus than in M. annandalei. 4In all species examined, the δ13C value increased in the sequence: old combs < termites < fungal nodules. Based on a two-source model of δ13C, the contribution of fungi to termite nutrition was considered higher in Odontotermes spp., H. makhamensis, A. pakistanicus and P. militaris than in Macrotermes spp. 5These results suggest that symbiotic fungi play different roles among fungus-growing termites. In Macrotermes spp., the main role of symbiotic fungi is to degrade lignin, so that the termites can utilize cellulose more efficiently, whereas in Odontotermes spp., H. makhamensis, A. pakistanicus and P. militaris, it is to serve as a food source.

1. Nitrogen and carbon isotope ratios and uric acid concentrations in fungus-growing termites (Isoptera: Termitidae: Macrotermitinae), sampled in Cameroon and Thailand, were determined in order to compare castes that are known to differ in behaviour and feeding habits. 2. Nitrogen isotope ratios (δ15N) were either not significantly changed or lower in workers compared with the diet (the fungus combs), whereas carbon isotope ratios (δ13C) were higher in worker termites than in the fungus combs. 3. In old workers, δ15N values were unexpectedly low and correlated negatively with whole-body uric acid concentrations. This indicates that older workers retain uric acid, which has a low δ15N value, to conserve nitrogen within the colony and, furthermore, that older colony members may ultimately be consumed by younger conspecifics.

Tissue N and C stable isotope ratios of two sympatric termite species were investigated in relation to those of their stored food materials at two savanna sites in north-eastern Australia (Woodstock and Oak Valley). At both sites, the δ 15 N values of termite tissues and the stored food materials of the litter-feeding species Amitermes laurensis were significantly greater than their equivalents in the grass-harvesting species Drepanotermes rubriceps . In addition, the carbohydrate/lignin and C/N ratios were higher and chitin/carbohydrate ratios were lower in food materials stored in the mounds of D. rubriceps . Both sets of analyses confirmed that D. rubriceps assimilates less decomposed food materials than A. laurensis . Tissue δ 13 C values indicated that both species assimilated similar proportions of C3 and C4 plant materials at the Woodstock site, but that the workers of A. laurensis assimilated a greater proportion of C3 plant material at the Oak Valley site. The δ 15 N values of termites from the Woodstock site were significantly greater than those from Oak Valley, although the differences in order at each site were consistent between species. The results indicate that N and C stable isotope ratios must be carefully compared across habitats.

Stable (δ13C and δ15N) and radio- (14C, 137Cs and 210Pb) isotopes were determined for termites that have been sampled from a dry evergreen forest in Thailand. A wood-feeding termite, Microcerotermes crassus, was separated from soil-feeders: Termes propinquus, Termes comis and Dicuspiditermes makhamensis by δ13C and δ15N values. The Termes group in Thailand had less diverse values in δ13C and δ15N than those in Australia, where the feeding habits of the ‘Termes’ group are more diverse. Other soil-feeding termites produced similar δ13C values, but a larger range in δ15N values. 14C-percent modern carbon (pMC) values suggest that the soil-feeding termites used younger carbon than the wood-feeding termites, and this was consistent with the termites from Cameroon, central Africa. Values of δ13C and 14C-pMC indicate that surface soil was used by a soil-feeding termite, D. makhamensis, in making the nest mounds, and deeper soil (10–30 cm) by a fungus-growing termite, Macrotermes carbonarius. 210Pb and 137Cs were scarcely incorporated into the termites, although 214Pb was recovered from the workers. The results suggest that stable- and radioisotopes are useful in the study of detritivorous animals, organic matter decomposition and ecosystem engineering.

Pycnotarsobrentus inuiae MaruyamaBartolozzi, gen. nov. and sp. nov. (Brentinae: Eremoxenini) is described from the Lambir Hills National Park, Borneo (Sarawak, Malaysia) based on specimens collected from Crematogaster difformis F. Smith, 1857 ant nests in the myrmecophytic epiphytic ferns Platycerium crustacea Copel. and Lecanopteris ridleyi H. Christ. A second species of Pycnotarsobrentus is known from Malaysia but is represented by only one female and consequently not yet described pending discovery of a male. Pycnotarsobrentus belongs to the tribe Eremoxenini and shares some character states with the African genus Pericordus Kolbe, 1883. No species of Eremoxenini with similar morphological modifications are known from the Oriental region.

In order to investigate the role of the mutualistic fungus, Termitomyces sp., in the fungus-growing termite, Macrotermes gilvus , we applied CP/MAS 13 C NMR and selected proximate analyses to fungus comb of different ages and degrees of maturation. We found evidence that lignin degradation took place progressively in the fungus comb. In vitro digestibility of cellulose in old fungus comb, on which the termites feed, was approximately 3-fold higher than that in the fresh part. These results confirm the ‘lignin degradation hypothesis’ that the role of the mutualistic fungi is to degrade lignin and enhance the digestibility of cellulose for the termites, suggesting the ability of the termite–fungus association to make extremely efficient use of plant material.

In the prolonged absence of catastrophic disturbance, ecosystem retrogression occurs, and this involves increased nutrient limitation, and reduced aboveground and belowground ecosystem processes rates. Little is known about how the nitrogen and carbon stable isotope ratios (delta(15)N and delta(13)C) of plants, soils and consumer organisms respond to retrogression in boreal forests. We investigated a 5000 year chronosequence of forested islands in the boreal zone of northern Sweden, for which the time since lightning-induced wildfire increases with decreasing island size, leading to ecosystem retrogression. For this system, tissue delta(15)N of three abundant plant species (Betula pubescens, Vaccinium myrtillus and Pleurozium schreberi) and humus all increased as retrogression proceeded. This is probably due to enhanced ecosystem inputs of N by biological fixation, and greater dependency of the plants on organic N during retrogression. The delta(13)C of B. pubescens and plant-derived humus also increased during retrogression, probably through nutrient limitation increasing plant physiological stress. Unlike the plants, delta(15)N of invertebrates (lycosid spiders and ants) did not increase during retrogression, probably because of their partial dependence on aquatic-derived prey that had a variable delta(15)N signature. The delta(13)C of the invertebrates increased as retrogression proceeded and converged towards that of an aquatic prey source (chironomid flies), suggesting increased dependence on aquatic-derived prey during retrogression. These results show that measurement of delta(15)N and delta(13)C of plants, soils, and consumers across the same environmental gradient can provide insights into environmental factors that drive both the aboveground and belowground subsystems, as well as the linkages between them.

Zygomorphic flowers are usually more complex than actinomorphic flowers and are more likely to be visited by specialized pollinators. Complex zygomorphic flowers tend to be oriented horizontally. It is hypothesized that a horizontal flower orientation ensures effective pollen transfer by facilitating pollinator recognition (the recognition-facilitation hypothesis) and/or pollinator landing (the landing-control hypothesis). To examine these two hypotheses, we altered the angle of Commelina communis flowers and examined the efficiency of pollen transfer, as well as the behavior of their visitors. We exposed unmanipulated (horizontal-), upward-, and downward-facing flowers to syrphid flies (mostly Episyrphus balteatus), which are natural visitors to C. communis. The frequency of pollinator approaches and landings, as well as the amount of pollen deposited by E. balteatus, decreased for the downward-facing flowers, supporting both hypotheses. The upward-facing flowers received the same numbers of approaches and landings as the unmanipulated flowers, but experienced more illegitimate landings. In addition, the visitors failed to touch the stigmas or anthers on the upward-facing flowers, leading to reduced pollen export and receipt, and supporting the landing-control hypothesis. Collectively, our data suggested that the horizontal orientation of zygomorphic flowers enhances pollen transfer by both facilitating pollinator recognition and controlling pollinator landing position. These findings suggest that zygomorphic flowers which deviate from a horizontal orientation may have lower fitness because of decreased pollen transfer.

1. Stable nitrogen (N) isotope has been widely used to disentangle food webs and to infer trophic positions of organisms based on an assumption that the stepwise enrichment occurs along trophic levels. The enrichment of N-15 in soil organisms with diet humification has also been reported, but the underlying mechanism has not been fully examined. 2. To examine the effect of diet humification on N-15, we estimated the stable N isotope ratios and diet ages of earthworms and termites. These organisms feed on organic matter with various degrees of humification, ranging from undecomposed plant materials to humified organic matter (soil organic matter), in a gallery forest and a savanna in the Ivory Coast. We defined diet age as the time elapsed since carbon (C) in the diet of earthworms and termites was fixed from atmospheric CO2 by photosynthesis; it was estimated by comparing the radiocarbon (C-14) content of these organisms to atmospheric (CO2)-C-14 records. 3. Stable N isotope ratios increased along the humification gradient of diets, and values for earthworms and termites varied from 1.8 parts per thousand to 9.9 parts per thousand and from -1.5 parts per thousand to 15.9 parts per thousand, respectively. Epigeic (litter-feeding) earthworms had younger diet ages (2-4 years), whereas endogeic (soil-feeding) earthworms generally exhibited older diet ages (5-9 years). Grass-feeding termites had young diet ages (2 years), and wood/soil-feeding termites had the oldest diet ages (c. 50 years). Soil-feeding termites were similar in diet age (7-12 years) to wood feeders (8-11 years), with the exception of one species (18-21 years) that consumes large-diameter wood. 4. A significant positive relationship was found between diet ages and stable N isotope ratios of the two groups in the savanna. This relationship held in the gallery forest when termites feeding on woody tissues were not considered. These results show that the stable N isotope ratios of organisms can increase with diet age, unless C in the diet has been stored as organic matter, such as woody tissue, that is able to age without being subject to humification processes. 5. Given that above-ground food webs are often sustained directly by material and energy flow from below-ground food webs, in addition to trophic interactions, gradual enrichment of N-15 with the humification of below-ground diets should be considered when interpreting stable N isotope ratios of terrestrial food webs.

Pycnotarsobrentus inuiae Maruyama & Bartolozzi, gen. nov. and sp. nov. (Brentinae: Eremoxenini) is described from the Lambir Hills National Park, Borneo (Sarawak, Malaysia) based on specimens collected from Crematogaster difformis F. Smith, 1857 ant nests in the myrmecophytic epiphytic ferns Platycerium crustacea Copel. and Lecanopteris ridleyi H. Christ. A second species of Pycnotarsobrentus is known from Malaysia but is represented by only one female and consequently not yet described pending discovery of a male. Pycnotarsobrentus belongs to the tribe Eremoxenini and shares some character states with the African genus Pericordus Kolbe, 1883. No species of Eremoxenini with similar morphological modifications are known from the Oriental region.

Nitrogen and carbon stable isotope ratios and uric acid concentrations in termites sampled from a dry evergreen forest in Thailand, were determined across three kinds of feeding habits. Feeding habits of Microcerotermes crassus, which is an abundant wood-feeder, and Dicuspiditermes makhamensis, a common soil-feeding termite, were confirmed by isotopic signatures. Lichen feeding termites (Hospitalitermes birmanicus, H. bicolor and H. ataramensis) were characterized by low delta15N values, suggesting that they assimilated nitrogen deposited from the atmosphere. There was also a significant difference in uric acid concentrations between termites representing different feeding habits. No significant relationships were found between uric acid concentrations and delta15N or delta13C in Hospitalitermes. However, delta15N values were correlated with C/N ratios in H. birmanicus, except in one colony of H. ataramensis. delta13C values in both species were negatively correlated with C/N ratios.

Trophic fractionation of carbon and nitrogen isotopes (Δδ13C, Δδ15N) was examined using previously complied databases for food chains in Lake Biwa, Lake Baikal, and Mongolian grassland. The following two features were clarified: (1) For each ecosystem, the ratios of trophic fractionation of carbon and nitrogen isotopes (Δδ15N/Δδ13C) throughout food chain could be obtained as the slope of linear regression line on the δ15N–δ13C plot. (2) Further, analysis of covariance (ANCOVA) revealed the slopes on δ15N–δ13C were not significantly different among these various ecosystems and allowed us to have the regression by setting δ15N as the response variable: δ15N = 1.61 δ13C + [ecosystem specific constant] with standard errors of [±0.41] and [±9.7] for the slope and the intercept, respectively. It was suggested that the slope of the regression (or the ratio Δδ15N/Δδ13C) could be applicable to more complicated food webs in case nitrogen and carbon isotope ratios of primary producers can be assumed constant in space and time within the ecosystems. The results from simple linear regression analyses coincided well with the ANCOVA results for these ecosystems, although there was some discrepancy between the results of the two statistical analyses. Possible factors that govern the linear relationship between δ15N and δ13C along a food chain are discussed together with a new scope for the stable isotope food chain analyses.